PERTflfllHA December 1988
Vol. 11 No. 3 Contents
Identification of Soybean Genotypes in Wide Rows for Use in Narrow-row Culture — Ghizan Saleh and Earl T. Griton. Seed-borne Infection and Development of Colletotrichum capsici in Naturally Infected Chili Seed — Sariah Meon. The Induction and Evaluation of Productive Semidwarf Mutants of Basmati-370 — M. Siddique Sajjad and M.A. A wan. Studies on Some Sooty Moulds on Guava in Malaysia — Lim Tong Kee. Media for Embryo Culture of Some Tropical Recalcitrant Species — H.F. Chin, B. KrishnapiHay and Z.C. A/ang
333
Progress of Crop in Some Rhizophora Stands before First Thinning in Matang Mangrove Reserve of Peninsular Malaysia — P.B.L. Srivastava, Saw Leng Guan and Ashari Muktar. Properties of Soils Derived from Some Metamorphic Rocks in Peninsular Malaysia — S. Zauyah. Some Changes in Chemical Characteristics of a Paddy and Mangrove Soil Sample during Submergence in Water — J. Marcus, M. Faridawati and M.L. Zalma. Evaluation of Potato Crisps Fried in Market Samples of Palm Olein, Corn Oil and Soya Oil — M.A. August in, L.K, Heng and Nor Aini Idris The Digestion, Absorption and Ultization of Refined Palm Oil, Palm Olein and Palm Stearin in the Rat — T.K. W. Ng, H.T. Khor and YM. Chong. Effect of Chemical Treatments on the Shelf Life of Rambutans (Nephelium lappaceum)II — Suhaila Mohamed, Eshah Othman and Fauziah Abdullah. Topographic Anatomy of the Abdomen of the Lesser Mousedeer (Tmgulus javanicus) — K.C. Richardson, M.K. Vidyadaran, N.H. Fuzina and T.I. Azmi. The Radiographic Anatomy of the Gastrointestinal Tract of the Lesser Mousedeer (Tragulus javanicus) — K.C. Richardson, M.K. Vidyadaran, TJ. Azmi and N.H. Fuzina. Preparation of Arsenobetaine hydrobromide — Hazar Ismail and Robert F. Toia. Malaysian Natural Rubber Market Model — Mohammed bin Yusof Real Money Balances in the Production Function of a Developing Economy: A Preliminary Study of the Malaysian Agricultural Sector — Muzafar Shah Habibullah. Some Stream Water Quality Characteristics of Two Small Logged Over Watersheds in Selangor — Lai F.S. and Norajiki A.J. Kesan Saliniti dan Keamatan Cahaya ke Atas Pertumbuhan Chlorella virginica — Abdullah Zaini Alias. Pengubahsuaian Bentuk Selang Kecerunan Dalam Kaedah Newton bagi Suatu Kelas Fungsi Satu Pembolehubah — Ismail bin Mohd. Communications Survival and Isolation of Avian Mycoplasmas from Drinking Water of Infected Chickens — Mohd Shah-Majid. Preliminary Estimates of Growth and Mortality in Nemipterus bathybus (Pisces: Nemipteridae) from the Coast off Sarawak, South China Sea — Mohd. Azmi Ambak, Abu Kair Mohammad Mohsin and Mohd. Zaki Mohd. Said.
365
A Scientific Journal published by UNIVERSITI PERTANIAN MALAYSIA
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483 487
ISSN 0126-6128
FniTORiAT ROAwn ILUI i CFI^IAI. BVJAKU
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PSRTflniHft Number 3, December 1988
Contents
Identification of Soybean Genotypes in Wide Rows for Use in Narrow-row Culture — Ghizan Saleh and Earl T. Griton. Seed-borne Infection and Development of Colletotrichum capsici in Naturally Infected Chili Seed — Sariah Meon. The Induction and Evaluation of Productive Semidwarf Mutants of Basmati-370 — M. Siddique Sajjad and M.A. Awan. Studies on Some Sooty Moulds on Guava in Malaysia — Lim Tong Kee. Media for Embryo Culture of Some Tropical Recalcitrant Species — H.F. Chin, B. Krishnapillay and Z.C. Alang Progress of Crop in Some Rhizophora Stands before First Thinning in Matang Mangrove Reserve of Peninsular Malaysia — P.B.L. Srivastava, Saw Leng Guan and Ashari Muktar. Properties of Soils Derived from Some Metamorphic Rocks in Peninsular Malaysia — S. Zauyah. Some Changes in Chemical Characteristics of a Paddy and Mangrove Soil Sample during Submergence in Water —J. Marcus, M. Faridawati and M.L. Zalma. Evaluation of Potato Crisps Fried in Market Samples of Palm Olein, Corn Oil and Soya Oil — M.A. August in, L.K. Heng and Nor Aini Idhs The Digestion, Absorption and Ultization of Refined Palm Oil, Palm Olein and Palm Stearin in the Rat — T.K. W. Ng, H.T. Khor and Y.H. Chong. Effect of Chemical Treatments on the Shelf Life of Rambutans (Nephelium lappaceum) II — Suhaila Mohamed, Eshah Othman and Fauziah Abdullah. Topographic Anatomy of the Abdomen of the Lesser Mousedeer (Tragulus javanicus) — K.C, Richardson, M.K. Vidyadaran, N.H. Fuzina and T.I. Azmi. The Radiographic Anatomy of the Gastrointestinal Tract of the Lesser Mousedeer (Tragulus javanicus) — K.C. Richardson, M.K. Vidyadaran, T.I. Azmi and N.H. Fuzina. Preparation of Arsenobetaine hydrobromide — Hazar Ismail and Robert F. Toia. Malaysian Natural Rubber Market Model — Mohammed bin Yusof Real Money Balances in the Production Function of a Developing Economy: A Preliminary Study of the Malaysian Agricultural Sector — Muzafar Shah Habibullah. Some Stream Water Quality Characteristics of Two Small Logged Over Watersheds in Selangor — Lai F.S, and Norajiki A.J. Kesan Saliniti dan Keamatan Cahaya ke Atas Pertumbuhan Chlorella virginica — Abdullah Zaini Alias. Pengubahsuaian Bentuk Selang Kecerunan Dalam Kaedah Newton bagi Suatu Kelas Fungsi Satu Pembolehubah — Ismail bin Mohd. Communications Survival and Isolation of Avian Mycoplasmas from Drinking Water of Infected Chickens — Mohd Shah-Majid. Preliminary Estimates of Growth and Mortality in Nemipterus bathybus (Pisces: Nemipteridae) from the Coast off Sarawak, South China Sea — Mohd. Azmi Ambak, Abu Kair Mohammad Mohsin and Mohd. Zaki Mohd. Said.
333 341 345 349 357 365 375 385 393 399 407 419 421 437 441 451 461 469 475
483 487
Pertanika 11(3), 333-340 (1988)
Identification of Soybean Genotypes in Wide Rows for Use in Narrow-row Culture GHIZAN[ SALEH^ a n d EARL T. GRITTON2 1 Department of Agronomy and Horticulture, Faculty of Agriculture, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia. 2
Department of Agronomy, University of Wisconsin, Madison, WI53706, U.S.A.
Key words: Breeding nurseries; response to row spacing; Glycine max (L.) Merr. ABSTRAK Tiga puluh warisan dan enam kultivar kacang soya fGlycine max (L.) Merr.) telah dinilaikan di dalam jarak barisan luas (76 sm) dan rapat (18 sm)t didua lokasipada tahun 1982 dan satu pada tahun 1983. Objektif kajian ini ialah untuk menentukan sama ada penilaian di dalam barisan luas boleh digunakan untuk mengenalpasti warisan-warisan yang unggul untuk kultur barisan rapat. Hasil di dalam barisan rapat adalah 27.0% lebih tinggi dari barisan luas di kedua-dua lokasi pada tahun 1982 dan 20.8% lebih tinggi pada tahun 1983. Pada amnya, entri-entri yang berhasil tinggi di dalam barisan luasjuga merupakan yang berhasil tinggi di dalam barisan rapat, walaupun ianya tidak semestinya merangkumi semua yang berhasil tinggi di dalam barisan rapat. Satu kultivar dan dua warisan, walau bagaimanapun, adalah kekal menunfukkan respons yang sederhana tinggi kepada penurunan jarak barisan di kedua-dua tahun dan lokasi. Tiada ciri telah dikenalpasti berkait dengan respons tersebut. Pada amnya, pemilihan untuk warisan-warisan yang unggul di dalam nurseri-nurseri barisan luas akan dapat mengenalpasti warisanwarisan yang akan berprestasi tinggi di dalam kultur barisan rapat.
ABSTRACT Thirty lines and six cultivars of soybeans (Glycine max (L.) Merr.) were evaluated in wide-row (76 cm) and narrow-row (18cm)spacingsat two locations in 1982 and one in 1983. The objective of the study was to determine if evaluation of lines in wide rows can be used to identify superior lines for narrow-row culture. Yield in narrow rows was 27.0% higher than wide rows at both locations in 1982 and 20.8% higher in 1983. In general, the high yielding entries in wide rows were also the highest yielding ones in narrow rows, although they did not necessarily include all the highest yielding ones in narrow rows. One cultivar and two lines were, however, consistent in exhibiting a moderate to high response to narrowing of row spacing, over years and locations. No trait was identified as being associated with the response. In general, selection of lines superior in wide row nurseries will identify lines that will perform best in narrow-row culture. INTRODUCTION Soybean [Glycine max (L.) Merr.] yields in the North Central U.S. have been shown to increase by narrowing the row width (Cooper, 1974; Costa
et al, 1980; Green et al, 1977; Hartung et al, 1980). More efficient light interception and wateruse was associated with narrow-row culture (Hicks et al, 1969; Peters and Johnson, 1960; Shaw and
GHIZAN SALEH AND EARL T. GRITON
Weber, 1967; Taylor, 1980). Cultivars were found to differ in the degree of response to narrowing of row widths, depending on the plant morphological and physiological characteristics (Cooper, 1977; Costa et alt 1980; Lehman and Lambert, 1960). Cooper (1977) and Costa et al (1980) found that early maturing cultivars exhibited a greater yield response to narrow row spacing than did late maturing ones. Small leaflet size and low leaflet mass in the late-maturing groups were also associated with high yields in narrow-row plantings (Metz et al, 1984). Weaver and Wilcox (1982), however, reported that the response was not affected by maturity. Short and lodging-resistant lines were also reported to be more responsive to narrow rows (Cooper, 1976); Costa et al, 1980; Green etal, 1977). The number of branches on soybean plants decreases as the plant spacing decreases (Costa et al, 1980; Lehman and Lambert, 1960; Weber et al, 1966). However, the reduction in the number of branches associated with narrow row plantings does not necessarily lead to a reduction in yield, because greater amounts of each of the yield components were produced by the main stem rather than the branches (Lehman and Lambert, 1960). Most soybean cultivar development and testing programmes are carried out based on the traditional wide-row (76 cm) culture. Consequently, lines which might do well in the narrow rows may not have been selected in the wide-row nurseries. Possible reliability of breeding and evaluating soybeans in the wide rows for use in the narrowrow culture has not been extensively studied. Identification of key morphological and physiological traits contributing to superior performance in narrow rows would be useful in breeding and selecting genotypes in wide rows for maximum production in narrow rows (Shibles and Weber, 1966). The objectives of this study were (1) to evaluate trie performance of experimental lines and cultivars under wide (76 cm) and narrow (18 cm) rows, so as to determine if the same lines were superior in both wide and narrow rows, and (2) to determine if keys traits can be measured in wide rows to predict for superior performance in narrow rows. MATERIALS AND METHODS
Thirty experimental lines and six adapted cultivars with diverse genetic background of maturity 334
groups ranging from Group I to III were used in the study. Pedigrees of the entries are shown in Table 1. The entries were evaluated in Wisconsin, U.S.A., at the University of Wisconsin Research Farm at Arlington in 1982 and 1983, and at the Rock County Farm, Janesville, in 1982. Soil types were Piano silt loam (typic Arguidoll) at both locations. The experiment was conducted in a split-plot arranged in a randomized complete block design with three replications at each environment. Entries were whole plots, while row spacings, wide (76 cm between rows) and narrow (18 cm between rows) were sub-plots. The plots were planted on May 8 and May 12 at Janesville and Arlington, respectively, in 1982, and on May 10 at Arlington, in 1983. The plots were each 3.66 m long and 3.05 m wide. A 4-row wide-row planter was used for planting in the wide-row plots and an 11-row narrow row planter for the narrow-row plots. The wide rows were seeded at the rate of 29.5 seeds per meter-row (387, 513 seeds ha" 1 ), while the narrow rows were seeded at the rate of 16.4 seeds per meter-row (592, 034 seeds ha"" 1 ). Harvesting was done on September 28 and October 16 ai Janesville and Arlington, respectively, in 1982, and on October 12 at Arlington in 1983, using a small-plot Hege combine. The harvest area was 4.18 m2 for each of the wide-row plots, and 3.41 m2 for each of the narrow-row plots. Data were taken on the following characteristics: yield (seed yield at 13.0% moisture content), plant height at maturity, lodging (score of 1 = all plants erect; 5 = all plants lodged), height of lowest pod, number of branches per plant, maturity (number of days from sowing to the date when 95% of pods were brown), days to canopy closure (number of days from planting to the date when plant canopy has occupied all inter-row spaces), days to flowering (number of days from planting to the date when 50% of plants have produced the first flower), number of seeds per pod, 100-seed weight, leaf area per plant (only at Arlington, in 1983), and seed quality (score given considering the amount and degree of wrinkling, defective seed coat, greenishness, and mouldy or rotten seeds, where 1 = very good; 5 = very poor). Analysis of variance was performed on the data at each environment. Data from the three environments were not combined in the analysis, because tests of homogeneity of error variance
PERTANIKA VOL. 11 NO. 3, 1988
IDENTIFICATION OF SOYBEAN GENOTYPES IN WIDE ROWS FOR NARROW-ROW CULTURE TABLE 1 Pedigrees and pedigree numbers of entries used in the study.
Entry
Pedigree
Pedigree
number 1 2 3 4 5
6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Hard in Corsoy 79 SRF 200 Hobbit Hodgson 78 Simpson
Hark X Disoy Corsoy X Disoy Corsoy X Disoy Salut X Blackhawk Salut X Grant Salut X Grant Blackhawk X Grant Salut X Amurskaja Salut X Amurskaja Corsoy X Harosoy Corsoy X Haxosoy Hark X Corsoy Hark X Corsoy Hark X Corsoy Hark X Corsoy Hawkeye X Disoy Hawkeye X Disoy Hawkeye X Disoy Blackhawk X Salut Grant X Salut Disoy X Hark Disoy X Hark Harosoy X Corsoy Corsoy X Hawkeye Corsoy X Hawkeye Blackhawk X Hark Blackhawk X Hark Blackhawk X Hark Hark X Harosoy Corsoy X Hawkeye Cultivar Cultivar Cultivar Cultivar Cultivar Cultivar
(Cochran and Cox, 1957) conducted showed that the environments were heterogeneous. Simple correlation coefficients were calculated between wide and narrow-row plots for all characteristics. Correlations between yield in narrow-row plots and other characteristics in wide row plots were also computed, to identify key traits measured in wide rows that may be used to predict superior performance in narrow rows. Yield response was defined as the yield of entry in narrow rows minus the yield of the same entry in wide rows, expressed as a percentage of the yield in wide rows.
10037 10064 10066 10091 10126 10132 10155 10175 10188 10209 10213 10239 10240 10245 10255 10265 10266 10287 10313 10345 10372 10379 10411 10483 10489 10501 10523 10525 10536 10490
RESULTS AND DISCUSSION The lines and cultivars used in the study differed significantly for yield and all other characteristics measured. Yields were higher in narrow rows than in wide rows in all three trials (Table 2). Average yields were 2990 kg h a " 1 , 3409 kg h a " 1 and 2836 kg h a " 1 , respectively, at Arlington in 1982, Janesville in 1982, and Arlington in 1983, for narrow-row plots, and 2372 kg h a " 1 , 2707 kg h a " 1 and 2359 kg h a " 1 , respectively, in the three environments, for wide row plots. Narrow-row
PERTANIKA VOL. 11 NO. 3, 1988
335
TABLE 2 Mean values for yield and other plant characteristics of soybean lines and cultivars grown in wide (76 cm) and narrow (18 cm) rows, in three environments. Row spacing
Yield
-cm-
-kg ruT 1 -
Plant
Lodging
height
m
Height of lowest pod
—cm—
Number of
Maturity
Canopy
Flower-
closure -days-
-days-
branches
-cm-
Leaf
Seeds
100-seed
Seed
ing
area*
per pod
weight
quality
—days—
-cm2-
-gO X
Arlington, 1982
z
2990*
77*
2.8*
11.1*
2.4*
150.9*
56*
72*
-
2.5
14.5
2.4
2372
73
2.2
8.7
3.9
152.6
98
73
-
2.5
14.5
2.4
7.2
4.6
10.8
19.2
23.0
0.5
3.4
1.7
4.3
2.5
16.5
Janesville, 1982 18
3409*
102*
3.1
13.9*
1.5*
150.4
54*
66*
-
2.6
15.8*
2.7
76
2707
95
3.1
11.4
3.1
150.4
86
67
-
2.6
15.3
2.6
7.8
5.1
1.0
18.3
34.3
0.8
4.1
1.9
4.5
2.9
15.9
3. 1988
O
a
m H
a 73
Arlington, 1983 18
2836*
90*
76
2359
85
3.0
6.7
5.4
10.3
c,<»)
on .EH AND
KA VOL.
18 76
3.3*
76
469.6*
2.5
14.7*
146.9
74
76
451.7
2.5
14.4
3.0
2.5
6.9
L5
12.8
4.8
3.5
16.4
147.2
15.2
2.4
27.8
34.5
Leaf area measurements were taken only at Arlington in 1983.
•Significant difference between wide and narrow-row plots (p < 0.05).
3.0
47*
1.4*
18.0*
H Z
TABLE 3 Ranking of soybean entries based on yield in narrow rows, and their respective yields in wide rows and yield responses, in three environments.
i
oo
Entry
Narrow-row (kg ha" 1 )
Wide-row (kg ha" 1 )
Response'7 (%)
Hardin Simpson Hobbit Corsoy 79 Hodgson 78 3 24 2 SRF200 28 19 7 23 6 20 16 17 27 1 14 9 18 8 29 5 11 4 10 15 25 12 22 21 13 30 26
3789 3487 3484 3477 3415 3346 3329 3299 3177 3165 3102 3076 3074 3065 3043 2993 2958 2948 2925 2914 2904 2886 2871 2854 2853 2838 2829 2802 2791 2783 2774 2681 • 2571 2528 2456 2180
2828 2576 2989 2412 2765 2319 2536 2587 2473 2456 2370 2645 2560 2266 2302 2219 2214 2202 2119 2334 2265 2463 2117 2322 2456 2436 2408 2468 2267 2412 2252 2163 1946 2251 2303 1718
34.0 35.4 16.6 44.2 23.5 44.3 31.3 27.5 28.5 28.9 30.9 163 20.1 35.3 32.2 34.9 33.6 33.9 38.0 24.9 28.2 17.2 35.6 22.9 16.2 16.5 17.5 13.5 23.1 15.4 23.2 23.9 32.1 123 6.6 26.9
Mean LSD (0.05)
2990 379
2372 379
27.0 7.9
Arlington, 1982
Yield
Entry
Narrow—row (kg ha" 1 )
Wide-row (kg ha" 1 )
Response3 (%)
Hobbit Hardin 2 Simpson Hodgson 78 3 SRF200 9 25 27 28 11 24 Corsoy 79 10 19 15 30 29 14 20 13 4 22 5 1 6 12 21 23 8 17 16 7 26 18
4584 4311 4163 3998 3933 3817 3771 3722 3701 3631 3601 3564 3546 3531 3527 3434 3390 3379 3367 3322 3299 3226 3195 3186 3176 3140 3128 3106 3069 3003 2959 2946 2941 2913 2634 2501
3723 3244 2984 3253 2724 2775 2915 2889 2958 2612 2831 2565 2982 2943 2535 2575 2797 2830 2574 2991 2625 2673 2500 2778 2636 2602 2773 2366 2106 2329 2787 2618 2518 2595 1976 2062
23.1 32.9 39.5 22.9 44.4 37.5 29.4 28.8 25.1 39.0 27.2 38.9 18.9 20.0 39.1 33.4 21.2 19.4 30.8 11.1 25.7 20.7 27.8 14.7 20.5 20.7 12.8 31.3 45.7 28.9 6.2 12.5 16.8 12.3 33.3 21.3
Mean LSD (0.05)
3409 468
2707 468
27.0 7.5
Yield Narrow—row (kg ha" 1 )
Wide-row (kg ha" 1 )
Response* (%)
28 24 25 Hodgson 78 2 Simpson 27 Hobbit 4 Hardin 15 SRF200 19 Corsoy 79 13 23 26 3 30 5 9 29 11 17 1 7 18 22 16 6 12 8 21 10 20 14
3175 3163 3144 3135 3113 3103 3102 3100 3072 3070 3066 3033 3010 3000 2972 2938 2932 2930 2867 2841 2801 2794 2721 2720 2716 2712 2678 2660 2638 2524 2496 2465 2403 2397 2313 2292
2470 2614 2449 2249 2601 2564 2462 2969 2436 2493 2552 2385 2530 2581 2323 2464 2158 2401 2256 2410 2333 2392 2255 2288 2256 2473 2491 2087 2514 2322 1996 2037 1946 2250 1989 1948
28.5 21.0 28.4 39.4 19.7 21.0 26.0 4.4 26.1 23.1 20.1 27.2 19.0 16.2 27.9 19.2 35.9 22.0 27.1 17.9 20.1 16.8 20.7 18.9 20.4 9.7 7.5 27.5 4.9 8.7 25.1 21.0 23.5 6.5 16.3 17.7
Mean LSD (0.05)
2836 352
2359 352
20.8 6.2
t \
5 o CD
tn > Z
om
1 *V
S3 z < 5 m o
2*o Z >
90 M
io i n
LTURE
"Defined as the difference in yield between narrow and wide-row' plots, expressed as a percentage of yield in the wide—row plots.
3 TION i
m >
Janesville, 1982
Yield
IDE1
Arlington, 1983 Entry
GHIZAN SALEH AND EARL T. GRITON
plots produced 27.0% higher yield than wide-row plots at both locations in 1982, and 20.8% higher at Arlington in 1983 (Table 3). Effects of row spacing were significant on plant height, lodging, height of lowest pod, number of branches and days to canopy closure in both years, at Arlington. At Janesville in 1982, the effects were significant for plant height, height of lowest pod, number of branches, days to canopy closure, days to flowering and 100-seed weight (Table 2). Plants in narrow rows were significantly taller, with the lowest pod being higher from the ground, with lesser branches and had earlier canopy closure than those in wide rows, in all trials. Number of seeds per pod and seed quality were not affected by row spacing. Taller plants have been found to be more prone to lodging (Cooper, 1976). In this study, this was true only at Arlington, where lodging was significantly higher in the narrow rows than the wide rows. No difference was found between them at Janesville. At Arlington, plants in narrow rows matured earlier than those in wide rows. No difference was however, exhibited at the other two environments. Plants were earlier-flowering in narrow rows than wide rows at both locations in 1982, but were not affected by row spacing in 1983. Leaf area per plant, which was only measured at Arlington in 1983, was significantly greater in narrow rows than wide rows. One hundredseed weight was higher in narrow rows than wide rows at Janesville in 1982 and Arlington in 1983, but no difference was exhibited at Arlington in 1982. Ranking of entries based on yield in narrow rows are shown in Table 3. In general, the high yielding entries in wide rows were also the highest yielding ones in narrow rows. However, the ten highest yielding entries in narrow rows did not include all the ten highest yielding ones in wide rows, and vice versa. This was caused by the difference in the degree of response of entries to row spacing. For example, at Arlington in 1982 (Table 3), Corsoy 79 was ranked fourth in the narrow rows, but was only sixteenth in the wide rows, for yield performance. This shows that the cultivar was very responsive to narrow-row spacing. A similar pattern was shown by Hardin and Simpson in the trial. Considering performance in both row spacings in all trials, Hardin, Hobbit and Simpson were the highest yielding cultivars, while Entries 2, 3, 24 and 27 were the highest 338
yielding lines, comparable to the performance of the cultivars. Entries which showed the most yield response to narrow row spacing were generally, inconsistent over environments (Table 3). However, three entries, namely Hardin and Entries 3 and 27 were consistent in exhibiting a moderate to high yield increase. It is interesting to note that Entries 3 and 27 performed well and ranked quite high in narrow rows, but not in wide rows, generally, in all trials. Entry 3 showed 44.3, 37.5 and 22.0% yield response in narrow over wide at Arlington in 1982, Janesville in 1982 and Arlington in 1983, respectively. These were obviously higher than the overall average response of 27.0, 27.0 and 20.8%, for the three trials, respectively. Entry 27 exhibited a response of 33.9, 39.0 and 26.0% for all the three trials, respectively, which were again, greater than the average over all entries in the respective trials. This clearly demonstrates that neither Entry 3 nor 27 would have been selected for superior yielding ability when grown in narrow rows. Entries showing patterns of response similar to these entries would have been missed out if selection were practised under wide rows only, for yield production in narrow-row culture. However, the superiority of a line in narrow rows could not be predicted as no yield-related plant trait was found unique to these lines. Simple correlation coefficients between wide and narrow-row plots were highly significant for seed yield and all other plant characteristics measured in all trials (Table 4), indicating that, the average general performance of the entries was similar in wide and narrow rows. Selection of traits for better performance in wide rows could be applied to those entries to be used in narrow-row culture, although some exceptions existed, such as Entries 3 and 27, as discussed. High genotypic correlation between wide and narrow rows was reported by Weaver and Wilcox (1982). Correlation coefficients between yield in narrow-row plots and plant characteristics measured in wide-row plots are shown in Table 5. As expected, yield was certainly highly significantly correlated between the two row spacings. None of the traits measured in wide rows showed good correlation with yield in narrow rows in all trial. Days to canopy closure and number of seeds per pod measured in wide rows, however, showed negative an positive correlation, respectively, with
PERTAN1KA VOL. 11 NO. 3, 1988
IDENTIFICATION OF SOYBEAN GENOTYPES IN WIDE ROWS FOR NARROW-ROW CULTURE TABLE 4 Simple correlation coefficients between wide and narrow-row plots for yield and other soybean plant characteristics, in three environments. Characteristics Arlington
Yield * Plant height*1 Lodging Ht. of lowest pod No. of branches Maturity Canopy closure
Correlation coefficients Janesville
1982
1982
1983
0.62**
0.69**
0.58**
0.84**
0.81**
0.84**
0.81**
0.80**
0.67**
0.59**
0.62**
0.33**
0.63**
0.59**
0.39**
0.97**
0.97**
0.81**
0.94**
0.85**
0.90**
0.43**
0.58**
0.49**
Days to flowering Leaf area Seeds per pod 100-seed weight
Arlington
0.53** 0.79**
0.74**
0.77**
0.89**
0.92**
0.92**
0.57**
0.63**
0.56**
Seed quality ! Significant at 0.01 probability level.
TABLE 5 Simple correlation coefficients between soybean yield in narrow rows and other plant characteristics measured in wide-row plots, in three environments. Characteristics measured
Correlation coefficient with yield
in wide rows
in narrow rows Arlington 1982
Janesville 1982
Arlington 1983
Yield
0.62**
0.69**
0.58**
Plant height
0.11
0.03
- 0.09
- 0.13
- 0.28
- 0.19
Lodging Ht. of lowest pod
0.26*
0.22
0.?.4*
No. of branches
- 0.23
- 0.22
- 0.20
Maturity
- 0.12
- 0.01
0.04
Canopy closure
- 0.35*
- 0.32**
- 0.06
Days to flowering
- 0.14
0.12
- 0.11
Leaf area Seeds per pod
0.01 0.30**
0.33**
0.18
100-seed weight
- 0.12
- 0.13
- 0.23
Seed quality
- 0.15
0.02
- 0.05
1
Significant at 0.05 and 0.01 probability levels, respectively. PERTANIKA VOL. 11 NO. 3, 1988
339
GHIZAN SALEH AND EARL T. GR1TON
yield in narrow rows only in 1982, but no correlation was exhibited in 1983. No plant characteristic was thus, found to be of absolute predictive value that could be measured in wide rows for superior narrow-row performance. CONCLUSION Selection for superior performance in wide-row nurseries, in general, will identify lines that will perform best in narrow rows. Some lines with superior yielding ability in narrow rows, however, may not be identified from wide row nursery trials. Key traits that could be measured in wide rows and would predict superior performance in narrow rows were not identified. This factor varied with environments. No separate selection programme or breeding nursery is thus, suggested for narrow-row culture, because wide-row nurseries allow better crop management. REFERENCES COCHRAN, W.G. and G.M. COX. (1957): Experimental designs. John Wiley and Sons, Inc., New York, N.Y. 611pp. COOPER, R.L. (1974): Solid seeding to maximize soybean yields. Soybean Digest. 34: 12-14. COOPER, R.L. (1976): Modifying morphological and physiological characteristics of soybeans to maximize yields, p. 230-236. In: Hill, L.D. (ed.). World soybean research. Interstate Printers and Publishers, Inc., Danville, IL. COOPER, R.L. (1977): Response of soybean cultivarsto narrow rows and planting rates under weed-free conditions. Agron. I 69: 89-92. COSTA, J.A., E.S. OPLINGER and J.W. PENDLETON. (1980): Response of soybean cultivars to planting patterns. Agron. / 72: 153-156.
340
GREEN, D.E., P.F. BURLAMQUI and R.M. SHIBLES. (1977): Performance of randomly selected soybean lines with determinate and indeterminate growth habits. Crop Sci. 17: 335-339. HARTUNG, R.C., J.E. SPECHT and J.H. WILLIAMS. (1980): Agronomic performance of selected soybean morphological variants in irrigation culture with two row spacing*. Crop. Set 20: 604-609. HICKS, D.R., J.W. PENDLETON, R.L. BERNARD and T.J. JOHNSON. (1969): Response of soybean plant types to planting patterns. Agron. J. 61: 290-293. LEHMAN, W.F. and J.W. LAMBERT. (1960): Effects of spacing of soybean plants between and within rows on yield and its- components. Agron. J. 52: 84-86. METZ, G.L., D.E. GREEN and R.M. SHIBLES. (1984): Relationships between soybean yield in narrow rows and leaflet, canopy and developmental characters. Crop Set 24:457-462. PETERS, D.B. and L.C. JOHNSON. (1960): Soil moisture use by soybeans. Agron. J. 52: 687-689. SHAW, R.H. and C.R. WEBER. (1967): Effects of canopy arrangements on light interception and yield of soybeans. Agron. J. 59: 155-159. SHIBLES, R.M. and C.R. WEBER. (1966): Interception of solai radiation and dry matter production by various soybean planting patterns. Crop Sci. 6: 5559. TAYLOR, H.M. (1980): Soybean growth and yield as affected by row spacing and seasonal water supply. Agron. J. 72: 543-546. WEAVER, D.B. and J.R. WILCOX. (1982): Heritabilities, gains from selection, and genetic correlations for characteristics of soybeans grown in two row spacings. Crop Sci. 22: 625-629. WEBER, C.R., R.M. SHIBLES and D.E. SYTH. (1966): Effect of planting population and row spacing on soybean development and production. Agron. J. 58: 99-102.
PERTAN1KA VOL. 11 NO. 3, 1988
(Received 13 July, 1988)
Pertanika 11(3), 341-344 (1988)
Seed-borne Infection and Development of Colletotrichum capsici in Naturally Infected Chili Seed SARIAH MEON & W. ZAINUN W. NIK
Department of Plant Protection Faculty of Agriculture Universiti Pertanian Malaysia 43400 UPMy Serdang, Selangor, Malaysia Key words: Colletotrichum capsici; chili seed; site of infection. ABSTRAK Pengujian biji benih menggunakan kaedah kertas serap dan agar dekstros menunjukan penjangkitan Colletotrichum capsici yang jelas di dalam dan di permukaan luar biji benih cili. Hirisan microtome menunjukkan miselium didapati dalam lapisan luar dan dalam kulit dan dalam endosperma. Pembentukan aservulus dimulakan di bawah kulit biji benih dan juga di kawasan endosperma dan keluar dengan memecahkan kulit biji. Tisu-tisu parenkima juga bertukar bentuk. Patogen akhirnya hidup di atas permukaan biji benih. ABSTRACT
The testing of seeds using both the blotter and potato dextrose agar showed infection of Colletotrichum capsici to be well established both within and on the external surfaces of chilli seeds. Microtome sections showed that mycelia were present in the outer and inner layers of the testa and in the endosperm region. Formation of the acervulus was initiated below the seed coat and also in the endosperm and emerged to the surface after disrupting the seed coat. Parenchymatous tissues were also distorted. The pathogen finally grows on the seed surface.
INTRODUCTION
Anthracnose or ripe fruit rot of chillies (Capsicum annuum, L) caused by Colletotrichum capsici (Syd.) Butler & Bisby has been found to be a serious disease in the chilli growing areas of Malaysia. The disease attacked the leaves, stems, flowers and was most damaging on mature ripe fruits. Disease symptoms were occasionally observed on green fruits as well. The first indication of the disease is the appearance of small, elliptical or oblong straw-coloured, slightly sunken lesions on the surface of the mature fruit. This is followed by the development of black acervuli arranged in concentric rings, giving a target board appearance. Current observations in Peninsular Malaysia shows that the reduction in yield of marketable fruits due to anthracnose generally ranges from 10%60% depending on certain seasons of the year (Mah, 1987).
The pathogen persists in the soil, infected crop residues and weeds (Saxena etal. 1982). Several workers reported that Colletotrichum capsici was seed-borne (Higgins, 1930; Grover and Bansod, 1970; Mordue, 1971; Rout and Rath, 1972; Holliday, 1980). Hence, the use of a-rithracnose infected seeds can give rise to weak seedlings which then become the primary inoculum source in a chilli field. From the primary inoculum source, spread of the disease can be through windborne spores or through rain splash. Although, control of C. capsici affecting foliage, branches and fruits of chillies had been well documented, no attention had been paid so far to the determination of the seed-borne nature of C capsici. Hence this study is initiated to determine the site of infection and subsequent development in the infected seed.
SARIAH MEON
MATERIALS AND METHODS
Chilli seeds used in this study were collected from diseased fruits of the variety MC-4 obtained from an experimental field in Universiti Pertanian Malaysia. Four hundred seeds were used for each method of isolation. External seed-borne infection was determined by directly plating the seeds on moist blotter. Internal seed-borne infection was determined by surface - sterilising the seeds with 10% chlorox for 3 minutes, followed by repeated washings with sterilised water and plating on potato dextrose agar. The plates were incubated for seven days at 25 C in alternating cycles of 12hr near ultraviolet-light (NUV) and darkness, and examined for fungal infection using a stereo and a compound microscope. The experiment was arranged in a completely randomised design and the mean percentage of isolation was based on 400 seeds. On the fourth day of incubation, naturallyinfected seeds were fixed in 3% glutaraldehyde in O.O25M phosphate buffer and vacuum extracted for half an hour. They were then dehydrated in eight graded alcohol series (30, 40, 50, 60, 70, 80, 90 and 100 % ethanol) and embedded in gelatin capsules with JB-4 embedding mixture (Polysciences, Inc. , Warington, PA 18976). Sections were cut on an ordinary rotary microtome and
stained with 0.1% toluidine blue in 0.1% sodium borate (O'Brien efa/, 1964). In addition, scanning electron microscopy was also used as an additional tool in this study. Seed samples were fixed in 6% buffered glutraldehyde, washed three times with phosphate buffer and post-fixed with 1% osmium tetroxide. The samples were then dehydrated in acetone series and dried in a Polaron Critical Point drying apparatus. The dried specimens were stuck to copper stubs and shadowed with gold in a Polaron Sputter Coater. Micrographs were taken in a JOEL JSM 35C Scanning electron microscope. RESULTS AND DISCUSSION
Colletotrichum capsici was isolated most frequently both by the blotter and potato dextrose agar, suggesting that the pathogen could be present on the seed surface as well as inside the seed (Table 1). Similar results were reported by Grover and Bansod (1970) and Rout and Rath (1972) on chilli seeds. In addition, eight genera of fungi were also isolated (Table 1). Rhizopus stolonifer, Chaetomium globosum and Pestalotiopsis sp. were isolated from chilli seeds and represent new records of chilli seed-borne infection. Transverse sections of seeds naturally infected by Colletotrichum showed a number of
TABLE 1 Percentage isolation of fungi from chilli seeds using the blotter method and potato-dextrose agar
* Percentage isolation * Blotter method
Potato-Dectrose Agar method
Colletotrichum capsici (Syd.) Butler & Bisby
37
29
Fusarium spp.
25
31
Aspergillus spp.
17
3
Drechslera rostrata (Drechsl.) Richardson & Fraser
16
19
3
11
Colletotrichum gloesporiodes (Penz.) sacc
0.3
0
Chaetomium globosum Kze
0.4
0
Rhizopus stolonifer (Ehrenb. ex Fr.) Lind
1.3
0
0
7
Curvularia lunata (Wakker) & Boedijin
Pestalotiopsis sp.
342
PERTANIKA VOL. 11 NO. 3, 1988
SEED-BORNE INFECTION AND DEVELOPMENT OF C. CAPSICI IN NATURALLY INFECTED CHILI SEED
acervuli over the seed surfaces. After infection, acervulus primordia were formed beneath the inner layers of the testa and within the endosperm region (Plate 1). Subsequently, the young acervulus enlarged and caused the testa to bulge outwards by exerting pressure on the testa (Plate
21
Plate 1.
of ooze. Mycelia were also detected in the endosperm region (Plates 4Af B) just below the inner testa. The mycelium could grow directly into the inner layer of the testa surrounding the endosperm, or could remain dormant until the seed germinated. Infected parenchymatous tissues of the seed appeared distorted.
Acervulus primordia (arrowed) beneath seed coat (Sc • seed coat; Ap - Acervulus primordia; P- parenchyma; e - endosperm).
Plate 3.
Plate 2.
Acervulus primordia emerging through seed coat (YA - young acervulus; AP - acervulus primordia)
Ultimately, the testa ruptured and the young acervulus emerged. Each mature acervulus bore numerous setae and masses of conida (Plates 3Ay B). On bursting, conidia were liberated in the form
(A) Tranverse section, (B) Whole mount (SEM micrograph) of mature acervulus showing numerous setae. (S • setae; MA • mature acervulus)
It is evident from these studies that C capsici was borne intraembryonal in chilli seeds. Disruption of seed tissues could be due to the activity of cellulolytic and pectionlytic enzymes produced by C. capsici (Sariah, 1980). Spores oozing out from acervuli can serve as a primary inoculum source for the spread of the disease. The fungus can then spread from the seed to the placenta of the fruit. The pathogen could then penetrate the developing ovules, or young seed with unlignified testa at any point on their surface. Infection of seeds could also occur directly from
PERTANIKA VOL. 11 NO. 3, 1988
343
SARIAH MEON (Neergaard, 1979). Therefore, with internal seedborne infection as was the case here, systemic fungicides will provide a more practical and effective control measure. Besides anthracnose infected seeds should not be used for planting. REFERENCES GROVER, R.K. and R.D. BANSOD. (1970): Seed borne nature of Colletotrichum capsici in chilli seeds and its control by seed dressing fungicides. Indian Phytopathol., 23:664-668. HIGG1NS, B.B. (1930); A pepper fruit rot new to the United States Georgia Expert. Stat. Bull, no. 162, 10 pp. Rev. Appl. Mycol. 10: 56-57 (Abstract). HOLLIDAY, P. (1980): Fungus Disease of Tropical crops. Cambridge University Press. 607 pp. MAH, S.Y. (1987): Anthracnose fruit rot of chilli (Capsicum annuum. L): Some aspects of its etiology, epidemiology and control in Peninsular Malaysia. M. Agric. Sc. Thesis. Universiti Pertanian Malaysia. MORDUE, J.E.M. (1971): Colletotrichum capsici CM.1. Description of pathogenic fungi and bacteria No. 317, Commonwealth Mycological Institute, Kew, NEERGAARD, P. (1979): Seed Pathology vols. 1 & 2. 2nd edition. The Macmillan Press Ltd. London. 840 pp & 843 pp. respectively.
Plate 4
Transverse section of infected seed showing the presence of hyphae in the endosperm region (A) light microscopy and (B) SEM micrograph (H - hyphae}.
the mother plant: through the pedicel, funiculus or integuments into the developing ovule. Also, conidia of C. capsici could also be mechanically attached to the surface of the testa, remaining dormant until the seed germinated. With seed-borne inocula, the way in which the inoculum was carried within or on the seeds influenced the type of control measure used
344
O'BRIEN, T.P., N. FEDER and M.E. MCCULLY. (1964): Polychromatic staining of plant cell walls by toludine blue. Protoplasma 59: 368-373. ROUT. R.K. and G.C. RATH. (1972): Note on seedborne desease of chilli (Capsicum annuum, L.) Indian Phytopathol. 25; 5 97-598. SARIAH, M. (1980). Pectinolytic and cellulolytic enzyme production by Colletotrichum capsici. (Syd,) Butler &Bisby. Malay. Appl Biol 9: 105-110. SAXENA, A.K. LJ. HOSHI and J.B. SAXENA. (1982): Additions to the host range of Colletotrichum capsici. India Phytopathol. 35: 166.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 29 April, 1988)
Pcrlanika 11(3), 345-348(1988)
The Induction and Evaluation of Productive Semidwarf Mutants of Basmati-370 M. SIDDIQUE SAJJAD and MA. AWAN Nuclear Institute for Agriculture and Biology (NIAB) Jhang Road, Faisalabad, P.O. Box 128, Pakistan. Key words: Induction; evaluation; productive semidwarf mutants; Basmati-370 ABSTRAK Bifi benih Basmati 370 yang kering dan seragam, dengan kandungan kelembapan lebih kurang 14% telah didedahkan kepada pancaran sinar gamma fdari sumber 60 Co) dosnya 0, 15, 20 dan 25 kR. Dua puluh satu muatan separuh kerdil telah dipilih dari 36,000 tumbuhan M2 yang berasal daripada lebih kurang 12,000 tumbuhan ML Perilaku pembaikbiak mutan-mutan tersebut telah dipilih di peringkat M3 dan kesudahannya 6 mutan telah dipilih. Ujian kadar hasil perbandingan mutan-mutan tersebut dengan induk kultivar Basmati-370, telah menunjukkan yangjenis mutan berpotensi mengeluarkan hasil yang tinggi dan mempunyai bentuk separuh kerdil Sifat-sifat lain dan juga trait fizikokimia mutan-mutan tersebut dan kultivar induk telah juga dibincangkan. ABSTRACT Dry, uniform seeds of Basmati 370 with about 14% moisture content were exposed to 0, 15, 20 and 25kR doses of gamma rays (60Co source). Twenty one semi-dwarf mutants were selected from 36,000 M2 plants originating from about 12,000 Ml plants. The breeding behaviour of these mutants was studied in M3 and consequently 6 mutants were selected. The comparative yield test of mutants with mother cultivar Basmati-370, established the high yield potential and semi-dwarf plant posture of mutant strains. The other plant attributes as well as physico-chemical traits of the mutants and parent cultivar are also presented. INTRODUCTION Basmati rices have been prized for their good cooking quality and aroma in the sub-continent of Indo-Pak from time immemorial. From these Basmati rices, Basmati 370 was isolated from the local germplasm and is currently cultivated predominantly in the province of Punjab. Though the cultivar is matchless in cooking quality and aroma, it possesses two detrimental aspects which are: (i) tall growing habit and (ii) weak straw. Both these attributes make the cultivar incapable of responding to fertilizer or withstanding lodging. The importance of short culm length and maximum yield potential is now readily recognized. But the hybridization with the world famous Dee-Gee-Woo-Gen dwarfing gene source may not accomplish the arduous task of rectifying the two
set-backs inherent in basmati rice. The mutagenesis approach may be the pragmatic approach to accomplish the job successfully. The technique may provide an alternative as it is a less disruptive approach inducing fresh variability into a well adapted local germplasm of basmati rices. The earlier strifes have been fruitful in the induction of short stature (Malik, 1982; Marie, 1981; Misra, 1979 and Sajjad, 1984), lodging resistance (Rutger et at., 1977) with high yield potential (Sajjad, 1984 and Rutger, 1982) and maintaining the original aroma of Basmati 370 (Ready etal, 1975). A large number of cultivars has already been developed using induced mutants as gene donors for desireable traits. A high frequency of recovering the semidwarf progenies resulting from hybridizing the induced semidwarf mutant lines
M. SIDDIQUE SAJJAD AND M.A. AWAN
have also been reported (Marie, 1981 and Reddy, 1975) which illustrates the even greater potential of mutants as parent for hybridization. In 1980, the Nuclear Institute for Agriculture and Biology (NIAB), Faisalabad, Pakistan embarked on this study to induce semi-dwarf mutants of Basmati 370. MATERIALS AND METHODS
Healthy and uniform seeds of Basmati 370 with moisture contents of about 14% were exposed to 9, 15, 20 & 25 kR doses of gamma rays from a Co source of NIAB 1979-80. The 15, 20 & 25 kR treatments consisted of 5000 seeds each. The transplantation was accomplished by using single seedling per hill, at a ear to row distance of 15 cm. On maturity of the crop, first three emerging panicles per plant were harvested and bulked dose wise. M2 generation was grown at a plant to row distance of 20 cm using a single seedling per hill during 1981-82. The plants of Basmati 370, grown from unirradiated seed as control, were transplanted after every 20th row of M2 plants. Some semi-dwarf mutants (DM) were selected and their breeding behaviour was studied during 1982-83. The true breeding mutants were selected
and yield tested in a micro yield trial (Gross plot size 8m 2 /entry) during 1983-84. The micro yield trial was conducted using a randomized complete block design with four replications. Single seedling per hill was transplanted with a plant to row distance of 20 cm. The crop was fertilized at the rate of 80:40:0, N:P:K, kg/ha respectively. The physico-chemical traits were studied and the data were statistically analysed. EXPERIMENTAL RESULTS
The breeding behaviour of the mutants selected from M2 generation was studied in M3 during 1982-83. The results are presented in Table.l. It is evident from the data that all the mutants were significantly shorter than the parent cuitivar Basmati 370 by about 16% to 19%. The performance of these semi-dwarf mutants for number of productive tillers per plant, number of days to flowering, panicle length, panicle fertility and yield per plant was not significantly different from that of Basmati 370. For Basmati 370, DM18, DM20, DM22, DM24 and DM25, number of grains per panicle was significantly higher than the rest of the mutants.
TABLE 1 Performance of semidwarf mutants and parent cuitivar Basmati-370 in M 3 (1982-83)
Name of mutant variety
Plant height reduction (%) over Bas. 370
No. of productive tillers/ plant
No. of grains/ panicle
Panicle fertility (%)
Yield plant (g)
-
11.7a
127.1a
93.4a
21.2a
DM18 (15kR)
16.0
11.6a
124.0ab
95.3a
20.8a
DM20 (15kR)
16.3
11.4a
120.6abc
95.0a
19.9a
DM22 (15kR)
16.4
11.3a
119.9abc
94.5a
19.5a
DM24 (20kR)
17.1
11.2a
118.4abcd
94.9a
19.3a
DM25 (20kR)
17.3
11.0a
116,4abcd
95.0a
15.3a
DM38 (25kR)
18.6
10.0a
110.3cd
93.8a
18.0a
Basmati 370
Figures followed by different letters are significant at 5% level of significance according to DMRT. 346
PERTANIKA VOL. 11 NO. 3, 1988
THE INDUCTION AND EVALUATION OF PRODUCTIVE SEMIDWARF MUTANTS OF BASMATI-370 TABLE 2 Performance of semi-dwarf mutants and parent variety Basmati-370 in a micro yield trial
Name of mutant/ variety
Plant height (cm)
No. of productive tillers/ plant
No. of grains/ panicle
Panicle fertility %
Yield (kg/ha)
Basmati 370
177.0a
9.3b
112.3a
85.3a
3564.0b
DM18 (15kR)
132.8b
10.9a
121.6a
89.1a
4464.0a
DM20 (15kR)
133.2b
11.8a
119.5a
88.9a
5085.0a
DM22 (15kR)
134.1b
11.3a
127.3a
89.1a
5154.0a
DM24 (20kR)
132.8b
11.8a
127.4a
89.6a
5373.0a
DM25 (20kR)
131.5b
11.3a
124.3a
89.5a
5245.0a
DM38 (25kR)
131.8b
12.0a
118.0a
89.3a
5250.0a
Figures followed by different letters are significant at 5% level of significance according to DMRT.
TABLE 3 Physico-chemical traits of rice mutants variety
Variety/ mutant
Length (mm)
Width (mm)
Length width
Quality Index
Elongation ratio
Amy lose %
Aroma
Basmati-370
6.9
1.8
3.8
2.3
1.8
22.8
3.2
DM-18
6.7
1.7
3.9
2.4
1.8
23.3
3.0
DM-20
6.7
1.7
3.9
2.3
1.9
22.9
3.0
DM-22
6.7
1.7
3.9
2.3
1.8
23.1
3.0
DM-24
6.8
1.8
3.8
2.3
1.9
23.5
3.5
DM-25
6.6
1.7
3.9
2.4
1.9
22.8
3.4
DM-38
6.6
1.7
3.9
2.4
1.8
23.2
3.0
The results of micro yield trial conducted during 1983-84 are presented in Table 2. It is evident from the data that all mutant lines were significantly shorter in height than Basmati 370. They were 24% to 26% shorter and had significantly higher number of productive tillers per plant. The performance of mutant strains for the plant
attributes of panicle length and thousand grain weight was significantly inferior to that of parent. The mutant strains were similar in performance for the plant attributes of number of grains per panicle and panicle fertility percent. The mutant strains surpassed Basmati 370 in yield potential by about 25-51%. The plant attributes which
PERTANIKA VOL. 11 NO. 3, 1988
347
M. SIDDIQUE SAJJAD AND M.A. AWAN
have enhanced the yield potential of mutant lines seem to be a higher number of productive tillers per plant. Such results have also been reported in rice, (Sajjad, 1984; Dat, 1978 and Rangaswamy, 1983) . The results of physicochemical traits of mutants and parent indicate (Tabel 3) that some characteristics of the mutants were at par while others were superior to those of the parent variety, Basmati-370. For instance, the mutants DM-24 and 25 had a stronger aroma than the parent. It may safely be concluded from the present study that all the mutants exhibited the property of semi-dwarf plant posture along with a higher yielding capability. They may be used as gene sources for short culm for basmati rices. The emphasis in future studies will be to isolate the number of mutants to avoid the potential risk of genetic vulnerability of semidwarf mutants and to use these as gene source for stem shortening of Basmati 370.
348
REFERENCES
DAT T.V., M.L. PETERSON, J.L. RUTGER. (1978): Crop Set 18:1. MALIK S.S. (1982): Internat. Rice Res. Newslett. 7:4. MARIE R. (1981): Mutation Breeding Newslett. 18:2. MISRA R.N. (1979): Proc. Symp., The Use of Induced Mutations in Crop Improvement, Hyderabad, India. RANGASWAMY S.R., C.R. ANANDAKUMAR. (1983): International Rice Res. Newslett. 8:4. REDDY T.P, A. PADMA, G.M. REDDY. (1975): Indian J. Genetics 35:31. RUTGER J.N., M.L. PETERSON, C.H. HU. (1977): Crop Set 17:978. RUTGER J.N., H.L. CHARNAHAN, CW. JOHNSON. (1982): Crop Sci. 22:164.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 9 April, 1988)
Pertanika 11(3), 349-355 (1988)
Studies on Some Sooty Moulds on Guava in Malaysia LIM TONG KWEE
Department of Plant Protection Faculty of Agriculture, Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia ABSTRAK
Kulat jelaga membiak dengan banyak di atas daun jambu batu, mendapat makanan dari bahan manisan yang dikeluarkan oleh serangga teritip, kutu daun, Mat putih dan koya. Sembilan spesies, lima spesies dari Ascomyetes iaitu: Phragmocapnias betle, Scorias philippensis, Trichomerium grandisporum, Limacinula musicola dan Aithaloderma clavatisporum, serta empat genus dari Deuteromycetes iaitu: Tripospermum sp., Polychaeton sp., Leptoxyphium sp. dan Conidiocarpus sp. adalah dihuraikan dan di~ laporkan sebagai laporan terbaru di Semenanjung Malaysia.
ABSTRACT
Sooty moulds proliferate in abundance on the foliage of guava, subsisting on the honeydew secreted by scale insects aphids, whiteflies, and mealy bugs. Nine species, five Ascomycetes species namely, Phragmocapnias betle, Scorias philippensis, Trichomerium grandisporum, Limacinula musicola and Aithaloderma clavatisporum; and four Deuteromycetes genera viz. Tripospermum sp., Polychaeton sp., Leptoxyphium sp. and Conidiocarpus sp., are described and reported as new records on guava for Peninsular Malaysia. INTRODUCTION
Sooty moulds are a commonsight on the leaves, twigs and sometimes fruits of man/ tropical fruit crops (Lim and Khoo, 1985). They form thin, black, papery films; black, velutinous growth, black pellicles, or psuedoparenchymatous crusts. Such growths represent unit communities of an assemblage of several diverse species of fungi belonging to the Deuteromycetes and Ascomycetes with dark-coloured somatic and fruit-body cells, living together saprotrophically in apparent harmonious and mutualistic association. Their ubiquity and abundant proliferation on such micro-habitats can be attributed to their adaptability to grow and reproduce in the high stress environment of the crop foliage. In accordance with Grime's ecological terminology (1979), they can be termed as stress-selected or S-selected fungi. Such fungi have culminated in a stresstolerant strategy for survival involving the development of adaptations which facilitate endurance of conditions of continuous environmental stress as
are experienced on the foliage. They can also be deemed as true foliage residents. Their diversity, temporal and spatial distribution on such niches depend on the supply of nutrients, availability of moisture, and thermal and radiation fluctuations. The sooty moulds obtain their water from free water films or water droplets on leaves from the rain or dew, atmospheric water vapour, and water exuded via guttation. They can obtain their nutrition from leaf diffusate or guttation fluid (Tukey, 1971); the cuticle which provide a rich and potential source (Baker, 1971, Holloway, 1971); honey dew secretions from insects such as scale insects, aphids, whiteflies and mealybugs; chemical contaminated rain; organic and inorganic dust particles; pollens; and spores of other microflora. Thus, they can be found on the plant foliage with or without the presence of the above-mentioned insects. However, on guava (Psidium guajava L.), sooty moulds are usually found in association with insects, subsisting on the rich, nutritive nabulum of the honey dew.
LIM TONG KWEE
Despite their ubiquity and plenitude, their significance is usually overlooked. Many of the hyphae of the sooty moulds are mucilaginous, absorbing moisture readily and maintaining a moist surface for prolonged periods. This creates a very humid environment within the tree canopy, facilitating the establishment and spread of other fungal diseases. Extensive growth of the sooty mould on the leaves can reduce the photosynthethic activity of the leaves and adversely affects the normal flowering and fruiting of the tree. Besides, trees heavily colonized by these fungi exhibit a decrease in growth vigour and tend to fruit poorly. On fruits, sooty moulds spoil the cosmetic appearance of the fruits and reduce their marketable value (Lim and Khoo, 1985). Information and studies on sooty mould of fruit trees in Malaysia are rather meagre and scanty, particularly on guava. In Peninsular Malaysia, only one sooty mould was listed on guavaTrichopeltis pulchella Speg. (Johnston, 1960). In Sarawak, four species namely : Atichia glomerulosa (Ach.) Flotow, Capnodium moniliforme Fraser, Capnophaeum sp. and Phaeochaetia annonicola (Hansf.) Bat. & Cif. were recorded by Turner (1971). In Sabah, Williams and Liu (1976) recorded three species viz. Aithaloderma clavatisporum Syd., Caldariomyces fumago Woron and a Chaetothyrium sp. This paper reports and describes nine previously unrecorded species of sooty mould found on guava in Peninsular Malaysia. The specimens are kept at the Plant Pathology Herbarium, Department of Plant Protection, Universiti Pertanian Malaysia at Serdang, Selangor.
attachment. Identifications were based largely on the recent concepts proposed by Reynolds (1979, 1982) and also on some of the concepts of Hughes (1976). Descriptions in the literature especially of Hansford (1946), Hughes (1951, 1976) and Reynolds (1971, 1978, 1979, 1982) were utilized in aiding specimen identification and in understanding the taxonomical position of the species concerned. Isolation of non-sterilized sooty mould mycelia and spores was also attempted on potato dextrose agar (PDA) and malt extract agar (MEA). No attempts were made to demonstrate holomorphic pleomorphy i.e. the relationship of any anamorphosis to a teleomorphosis.
Fig. la; Sooty mould occurring as disrete, spongy, velvety, black spots on the guava leaf.
MATERIALS AND METHODS
Observations including morphometric measurements were made on all possible taxonomically significant characters. Tissues of sooty moulds were gently removed from guava leaf surfaces (Fig. la & b) and mounted in lactophenol clear on glassslides and sealed with nail polish. Colloidin impressions of sooty mould in-situ on the leaves were made, mounted on glass slides and the colloidin materials were subsequently removed with acetone. Free-hand sections mounted in lactophenol clear or lactophenol cotton-blue were also prepared. The slides were studied and microphotographed using the Lietz Orthoplan fitted with a Nomarskii interference contrast 350
Fig. lb: Sooty mould occurring as a thin papery, black film on guava leaves (arrowed). RESULTS AND DISCUSSION
The following species of sooty mould listed and described below represent new records of sooty mould for Psidium guajava (guava) in Peninsular Malaysia.
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ascocarp in the latter (Reynolds, 1979). On guava, Phragmocapnias can be found together with Leptoxyphium, Conidiocarpus and Polychaeton in mixed colonies. Scorias philippensis Mendoza emend. Reynolds
KHU
Fig. 2:
mm.
I
Stalked, setose, subglobose ascocarp of Phragmocapnias betle. Arrow denotes ascospores inascus. Bar = 18\Jm.
Phragmocapnias betle (Sydow & Butler) Theissen & Sydow emend. Reynolds. This is a stalked capnodiaceous species with brown mycelia, epiphyllous and pelliculose on the leaf surfaces. Hyphae are dark brown, reticulate with cylindrical cells slightly constricted at the septa and somewhat mucilaginous. Mycelial setae are black, with a rounded, blunt tip and measure on the average 4iO fjtxn by 75.6 fjm at its broad base. Ascocarp are subglobose, blackish-brown, setose, unilocular and are stalked (Figure 2). The setae are acuminate, 53.6-107.2 jum long, found usually on the upper half of the ascocarp. The ascocarps measure 75-165 x 70-125 jjm (mean 107 x 112.6 jum). The asci bitunicate with eight hyaline ascospores. The ascospores are elliptical with blunt acute apices, 16—18.76 x 3.2—4.8 jum and have three transepta. Phragmocapnias was established by Spegazinni in 1918 as a new genus on the basis of Capnodium betle Syd. & Butl. Reynolds (1979) emended the species to P. betle and listed the following species (teleomorphosis only) to be synonyms:- Antennellopsis elegans Bat. & Cif., A. formosa Bat. & Clif., A. vulgaris (Yamamoto) Bat. & Cif., Antennella citri Swada, Capnodium tanakae Shirae and Hara emend. Sawada, Neocapnodium tanakae (Shirae and Hara) Yamamoto, Scorias communis Yamamoto, S. cylindrical Yamamoto, S. vulgare Yamamoto, Chaetoscorias vulgaris Yamamoto, Trichomenum jambosae Bat. &Cif. Phragmocapnias can be distinguished from species of Scorias on the absence of setae on the
The fungus forms a black, epiphyllous, spongy subiculum on guava leaves. The hyphae are more or less cylindrical, 10.72 x 5.4 jzm, brown to dark-brown, mucilaginous and they anastomise to form a much-branched network. The ascocarp is dark-brown, subglobose (39-41 x 41-43 fjtm) to ellipsoidal, nonsetose and subtended on a columnar stalk, 10-15 jum high (Figure 3). The ascocarp is uniloculate and ostiolate at maturity, Asci are bitunicate and eight-spored. The ascospores are hyaline, clavate, tapering more at one end, 22 x 4.82 jum, with three transepta.
Fig, 3:
Stalked, nonsetose ascocarp of Scorias philippenses. Bar = 10.5 jJm.
On guava, Scorias can be found together with Polychaeton on the same mycelial network. Reynolds (1979) listed Hyalocapnias psidii Bat. & Cif. which was reported on guava to be synonymous. Trichomerium grandisporum (Ellis in Ellis and Everhart) Bat. & Cif.
and
Martin
This is a anon-stalked, capnodiaceous, foliicolous genus in the Ascomycetes, occurring as superficial, pelliculose, growths on the leaf surfaces. Mycelia are brown to olivaceous, much-branched with cylindrical hypae. Mycelia setae are blackish-
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brown with acuminate apices and 85-320 pn long. The ascocarp are subglobose to ampuliform, 75-102 x 91 160 jttm, olivaceous to dark brown, setose, apically ostiolate and unilocular (Figure 4a). The setae are 107-130 x 4 - 7 jum, darkbrown with rounded apices. The ascospores are hyaline, fusiform to elliptical with blunt acute apices, 18.6-24.12 /zm in length and with three transepta (Figure 4b).
Fig. 4;
Setose,
ampulliform,
sessile ascocarp of
Trichomerium grandisporum. Bar • 20 fJm.
Fig. 4b: Ascocarp of T. grandisporum crushed to show the asci (arrowed). Bar • 18 \Mn.
The genus Trichomerium was established in 1918 by Spegazzini with T coffeicola as the type species (Reynolds, 1982). After examining over a hundred species specimens cited in the literature and scores of additional field collections, Reynolds (1982) reduced the many names to a monotypic genus T. grandisporum with the basionym as Capnodium grandisporum Ellis and Martin in Ellis and Everhart. This species comprises fungi having a capnodiaceous centrum with hyaline, trans-septate ascospores in a sessile, 352
setose ascocarp and the synonyms listed in chronological sequence by Reynolds include :Limacinia grandispora (Ellis and Martin) Arnaud, Limacinia coffeicola Puttemans, Trichomerium coffeicola (Puttemans) Spegazzini, Limacinia spinigera Hohnel, Aithaloderma spinigera (Hohnel) Hohnel, Phaeochaetia spinigera (Hohnel) Batista and Ciferri, Chaetothyrium spinigerum (Hohnel) Yamamoto, Triposporiopsis spinigera (Hohnel) Yamamoto, Limacinia imperspicua Saccardo, Phragmocapnias impersicua (Saccardo) Batista and Ciferri, Trichomerium postericense Spegazzini, Chaetothyrium syzgii Hansford, Phaeochaetia ansyzigii (Hansford), Batista and Ciferri, Chaetothyrium annonicola Hansford, Phaeochaetia annonicola (Hansford) Batista and Ciferri, Trichomerium abhorrens Batista, Trichomerium psidii Batista, Trichomerium hirtellum Batista, Chaetopotius commistus Batista, Aithaloderma citri (Sawada) Sawada, Setella citricola Batista and Peres, Gilmania xylopia Batista, Nascimento and Ciferri, Vitalia averrhoae Batista, Vitalia cercropia Batista, Vital and Ciferri, Vitalia jaboatonensis Batista, Nascimento and Ciferri, Vitalia multiseptata Nascimento and Ciferri, Vitalia phormicola Bastita, Nascimento and Ceferri, Vitalia plumierae Batista and da Matta, Capnobatista serrulata Cifferri and Leal, Trichomerium abhorrens Batista var. coffea Batista and da Matta, Trichomerium crinosporum Batista and Cifferri, Trichomerium didymopanacis Batista and Ciferri, Trichomerium inconditum Batista and Cifferi, Trichomerium mangifericolum Batista and Cifferi, Trichomerium ornatum Batista and Ciferri. Although the ascospores of T. grandisporum are similar to those of Phragmocapnias and Scorias in septation and the absence of pigmentation at maturity and before germination, it can be differentiated from the latter two by the presence of the sessile ascocarp (Reynolds, 1982). Tripospermum sp. Speganizzi.
A dematiaceous Hyphomycetes fungus that occurs together with other sooty mould as a superficial, pellicullose or crust-like black growth on leaf surfaces (Figure 5). Hyphae are olivaceous to pale-brown, reticulate, consisting of long cylindrical cells 26 um long by 7 jam wide. Conidia are brown, star-shaped (stauroconidia) with four divergent arms which are wide at the base and
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STUDIES ON SOME SOOTY MOULDS IN GUAVA IN MALAYSIA.
tapering to a rounded apex. Each arm can have 4 - 8 cells i.e. 3 - 7 septa and are slightly constricted at the septa. Arms with 4 - 5 septa are the most common {Figure 5). Conidia can also be diradiate although triradiate or tetraradiate are the most common. Each conidium arises from a pyriform, stalked cell 5.5-8.8 /zm long. Each arm measures 26-75 /zm long by 7.65-10.5 jLtm at the base. The teleomorphosis is uncertain. The genus Tripospermum was established by Spegazzini in 1918 as a segregate of Triposporium and T. acerinum was the type species (Hughes, 195 l){Figure 5). Fig. 6:
Long-necked pycnidium of Polychaeton sp, with hyaline, ellipsoidal conidia (arrowed).
He suggested that Microxiphium aciculiforme Cif., Bat. and Nasc, M. coffeanum Bat., & Matta, M. pinicola Bat., Nasc, & Cif. and Astragoxyphium plumeriae Bat., & Matta should be placed in Polychaeton.
Fig. 5:
Tetraradiate stauroconidium of Triposperium sp. Bar = 15 (Jm.
Polychaeton sp. (Pers.) Lev. The broad anastomising hyphae are made up of irregular, cylindrical (18.76 x 4.8 /im), dark brown cells immersed in a mucilaginous matrix. Pycnidia produced have broad, robust simple or branched stalks which can reach lengths of 109 to 345 /im and widths of 22 /im at the stalk base and 28 /im at the broadest portion of the pycnidia. The pycnidium is extended into a subulate or cylindrical neck and terminates in a fringe of hyaline, subulate extensions around the ostiole {Figure 6). The pycnidial cavity is ellipsoidal but there is no obvious swelling in the stalk to indicate its position. Conidia extruded through the ostiole and gather in a terminal droplet. They are minute (1.88 x 3.8 /an), hyaline and ellipsoidal. Hughes (1976) designated Fumago quercina Pers. as the lectotype species of Polychaeton.
On guava, Polychaeton usually occurs together with Phragmocapnias betle and Scorias philippensis. Yamamoto (1954) showed that Polychaeton was connected to Neocapnodium tanakae which is a synonym of P-betle. However, Reynolds (1979) regarded Yamamoto's taxonomical studies on the Taiwan Fungi to be intuitive and unsupported by systematic experimental work which he purported to have undertaken. Hence, the teleomorphosis of Poly-chaeton is still uncertain, Conidiocarpus sp. Woronichin. This fungus was found on the same type of hypae in a spongy subiculum as Scorias. The species on guava produces a lon^ stalked 170-230 /im and short stalked, 48-110 jum pycnidia without a neck. The pycnidium can be seen as a terminal, oval to hemispherical (59 x 55 /im) swelling with an ostiole fringed with hyaline subulate extensions of the synematous hyphae. {Figure 7) The conidia are hyaline, ellipsoidal, minute, 5.36 x 1.6 /im. This form genus was proposed in 1917 for the single species of C caucasicus (Hughes, 1976). Yamamoto (1954) demonstrated that the pycnidia of the Conidiocarpus type were connected to
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L musicola was emended by Reynolds (1971) as a new combination and he listed 5 synonyms viz.: Phaeosaccardinula musicola Batista., Phaeosaccardinula guajavae Bat. & Vital, Phaeosaccardinula vera Bat. & Cif., Phaeosaccardinula guajavae Bat. & Vital var. citrina Bat. and Paracapnodium brevistipitatum Bat., Cif. and Maia. Aithaloderma clavatisporum H. & P. Sydow
Fig. 7:
Long stalked pycnidium of Conidiocarpus sp. Bar = 20 fim
Scorias commune Yamamoto which is a synonym of Phragmocaphias betle (Reynolds, 1979). Limacinula musicola (Batista) Reynolds. This foliicolous species grows saprotrophically with other fungal species on leaf surfaces. The fungus mycelia consist of brown hyphae made up of short to elongate-rectangular cells forming a thin subiculum and a pale brown hyphae extending from the fruit body wall, collectively forming a loose, distinctive weft. The fruit body is a psuedothecium, brown, epipellicle, nonsetose, sessile, globose (150-320 /im in diameter) when immature becoming collabent and ostiolate at maturity. Asci are bitunicate, form in a basal hymenium in the locule of the psuedothecium. Ascospores are muriform, hyaline to olivaceous, large, 36-62.5 x 12.6-24/zm, with a polystichous longisepta and 7-12 transepta. (Figure 8)
Fig. 8:
Close-up of the large, muriform ascospores of L. musicola. Bar = 12 fJim.
354
On guava, this sooty mould species produces brown, cylindrical-celled (8 x 4.02 um), mucilaginous hyphae which appear slightly constricted at the septa. No hyphal setae are produced. It produces brownish-black, subglobose to conical-globulose pycnidia which can measure up to 101-107.2 x 80.4-93.8 um. The latter pycnidia show some hyphae radiating from their base. Most of the pycnidia bear short, conical, dark-brown, thickwalled seta with tapering apices, measuring 8.0422.95 jttm; although some are devoid of setea (Figure 9). The pycnidia are usually produced in clusters. The conida are hyaline, minute, 3-4 x 22.5 jLtm, ellipsoidal and are produced in abundance. The teleomorphic ascigerous state was encountered with Leptoxyphium on some guava leaves infested with the spiral whiteflies. The ascocarps produced are subglobose (64 x 68 jzm) to domeshaped, brownish-black, ostiolate, uniloculate and have several (6) thick-walled, dark brown setae 11-27 jum long. The acid are bitunicate, and the ascospores are hyaline, clavate, 23 x 5 pern, with 3-5 septa.
Subglobose pycnidium of Aithaloderma clavatisporum with distinct, short conical seta (a) and minute, hyaline conidia (b).Bar = 12. 5 JAm.
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STUDIES ON SOME SOOTY MOULDS IN GUAVA IN MALAYSIA. Synonyms of A. clavatisporum are Phaeochaetia clavatispora (H. & P. Syd.) Hansf. Hughes (1976) listed the following as congeneric to A. clavatisporum; A. ferruginea Fraser, A, viridis Fraser, A. capensis Doidge (Phaeochaetia capensis Doidge) Bat. and Aithaloderma sp 1 (Herb. DAOM) which is the ascigerous state of Ciferrioxyphium chaetomorphum Speg.
BAKER, E.A. (1971): Chemical and physical characteristics of cuticular membranes, In Ecology of Leaf Surface Micro-organisms, (eds. T.F. Preece and C.H. Dickinson) pp. 55-65. Acad. Press, London and New York.
Leptoxyphium sp. Speg.
HANSFORD, C.G. (1946): The foliicolous Ascomycetes, their parasites and associated fungi. My col Pap. 15: 1-240.
This fungus forms an epiphyllous, black, effuse, pelliculous crust on leaf surfaces. The hypae are more or less sylindrical, brown, mucilaginous, slightly constricted at the septa and form a tightly anastomising network. It produces synemata which are made up of closely adpressed erect hyphae with a helical twisting in its axis {Figure 10). The synema is covered with a mucilaginous layer and is cylindrical or subulate, reaching lengths of 200 jum with a wide base. The apex terminates in a fringe of sterile hairs enclosing the opening through which the conidia are extruded. The conidia are broadly ellipsoidal, and hyaline when immature, but larger (5.36 x 2.68 /urn), may be one-septate and brown when mature.
REFERENCES
GRIME, J.P. (1979): Plant Strategies and Vegetation Processes. John Wiley, Chichester and New York.
HOLLOWAY, P.J. (1971): The chemical and physical characteristics of leaf surfaces. In Ecology and Leaf Surface Micro-organisms, (eds. T.F. Preece and C.H. Dickinson) pp. 39-53. Acad. Press, London and New York. HUGHES, SJ. (1951): Studies on micro-fungi XII. Triposporium, Tripospermum, Ceratosporella, and Tetraposporium (gen. nov.). Mycol Pap. 46: 1-35. HUGHES, S.J. (1976): Sooty molds. Mycologia 68: 693-820. JOHNSTON, A. (1960): A supplemental to a host list of plant diseases in Malaya. Mycol. Pap. 77. LIM, T.K. and KHOO, K.C. (1985): Diseases and Disorders of Mango in Malaysia, Tropical Press, Kuala Lumpur REYNOLDS, D.R. (1971): The sooty mold Ascomycete genus Limacinula. Mycologia 63: 1173-1209. REYNOLDS, D.R. (1978): Foliicolous Ascomycetes 2: Capnodium salicinum Montagne emend. Mycotaxon 7: 501-507. REYNOLDS, D.R. (1979): Foliicolous Ascomycetes 3: The stalked capnodiaceous species. Mycotaxon 8: 417-445. REYNOLDS, D.R. (1982): Foliicolous Ascomycetes 4: The capnodiaceous genus Trichomerium Spegazzini emend. Mycotaxon 14: 189-220. TUKEY, H.B. JR. (1971): Leaching of substances from plants. In Ecology of Leaf Surface Micro-organisms (eds. T.F. PREECE and C.H. DICKINSON) pp. 39-53, Acad. Press. London and New York.
Fig, 10: Cylindrical synemata of Leptoxyphium sp. Bar = 12.5 [Xm. Hughes (1976) listed L. graminum (Patouillard) Speg (= Capnodium graminum Pat) as the type species and the following genera as synonyms: Caldariomyces Woron., Megaloxiphium Cif., Bat. & Nasc. and Astragoxyphium Bat., Nasc. &Cif.
TURNER, G.J. (1971): Fungi and plant disease in Sarawak, CMIPhytop. Pap. 13: WILLIAMS, T.H. and LIU, P.S.W. (1976): A host list of plant disease in Sabah, Malaysia. CMI Phytop. Pap. 119. YAMAMOTO, W. (1954): Taxonomic studies on the Capnodiaceae. II. On the species of the Eucapnodiae. Ann. Phytopath. Soc. Japan 19: 1-5.
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(Received 20 August, 1988)
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Pertanika 11(3), 357-363 (1988)
Media for Embryo Culture of Some Tropical Recalcitrant Species H.F. CHIN1, B. KRISHNAPILLAY2 AND Z.C. ALANG3 1 &2
Agronomy and Horticulture Department Faculty of Agriculture Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia
3 Biotechnology Department Faculty of Food Science and Biotechnology Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia Keywords: embryo culture, recalcitrant species, media formulation. ABSTRAK Dalam kajian init embrio yang telah diasingkan dari bijibenih 11 spesis "recalcitrant" telah dikaji untuk menentukan media dan keadaan pengkulturan yang sesuai bagi perkembangan menjadi pokok yang lengkap. Media Murashige dan Skoog yang diubah melalui penambahan 0.17 gl"1 NaH PO telah didapati sesuai sebagai media dasar bagi semua spesis yang dikaji. Media dasar yang ditambah dengan 2gl~l 'activated charcoal* dan 1.0 mgl~~l NAA atau IAA bersama dengan 1 mgV~x Kinetin, BAP atau 2iP didapati sesuai bagi semua 11 spesis di mana embrio yang diasingkan itu dapat berkembang secara normal untuk menjadikan pokok yang lengkap. Bagi beberapa spesis, rendaman embrio yang terasing steril dalam larutan 'antioxidant* (10 mgl~l asid askorbik dan 5 mgl~l asid sitrik) selama 10-45 minit sebelum dikulturkan didapati menggalakkan perkembangan normal bagi embrio-embrio. ABSTRACT In this studyf the excised embryos of 11 recalcitrant species were studied to determine suitable media and cultural conditions for their development into whole plants. Murashige and Skoog's medium formulation modified slightly by the addition of 0.17 gl~l NaH2PO4 was found to be a suitable basal medium for all the species studied. The basal medium supplemented with 2 gl"1 activated charcoal and 1.0 mgl"1 NAA or IAA in combination with 1 mgl"1 Kinetin, BAP or 2iP was found to be suitable for all the 11 species where the embryos developed normally to form whole plants. For some of the species, soaking the sterile excised embryos in an antioxidant (10 mgl~l ascorbic acid and 5 mgl~^ citric acid) solution for 10-45 minutes prior to culture enhanced the normal development of embryos. INTRODUCTION The storage of seeds is the traditional means of conservation and international exchange of germplasm. For many crops, including most of the major world food resources, conservation can be carried out very efficiently in seed banks (Ellis et al.t 1985a, 1985b; Williams, 1984). The orthodox (i.e. desiccation tolerant) nature of these
seeds enables them to be dried to very low moisture levels, making them very amenable to long-term storage. However, in a number of tree species and plantation crops found especially in the tropics and subtropics, the seeds produced are recalcitrant (Roberts and King, 1980). Such seeds are sensitive to desiccation and temperature and thus cannot be preserved by conventional
H.F. CHIN, B. KRISHNAPILLAY AND Z.C. ALANG
seed storage methods for long periods due to degeneration of the embryos (Bajaj, 1985). Their maximum longevity varies from a few days to a few months. Orthodox seeds are relatively simple to store for long periods, in addition their excised embryos can be stored and cryopreserved. Culture of excised embroys of soybeans (Chin et ai, 1974, 1977), frenchbeans (1975), pea and wheat (Chin and Atiken, 1975) were easily cultured on sterilized soil. The general effect of cotyledon excision is slower growth and delay in senescence. However with the excised embryos of recalcitrant species which cannot tolerate desiccation, there was great difficulty raising seedlings from excised embryos in particular after a period of stress in storage at extremely low temperatures. These low vigour embryos need aseptic conditions and enriched media for their culture. In this paper, the objective is to find suitable media that support normal growth of excised embryos of recalcitrant species. This paper reports the findings of the first phase of the study carried out over a one year period, on the screening for suitable media for the culture of excised embryos of some of the recalcitrant species found commonly in this region. The carrying out of this part of the study prior to the actual work on finding storage techniques was necessary because a literature search revealed that very little work had been done to establish suitable media and cultural conditions for the normal development of excised embryos of the tropical recalcitrant species to be studied for long-term storage. MATERIALS AND METHODS
The excised embryos of 11 recalcitrant species were screened for suitable medium and cultural conditions. They were: jackfruit (Artocarpus heterophyllus), cempedak {Artocarpus champeden), rambutan {Nephelium lappaceum), langsat (Lansium domesticum), durian (Durio zibethinus), mango (Mangifera indica), kundang (Bouea ganadaria), avocado (Persea americana), cocoa (Theobroma cacao), rubber (Hevea brasiliensis) and dryobalanops {Dryobalanops aromatica). During the respective fruiting seasons, fresh ripe fruits were brought back to the laboratory and processed for their seeds. Embryos 358
from the seeds were excised and cultured within two days of the arrival of fruits in the laboratory. Prior to aseptic excision of the embryos, the seeds were either surface-sterilized with 15% Clorox solution or 80% alcohol, depending on the seed types. For mango and kundang, in which the flesh could not be completely removed from the testa, surface-sterilization was carried out using 15% Clorox containing 1 — 2 drops of 7-x detergent for 20 minutes followed by 3-4 rinses with sterile distilled water. For jackfruit, cempedak, rambutan, langsat, durian, avocado, cocoa, rubber and dryobalanops, individual seeds were swabbed with 80% alcohol and allowed to dry in a laminar flow cabinet for about 20 minutes before aseptically excising the embryos. Embryos from those species that exhibited browning during excision were immersed in a filter-sterilized antioxidant solution (10 mgl~ 1 ascorbic acid and 5 mgl" 1 citric acid) until they were cultured. This reduced browning of the cut ends, excessive secretion of polyphenols into the medium and increased the survival rate of the embryos in culture. Murashige and Skoog's (1962) medium formulation modified slightly by the addition of 0.17 gl" 1 of NaH2PO4 was used as the basal medium for the entire study. Various growth hormones at different concentration and combination were added to this medium. The auxins tested were NAA (a-napthalene acetic acid) and IAA (indole-3-acetic acid) at concentrations of 0, 0.1, 0.5 and 1.0 m g P 1 while the cytokinins examined were Kinetin, BAP (benzylaminopurine) and 2iP (2-isopentyladenine) each at concentrations of 0, 0.1, 0.5 and 1.0 m g P 1 . The effect of activated charcoal at concentrations of 0 and 2 gl" 1 , on normal development of the excised embryos was also studied. Difco Bacto agar at 6 gl" 1 was used to solidify the medium. For each treatment three replicates of ten excised embryos were employed. Each experiment was repeated twice over the fruiting season, once at the beginning of the season and the other towards the end. The number of normally-developing embryos (ie with a good shoot and root system) was scored as a percentage over th£ total number of cultures in each treatment. Evaluations were carried out at the end of six weeks of culture.
PERTANIKA VOL. 11 NO. 3, 1988
MEDIA FOR EMBRYO CULTURE OF SOME TROPICAL RECALCITRANT SPECIES TABLE 1 Mode of embryo development on a non-charcoal and charcoal medium supplement with various auxins and cytokinins of suitable levels
Modified Murashige & Skoog's Medium Without Charcoal + 0.1 mgl" 1 NAA orlAA and 0.1 mgl Kinetin, BAP or 2iP
Species
With Charcoal (2 gl l) + 1.0 mgl" 1 NAA or IAA —l and 1.0 mgl Kinetin BAP or 2iP
Artocarpus
heterophyllus
Normal Development
Normal Development
Artocarpus
champeden
Normal Development
Normal Development
lappaceum
Normal Development
Normal Development
Normal Development
Normal Development
Nephelium Lansium Durio
domesticum
zibethinus
Slow Growth
Enhanced Growth
Mangifera indica
Inhibited Growth
Normal Development
Bouea ganadaria
Inhibited Growth
Normal Development
Persea americana
Inhibited Growth
Normal Development
Theobroma cacao
Normal Development
Normal Development
Hevea brasiliensis
Slow Growth
Enhanced Growth
Dryobalanops
No development
Normal Development
aromatica
All cultures were maintained at 28°C ± 1°C in a temperature-controlled room with a 12-hour photoperiod of 8,000 to 10,000 lux intensity, provided by flourescent tubes. RESULTS The best results obtained for the various embryo types are summarised in Table 1. The optimum cultural conditions and problems encountered for individual species are reported below: (a) Artocarpus heterophyllus and Artocarpus champeden: The embryos in these two species could easily be excised and showed no browning during excision or in culture. Normal development was obtained both on a non-charcoal and charcoal added medium. On the non-charcoal medium, the best development was seen when 0.1 mg/~ ! NAA or IAA was used with 0.1 mg/" 1 Kinetin, BAP or 2iP. On charcoal medium, 1.0 mgl" 1 NAA or IAA when used with 1.0 mgl" 1 Kinetin, BAP or 2iP appeared optimum. Within six weeks, robust plants with good shoot and root system were obtained. (b) Nephelium lappaceum: The tiny embryo in this species was difficult to excise
because it is tightly held between the cotyledons at one end of the seed. It was impossible to excise it without causing serious damage. In this case, a small block 2 x 4 x 4 mm of the cotyledonary tissue enclosing the embryo was treated as the explant material. This tissue exhibited no browning and was easy to handle in culture. Growth of the embryo was observed in all the treatments both on a non-charcoal and charcoal added medium. However, the best development was observed with a combination of 0.1 mg/" 1 NAA or IAA and 0.1 mg/" 1 Kinetin, BAP or 2iP on a non charcoal medium and with a combination of 1.0 mgl" 1 NAA or IAA and 1.0 mgl" 1 Kinetin, BAP or 2iP on a charcoal added medium. (c) Lansium domesticum: The seeds of this species are polyembryonic. The seed could be fragmented into 2 or 3 pieces, each piece carrying an embryo embedded between the cotyledons. If the cotyledons were split open, the embryo embedded between them was torn into two. To avoid damage to the embryo, small blocks measuring 2 x 4 x 4 mm of the cotyledonary tissue enclosing the embryo was taken as the culture material. The excised tissues showed
PERTANIKA VOL. 11 NO. 3, 1988
359
H.F. CHIN, B. KRISHNAPILLAY AND Z.C. ALANG
no browning and grew satisfactorily to form plants. Best development was obtained in either a non-charcoal medium containing 0.1 mgl" 1 NAA or IAA in combination with 0.1 mgl" 1 Kinetin, BAP or 2iP or on a charcoal added medium containing 1.0 mgl" 1 NAA or IAA in combination with 1.0 m g T l Kinetin, BAP or 2iP. (d) Durio zibethinus: The embryos in this species were large measuring on the average 4 x 4 x 12 mm. The plumular end of the embryo is attached at the centre of the seed and when excised, exuded a mucous type secretion which was highly oxidative. Slight injury to the surface of the embryo caused strong browning reactions of the injured parts. Direct culture of these embryos on culture medium resulted in excessive secretion of phenols into the medium that slowed the growth and development of the embryo. However, when the embryos were soaked in antioxidant solution for 30-45 minutes, both the browning reaction and the secretion of phenols into the medium was greatly reduced and development was enhanced. Best development was observed in a charcoal medium containing i.O mgl" 1 NAA or IAA in combination with 1.0 mgl" 1 Kinetin, BAP or 2iP. The charcoal in the medium appeared to enhance normal and early development, compared to the non-charcoal medium in which growth was slow and plantlet development appeared weak and abnormal. (e) Mangifera indica and Bouea ganadaria: These two species have a fibrous mesocarp. The embryos excised from both species exhibited intense browning and phenols were secreted into the medium within hours of culture. Soaking the embryos in sterile antioxidant solution for about 30-45 minutes reduced the oxidative process of the polyphenols. Best growth and development was observed on a charcoal medium containing 1.0 mgl" 1 NAA or IAA in combination with 1.0 mgl" 1 Kinetin, BAP or 2iP. On a noncharcoal medium, inhibition of either the shoot or root system was commonly observed. (0 Persea americana: In this species, the embryo is located in the centre of the seed attached to both the cotyledons. Attempts to split the cotyledons to expose the embryo resulted in tearing the embryo into two. Hence, a small block of cotyledonary tissue measuring 4. x 4 x 4 mm and enclosing the embryo was excised and 360
treated as the culture explant. During excision, browning of the cotyledonary tissue was intense. Soaking the excised blocks in sterile antioxidant solution for 25—30 minutes prior to culture was found to be beneficial in reducing the oxidative effects of the polyphenols. For this species, charcoal medium containing 1.0 mgl" 1 NAA or IAA in combination with 1.0 mgl" 1 Kinetin, BAP or 2iP was found to be most promotive for normal development of the embryos. On a noncharcoal medium, inhibition of normal development was clearly evident. (g) Theobroma cacao: Excision of the embryonic axis in this species was easily effected, however browning of the embryo after excision occurred very quickly. Treatment of the embryos in an antioxidant solution for 10-15 minutes prior to culture appeared beneficial but not essential. On a non-charcoal medium, a hormone combination of 0.1 mgl" 1 NAA or IAA and 0.1 mgl" 1 Kinetin, BAP or 2iP was optimum for normal development. On charcoal medium, a hormone combination of 1.0 mgl" 1 NAA or IAA and 1,0 m g P 1 Kinetin, BAP or 2iP was best for normal shoot and root development. (h) Hevea brasilliensis: Embryonic axes of this species was easily excised and grown on medium. They exhibited no browning and were amenable in culture. Normal growth and development was much slower on a non-charcoal medium compared to a charcoal medium. On a charcoal medium containing 1.0 mgl" 1 NAA or IAA in combination with 1.0 mgl" 1 Kinetin, BAP or 2iP, a well developed shoot and root system was obtained after 6 weeks in culture. (i) Dryobalanops aromatica: This forest species set seeds only once in two or three years. The embryonic axes were easily excised from the seeds and emitted a sweet smell of camphor. No browning of the embryonic axes occurred during excision. However, after 2 - 3 days in culture, the embryo developed necrotic spots either at the plumular or radical region and failed to develop on any of the medium tested. When the excised embryos were dipped in an antioxidant solution for 10-15 minutes prior to culture, those embryos on the charcoal medium started to develop while none of the embryos on the noncharcoal medium developed normally. For best development of these embryos in a charcoal
PERTANIKA VOL. 11 NO. 3, 1988
MEDIA FOR EMBRYO CULTURE OF SOME TROPICAL RECALCITRANT SPECIES
medium, a hormone combination of 1.0 mgl - 1 NAA or IAA and 1.0 mgl" 1 Kinetin, BAP or 2iP was found to be suitable. DISCUSSION
A vital prerequisite to in vitro storage of embryonic axes for long term conservation is the elucidation of a reliable method for normal development of embryonic axes into seedlings under in vitro conditions when required, In the past, embryo culture techniques were developed principally for use in plant breeding, horticulture and for basic studies on embryo physiology and biochemistry (Collin and Grosser, 1984; Dunwell, 1986). Moreover, most of the studies on embryo culture have been confined to temperate crops and very little work has been carried out for tropical species. With the advent of the concept that excised embryos have the potential to be used as genetic material for difficult-to-store (recalcitrant) seeds, there now arises an urgent need to establish suitable in vitro practices for the regeneration of the embryos of recalcitrant species into whole plants. This study therefore aimed to fulfill this need and suitable media and cultural conditions for 11 recalcitrant species have been elucidated. From the study, it was found that Murashige & Skoog's (1962) basal medium modified slightly with the addition of sodium dihydrogen phosphate was found to be suitable for all the species screened. Monnier (1978) states that embryos that have their own endogenous hormones can be considered as plant initials. However, in several cases embryo culture has been facilitated by exogenously supplied hormones. In our study too we have found that in media without hormones, the embryos began to grow but development was either very slow or abnormal. However, growth and normal development was enhanced in medium containing low levels of growth hormones. There appeared to be no significant differences between the types of hormone combination used. Either NAA in combination with Kinetin, BAP or 2iP or IAA in combination with Kinetin, BAP or 2iP was found to promote normal development. However, the concentration of the hormones used was critical. On media without charcoal, 0.1 mgl" 1 of both auxin and cytokinin was found to be optimum while a charcoal media
containing 1.0 mgl 1 was found to be ideal for normal development. The use of activated charcoal at 2.0 gl" 1 was found to be promotive for the embryo development for all the species screened. Fridborg and Erickson (1975) suggested that the presence of activated-charcoal in the medium tends to remove those substances produced in the medium that promote unorganized growth, inhibit embryogenesis, root formation and elongation. In our study, we have observed a similar effect. Embryos developing on a charcoal-medium showed wellorganized and robust growth for all the species. In the case of Durio zibethinus and Hevea brasiliensis growth and development was better in a charcoal medium as compared to a non-charcoal medium. Moreover, for Dryobalanos aromatica, only the charcoal medium appeared to enhance normal development. This suggest that some of the species tend to release some toxic substances into the medium owing to injury during excision and these substances, if not removed, tend to be inhibitive to development of the embryos. Figures 1 (a)-(i) show some of the developing embryos in culture. Having established a system for normal development of plants from the embryos of these species, we have now moved on to working out a protocol for the long term storage of these species by cryopreservation of aseptic embryos in liquid nitrogen, (-196°C). Withers (1987) made a cautioned statement that after cryopreservation, tissues tend to undergo a considerable amount of damage and hence the composition of the initiallyestablished medium for non-cryopreserved embryos may not be sufficient as a recovery medium and may have to be further enriched with other growth promoting substances. It is therefore anticipated that the above established media for the different species will probably need to be modified in the course of the study to cater for the need of the recovering embryos after cryopreservation. ACKNOWLEDGEMENT
The authors thank Mr. Ong Choon Hoe for technical assistance and IBPGR Grant for this part of the study.
PERTANIKA VOL. 11 NO. 3, 1988
361
Fig. la: Jackfruit
Fig. 1b: Cempedak
Fig. Id: Langsat
Fig. le: Mango
Fig. If: Kundang
Fig. Ig:Avocado
Fig. Ih: Cocoa
Fig. li: Rubber
FIGURE 1: Embryo culture of some of the recalcitrant species 362
Fig. 1c: Rambutan
PERTANIKA VOL. II NO. 3, 1988
MEDIA FOR EMBRYO CULTURE OF SOME TROPICAL RECALCITRANT SPECIES REFERENCES BAJAJ, Y.P.S. (1985). Cryopreservation of embryos. In: Cryopreservation of plant cells and organs. Ed. by K.K. Kaitha, CRC Press Inc. Boca Raton, Florida, pp 227-242. CHIN, H.F., T.F. NEALES and J.H. WILSON. (1974). Effect of mechanical damage in soyabean seeds on seedling growth and final seed yield. Mai Agric. Res. 3: 184-189. (1975). The effects on seedling growth of simulated seed damage in french beans. Seed Set & Technol 3: 837-841. (1977). The effects of cotyledon excision on growth and leaf senescence in soyabean plants. Ann Bot. 41: 771-777. CHIN, H.F. and Y. AITKEN. (1975). Effect of seed damage on growth and development in pea {Pisum sativum L.) and wheat {Triticum aestivum L.). Ann Bot. 40, 165: 91-98. COLLINS, G.B. and W. GROSSER. (1984). Culture of embryos. In: Cell culture and somatic cell genetics of plants, Vol. I: Laboratory procedures and their applications. Ed. by I.K. Vasil, Academic Press, Inc. New York pp 241-257. DUNWELL, J.M. (1986). Pollen ovule and embryo culture as tools in Plant breeding. In: Plant tissue culture and its agricultural application. Ed. by L.A. Withers and P.G. Alderson, Butterworth, London pp 375-404. ELLIS, R.H., T.D. HONG and E.H. ROBERTS. (1985a). Handbook of Seed technology for genebanks. Vol. I. Principles and Methodology, IBPGR, Rome.
Compendium of specific germination information and test recommendations. IBPGR, Rome. FRIDBORG, G. and T. ERICKSON. (1975). Effect of activated charcoal on growth and morphogenesis in cell cultures.Physiol Plant. 34: 306-308. GROUT, B.W.W. (1986). Embryo culture and cryopreservation for the conservation of genetic resources of species with recalcitrant seed. In: Plant tissue culture and its agricultural application. Ed. by L.A. Withers and P.G. Alderson, Butterworth, London, pp. 303-309. MONNIERS, M. (1978). Culture of zygotic embryos. In: Frontiers of plant tissue culture 1978. E. by T.A. Thorpe. Univ. Calgary Press, Calgary, Canada pp 227-278. MURASHIGE, T. and F. SKOOG. (1962). A revised medium for rapid growth and bioassays with tobacco tissue culture.Physiol. Plant. 15: 473. ROBERTS, E.H. and M.W. KING. (1980). Storage of recalcitrant seeds. In: Crop Genetic Resources - the Conservation of Difficult Materials. Ed. by L.A. Withers and J.T. Williams, Int. Union Biol. Sci. Ser. B 42, IUBS, Paris pp 39-48. WILLIAM, J.T. (1984). A decade of crop genetic resources research. In: Crop Genetic Resources: Conservation and Evaluation. Ed. by J.H.W. Holden and J.T. Williams, Allen and Unwin, London pp 1-17. WITHERS, L.A. (1987). Long-term preservation of plant cells, tissues and organs. In: Oxford survey of plant molecular and cell biology Vol. 4 (In Press).
ELLIS, R.H., T.D. HONG and E.H. ROBERTS. (1985b). Handbook of Seed technology for genebanks. Vol. 2.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 4 November 1988)
363
Perlanika 11(3), 365-374 (1988)
Progress of Crop in Some Rhizophora Stands before First Thinning in Matang Mangrove Reserve of Peninsular Malaysia *P.B.L. SRIVASTAVA, **SAW LENG GUAN and ASHARI MUKTAR Faculty of Forestry Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia Key words: Mangrove; stocking density; diameter; height ABSTRAK Kajian ini melaporkan tentang komposisi dan pola pertumbuhan tanaman yang didominasikan oleh spesis Rhizophora berasaskan kepada stok pertumbuhan, diameter dan ketinggian pada dirian bakau yang terurus di Hutan Simpan Paya Laut Matang, Semenanjung Malaysia. Sejumlah 10 dirian telah dikaji terdiri dari umur 6 tahun (4), 9 tahun (3) dan 12 tahun (3). Secara purata terdapat 8371, 4661 dan 4181 batang sehektar tumbuhan Rhizophora untuk umur 6, 9 dan 12 tahun. Min diameter aras dada (DPD) dan ketinggian ialah 3.26 cm dan 6.34 m bagi umur 6 tahun, 5.50 cm dan 10.96 m bagi umur 9 tahun, dan 6.91 cm dan 12.62 m bagi peringkat umur 12 tahun. Kadar kematian yang tinggi berlaku pada umur di antara 6 hingga 9 tahun dan diperingkat umur ini juga terdapat pertumbuhan yang terbaik dari segi diameter dan ketinggian. Pepanjat Derris trifoliata mungkin memberi kesan kepada tumbuhan dan kemandirian dibeberapa kawasan. ABSTRACT The study reports the composition and growth pattern of the crops dominated by Rhizophora species in terms of stocking density, diameter and height growth in some wellYnanaged stands in Matang Mangrove Reserve of Peninsular Malaysia. In all, 10 stands representing 6 years (4), 9 years (3) and 12 years (3) were studied. On an average there were 8371, 4661 and 4181 stems/ha of Rhizophora in 6, 9 and 12 year-old crops respectively. Mean DBH and height for the respective age crops were 3.26 cm and 6.34 m, 5.50 cm and 10.96 m and 6.91 cm and 12.62 m. Highest mortality occurred in 6-9 year-old crop. This period also showed best growth both in diameter and height. These studies indicate the need for silvicultural thinning in 6-9 year-old crop. The climber Derris trifoliata may seriously affect growth and survival in some areas. INTRODUCTION In an earlier communication (Tay and Srivastava, 1982), some silvicultural aspects of the Mangrove Forest in Matang Reserve, such as, crop composition, stocking density before thinning I, II, III and final felling, DBH and height at these successive stages, were reported and their implications on the current management practices were discussed.
Present address :
The studies had indicated that there was a possibility of large scale mortality between year 2 and 15 when the crop is ready for first thinning. The present study reports observations on the progress of the crop in the early period of rotation i.e. after the final felling till the crop is ready for thinning I. As in the earlier studies it aimed at determining, (i) the species composition, (ii) stocking
*
Department of Forests, Papua New Guinea
**
Forest Research Institute, Kepong, Malaysia
P.B.L. SR1VASTAVA, SAW LENG GUAN AND ASHARI MUKTAR
density of different aged crops, and (iii) dynamics of size class distribution. On the basis of this information, it was hoped that comments can be made on, (i) rate of growth and survival/mortality, and (ii) realistic stocking and its effect on present thinning schedule, especially the first thinning. MATERIALS AND METHODS
Study Area The study was undertaken in Kuala Sepetang and Kuala Trong ranges of Matang Mangrove Reserve in Perak. These are amongst the best and the oldest managed mangrove forests in the world. Numerous publications (e.g. Watson 1928, Noakes 1952, Dixion 1959, Mohd. Darus 1969, Srivastava 1977, Srivastava and Abdullah Sani 1979, Srivastava and Daud 1978, Srivastava and Singh 1980, Tang et ah, 1980) have discussed various aspects of this Reserve. Haron (1981) has written the current working plan. Six stands representing the three age classes of 6-year, 9-year and 12-year old Rhizophora dominated stands were located in Kuala Trong Range. In Kuala Sepetang Range, four stands were choosen to represent the three different age classes
(Table 1). All sites included in the present study belong to Watson's inundation class I and II which are inundated 45 to 62 times in a month by normal tides. The prevailing conditions are suitable for the growth of Rhizophora spp., R. mucronata on soft mud along the streams and channels and R. apiculata showing optimum development in almost pure stands away from the streams. Sampling Procedures Systematics strip sampling, as followed by Tay and Srivastava (1982) was used in this study. Systematic sampling was choosen because of the simplicity of design/low cost, convenience on difficult sites and greater control over field work. Besides, in a systematic sample, the relative position in the population of different units included in the sample is fixed. In fact, the method gives a more precise estimate of the population than a random sampling (Sakhatame and Sukhatame, 1970). In selecting the representative coupes, a preliminary study of the whole Reserve was carried out with the help of maps and files provided by the District Forest Department, Larut Matang. Suitable compartments were made on the basis of the following criteria :
TABLE 1 Locality Features Range
1
Site
Age
Compt.
No.
(Yrs)
No.
2
3
4 69
KT KT
Total Area (ha)
Area Surveyed (ha)
5
6
7
27.1 27.1
20.0 20.0
1.00 1.00
16.2 20.2 44.5
10.8
0.54
10.8
0.54
72 KS 7 KS 21
Actual Area Sampled (ha)
K.T K.T K.S
5 6 7
9
55 72 23
40.5 41.3 31.0
16.0 20.0 10.0
0.80 1.00 0.54
K.T K.T K.S
8 9 10
12
60 51 17
38.0 37.2 16.2
20.0 20.0 10.8
1.00 1.00 0.54
Source : Larut Matang District Forest Office Taiping, Perak Note:
366
KS KT
-
Kuala Sepetang Kuala Terong
PERTANIKA VOL. 11 NO. 3, 1988
PROGRESS OF CROP IN SOME RHIZOPHORA STANDS BEFORE FIRST THINNING
infested by this climber. Their presence was approximated in the following grades.
(i)
coupe selected must be a Rhizophora dominated stand, (ii) the coupe must be of a pre-determined age, (iii) the coupe must be within Watson's (1928) inundation classes I and II, (iv) only coupes designated for charcoal were selected; those earmarked for firewood have shorter rotation. The sampling plots were selected based upon the following methods. A base line was established along a convenient boundary. Sampling lines, spaced 200 m apart were drawn straight and at right angles to the base line. Sample plots of 20m x 10m were marked contiguously along the sampling line, thus making a continuous strip of 10m width. This yielded a sampling intensity of 5 per cent. All the trees encountered in the plots were counted by species and their diameter at breast height (DBH) recorded. In each plot, height of one tree was recorded with the help of clinometer. Trees occurring on forward and right side boundaries of the plot were included in the enumeration while those on the left and rear end boundaries were excluded. Other details of the plot, like presence of fern weed (Acrostichum aureum) and climbers which could affect the survival and growth of the trees were also recorded. While the presence of fern was estimated ocularly, the occurrence of Derris trifoliata was recorded in terms of the trees
Grades
Value
0 1
Nil Rare
2
Moderate Abundant Infested
3 4
% infestation (trees/plot) 0
1-25 26-50 51-75 75-100
Statistical Method A single classification ANOVA, in the form of a nested ANOVA design was used in this study. When there was significant variance between the means, Student - Neuman - Keul's test was used to determine location of the significant difference. Nintyfive percent confidence level was used in both the nested analyses of variance and the significance test. RESULTS
Species Composition Table 2 presents the composition of tree species recorded on different sites. With the exception of Site 7, all sites had Rhizophora apiculata as the most dominant species (89.5 to 99.4%). Site 7, however, had only 46.0 of the total stems of this species. R. mucronata occurred in generally quite low densities (i.e. less than 4%). Site 7, however, had the highest percentage of this species (50.5%).
TABLE 2 Species composition expressed as percentage of total number of stems
Class
Sites
Rhizophora apiculata
Rhizophora mucronata
6
1 2 3 4
92.4 97.7 94.2 99.4
3.8 0.1 3.5 0.4
5 6 7
91.0 96.9 46.9
1.0 0.8 50.5
8 9 10
93.0 95.9 89.5
3.9 2.8 3.0
Age
Years
9 Years
12 Years
Bruguiera parviflora
PERTANIKA VOL, 11 NO. 3, 1988
Others
Total
3.8 2.2 2.2 0
0 0 0.1 0.2
100 100 100 100
7.9 2.3 0.4
0.1 0 2.2
100 100 100
2.8
0.3 0 3.0
100 100 100
1.2 4.5
367
P.B.L. SRIVASTAVA, SAW LENG GUAN AND ASHARI MUKTAR
On all the sites, Bruguiera parviflora comprised of less than 8% of the total stems; The other species recorded rarely were trees of Bruguiera gymnorrhiza, B. cylindrica, Xylocarpus sp:, Avicennia sp. and Sonneratia sp. Stocking, DBH and Height The summary of the results of this study is shown in Table 3. In general, the plots in the age class 6 years showed highly variable results. Statistical tests on this results revealed the following. a. Stocking (Table 4) There was no significant difference in stocking between the age classes of 9 and 12 years. However, when tested against the 6 years class there were significant differences. b. DBH (Table 5) The DBH showed significant differences in all sites and ages. c. Height (Table 6) The were significant differences in height between all the age classes. In general there was also a certain amount of uniformity in heights within the age classes as reflected in no significant differences between some of the plots of the same age classes. Weeds Infestation - Acrostichum aureum (fern) and Derris trifoliata Lour, (climber) Sites 6 and 7 were free from fern infestation; Sites 5 and 9 were found to have heavier fern infesta-
tion than other sites (Table 7). On the whole fern infestation was low. Six out of ten sites showed no infestation by climber. Of the remaining four sites, Site 9 showed heaviest infestation. Site 5 also showed heavy infestation. However, sites 2 and 6 had relatively light infestation by the climber. DISCUSSION
Species Composition On all the sites, Rhizophora spp. formed more than 90% of total standing trees. The most common associate of Rhizophora spp. was Bruguiera parviflora. This species is not preferred mainly because it is inferior as a fuel wood. In the present study, however, it never exceeded 8 per cent in any stand. It has been reported by many workers that there is an increase in the frequency of B. parviflora after clear felling (Dixon 1959, Mohd. Darus 1969). It appears that the present management system has ample control over the distribution of other inferior species. Among Rhizophora spp., R. apiculata was dominant on all the sites except site 7 where R. mucronata was more abundant. Stocking Srivastava and Abdullah Sani (1979) projected that after Thinning I, there should be 6726 stems per ha. If it is assumed that this is the minimum
TABLE 3 Summary of results
3A. Age classes Age
Stocking (Stems/ha)
6 Years
9 Years
12 Years
8371
4661
4181
DBH
3.26
5.50
6.91
Height (m)
6.34
10.96
12.62
3B. Sites 6 years
Age
12 years
9 years
1
2
3
4
5
6
7
8
9
10
Stocking (Stems/ha)
8734
5739
11989
8954
4347
5437
3648
4258
4170
4059
DBH (cm)
3.35
3.69
2.81
3.19
5.52
5.39
5.80
7.23
6.63
6.80
Height (m)
6.28
5.81
6.16
7.10
10.30
10.43
12.15
12.73
12.37
12.77
Sites
368
PERTANIKA VOL. 11 NO. 3, 1988
PROGRESS OF CROP IN SOME RHIZOPHORA STANDS BEFORE FIRST THINNING TABLE 4 Test of significance - Stocking 4A. Age classes (stems per plot)
2
1
Rank Age (years) Mean stocking (x) Replicates (n)
3
12
9
6
83.62
93.21
167,42
127
117
154
Significant Test
4B. Sites (stems per plot)
7
9
10
1
2
3
4
5
Site No.
7
10
9
8
5
6
2
1
4
3
50
50
27
27
114.78
174.68
179.07
Replicates Mean
6
8
Rank
27
27
50
50
40
50
72.96
81.19
83.40
85.16
87.48
108.74
239.78
Stocking (x) Test Note 1 : Means that have no significant difference with each other are indicated with a common line.
TABLE 5 Test of significance -Diameter at Breast Height (DBH) 5 A. Age classes (cm) Rank
1
2
3
Age (years)
6
9
12
Replicates (n)
25782
10906
10617
MeanDBH (x)
3.26
5.50
6.91
Significant Test
5B. Sites (cm) Rank
1
Site No.
3
4
Replicates (n)
6474
MeanDBH (x)
2.81
2
3
4
5
6
7
8
9
10
1
2
6
5
7
9
10
8
4835
8734
5739
5437
3499
1970
4170
2189
4258
3.19
3.35
3.69
5.39
5.52
5.80
6.63
6.80
7.23
Significant Test
Note 1 :
Means that have no significant difference with each other are indicated with a common line.
PERTANIKA VOL. 11 NO. 3, 1988
369
P.B.L. SRIVASTAVA, SAW LENG GUAN AND ASHARI MUKTAR TABLE 6 Test of significance - Height 6A. Age classes (m) Rank
1
2
3
Age (years)
6
9
12
Replicates (n)
154
117
127
Mean Height (x)
6.34
10.96
12.62
Significant Test 6B. Sites (m) Rank
1
2
3
4
5
6
7
8
9
10
Site No.
2
3
1
4
5
6
7
9
8
10
50
27
50
27
40
50
27
50
50
27
5.81
6.16
6.28
7.10
10.30
10.43
12.15
12.37
12.73
12.77
Replicates Mean Height (x) Significant Test
Footnote 1 : Means that have no significant difference with each other are indicated with a common line. TABLE 7 Weed infestation (expressed in percentage of plots affected) 7A, Acrosticlium aureum Site No. 1
2 3 4
5 6 7 8 9 10
Infestation - Grades 0
1
2
3
4
42 96 78 96 53 100 100 78 12 78
36 4 19 4 7 16 44 8
18 -
3 15 6
2 5 -
2 _ — 20 _ _ —
28 7
14 7
2 -
100 100 100 100
100 100 100
100 100 100
7B. Derris trifoloata Lour. Site No. 1 2 3 4 5 6 7 8 9 10 370
0
1
100 88 100 100 40 84 100 100 6 100
— 6 3 8 _ 22 -
Infestation - Grades 2 _ — 12 2 _ _ 10 -
3
4
_ 4 _ 10 4 _
«. 2 35 2 — _ 46
16 -
PERTANIKA VOL. 11 NO. 3, 1988
100 100 100 100 100 100 100 100 100 100
PROGRESS OF CROP IN SOME RHIZOPHORA STANDS BEFORE FIRST THINNING
density for adequate stocking before Thinning I, then all the sites of six-year-old crop with an average of 8371 stems per ha, with the exception of Site 2 were adequately stocked. The stocking of 6-year crop, in the present study, appears to agree closely with studies conducted by the earlier workers on the younger age crops (Liew etal, 1977, Srivastava & Daud Khamis 1978, Srivastava & Abdullah Sani 1979). They obtained densities ranging from 8,300 to 9,100 stems per ha within 24 months after final felling. From this comparison, it appears that there was little net recruitment or mortality during 24 months to 6 years after felling. A possible reason for low recruitment could be the dense crop near the streams preventing water-borne seedlings from reaching the interior blank areas. Net mortality is also presumably low because the crop has yet to reach a size where competition becomes a major factor. At the same time, 24 month-old plants are strong enough to withstand environmental stresses. However, the average density by itself can be misleading. During the course of the survey, it was observed that there were large patches of blank areas. The sites at 6 years age could definitely support a density higher than the average number of 8371 stems per ha. This was evident on site 3, where the stocking was as high as 11,989 stems per ha. To get an index of the distribution pattern of the trees, the following exercise can be done.
Using 6726 stems as the minimum stocking per ha, the minimum stocking per plot can be calculated in the following manner: Area of each plot Minimum density per ha Therefore, the minimum stems per plot
: 10x20m2 = 200m2 : 6726 stems per ha = 6727 x 200 10 000 = 134.54 stems per plot.
Using this figure, the distribution pattern of stem density per plot was calculated (Table 8). The table shows the cumulative percentage distribution of stems per plot expressed at intervals of approximately one quarter, half, three quarters, one, one and one quarter and one and a half. All the plots stocking less than 134 stems are considered inadequately stocked. As is evident from this table, between 37% and 64% of the plots in the 6-year old crop were inadequately stocked though the average density was more than the minimum stocking. Apparently a large number of plots were poorly stocked. Thus if a area has high average stocking but has irregular and patchy distribution with large blanks, it would still be considered poorly regenerated and may require planting. However, another equally important consideration is whether 6726 stems per ha is too high a value for adequate stocking. This will be discussed later in this section.
TABLE 8 Cumulative stocking of stem density per plot (percent)
Age
Site
(years)
No.
less than
less than
less than
less than
less than
less than
32
66
100
134
168
202
4
2 10 0 11
10 32 7 26
32 54 22 30
46 64 48 37
56 76 55 52
60 84 59 59
5 6 7
15 0 7
38 12 37
60 40 85
83 76 100
95 94 100
100 100 100
8 9
0 0 11
28 26 33
72 72 66
90 92 100
100 98 100
100 100 100
1 6
9
12
Cumulative stocking (stems/plot)
2 3
10
PERTANIKA VOL. 11 NO. 3, 1988
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P.B.L. SRIVASTAVA, SAW LENG GUAN AND ASHARI MUKTAR
The density in the 9-year old stands was, on an average, 4661 stems per ha which compared well with Noakes' (1952) Tentative Yield Table. Noakes estimated a density of 3954 stems per ha at 8 years of age and 3188 stems per ha at 9 years. During this present study with an average stocking of 4661 stems per ha, the 9-year old crop appears to be inadequately stocked compared to the standard of 6726 stems per hectare. Based on this criteria, Table 8 indicates that between 76 to 100% of the plots fall below the adequately stocking standard. Although the results showed no significant difference at 95% confidence level between the sites, site 7 of the 9-year old crop had the lowest density. This was probably due to the higher composition of R. mucronata in the stand. Site 7 had 50.50% of/?, mucronata, whereas, the other sites had more than 89% of R. apiculata. R. mucronata is planted at 1.83m x 1.83m espacement to yield a density of 2990 stems per ha. Therefore, with such a planting distance, higher composition of this species results in a lower density. Moreover, R. mucronata is a larger tree than R. apiculata and requires more growing space. Comparing the density of 9-year old crop with that of 6-year old crop, there is an observed drop of about 44%. This is a high rate of loss considering that the net mortality between 24 months and 6 years was negligible. It appears that during this period i.e. between 6 and 9 years, the factors affecting survival are different from those that prevailed during the earlier period. Competition probably is the most important factor as the space between stems is much reduced and the ability of the site to support a large number of stems per unit area becomes limiting. Watson (1932) had observed that the Rhizophora spp. are extremely susceptible to competition. The density of the crop at 12 years of age was 4181 stems per ha. There was no significant difference with the 9-year old crops at 95% confidence level. It can, therefore be infered that mortality between the ages 9 and 12 years is negligible or has slowed down to a level that is statistically not significant. It appears that after the high rate of mortality between the 6 and 9 years, the stands begin to stablise. At 12 years age, Noakes predicted the density of crop at 2743 stems pei ha. This value is lower than the average density observed presently. 372
The present study compares well with Tay and Srivastava's (1982) figures of 15-year old crop in Kuala Trong (3922 stertis per ha). It also indicates that the mortality between 12 and 15 years (Tay's observation compared with the present findings) is low compared to the period between 9 and 12 years. On the basis of the projected figure of 6726 stems per ha, the 12-year crop appeared to be inadequately stocked. Table 8 indicates that more than 90% of the plots have stocking rate of below 134 stems per plot. Based on the present finding, the apporximation made by Srivastava and Abdullah Sani (1979) appears to be too high. This study, as indicated above, agrees well with Tay and Srivastava's findings. The stocking of the crop at Thinning I should be within a range of 2500 to 4000 stems per ha. Srivastava and Abdullah Sani (1979) did not take into consideration the high rate of mortality between 6 and 9-year old crop and based their projections on 1.22 x 1.22 m initial planting space. Tay and Srivastava (1982) estimated that 3363 stems per ha would be a realistic figure after Thinning I. If we consider this to be the minimum stocking, then the new stocking per plot for the purpose of the present study would be 67 stems per plot (336x200). 10 000 On re-examination of Table 8, it may be seen that only 7 to 38% of the plot were inadequately stocked. On the basis of the above discussion, the following points can be summarised; (1) low mortality between 24 months and 6 years, (2) highest mortality in 6 and 9-year old crop, (3) crops stabalise after 9 years, mortality being low between 9 and 15 years, (4) stands with higher percentage of R. mucronata have lower density; and, (5) a better approximation of density of crop at Thinning I would be about 3363 stems per ha. The implications of the above observations would be that Thinning I would be of greatest value to the stand when competition between stems is most intense, i.e. between 6 and 9 year period. It is the author's contention that the 1.22m (4ft) stick thinning would benefit the crop
PERTANIKA VOL 11 NO. 3, 1988
PROGRESS OF CROP IN SOME RHIZOPHORA STANDS BEFORE FIRST THINNING
most during this period. At 6 years, the density of the crop is generally greater than 6726 stems per ha. Hence, stem release would be more appropriate at this stage. However, due to the non-commercial aspect of such a thinning, this practice is not being currently followed in Matang. The above discussion indicates that systematic studies must be carried out on the effect of thinning when the crop is 6 and 9 years old to determine its effect on the final crop. The above discussion indicates that systematic studies must be carried out on the effect of thinning when the crop is 6 and 9 years old to determine its effect on the final crop. The benefits of such a thinning on the subsequent crop might make it worthwhile in spite of little revenue at this age. If the end product are chips when even the small diameter stems can be chipped, as in Sabah and Sarawak, this should become a worthwhile consideration. Diameter Growth The results show that there is*significant difference at 95% confidence level between the three age classes and between ^11 the sites. This is due to the large sample size used in the computation of the average diameter. The sample size of the sites was generally above 2000 replicates. This probably resulted in the individualistic nature of each site. Nevertheless, an examination of the values in Table 4 will reveal that the average diameters of the same age class identifies better with each other than with different age classes. The present study showed a much lower average diameter than predicted by Noakes (1952); at ages 8 and 10 years, 6.5 and 8.1 cm respectively or an average of 7.2 cm for a 9-year old stand and 9.5 cm for a 12 year old crop as against 5.5 cm and 6.9 cm in the present study. However, these results compare well with Tay and Srivastava (1982).They recorded the diameter of Rhizophora dominated stands to range from 6.6 cm to 9.9 cm with an average of 9.25 cm. The annual growth between 9 and 12 years and between 12 and 15 years is more or less the same, 0.47 cm per year and 0.45 cm per year respectively. In this study, the growth pattern of diameter between 6 and 12 years is shown in Figure 1. The curve is sigmoidal. Highest growth increment occurred during the period between 6 and 9 years. In absolute values, the growth between 6 and 9
years was determined to be 0.75 cm per year as compared to 0.47 cm per year between 9 and 12 years. Both in terms of diameter and height, growth appear to be the highest during the period of 6 and 9 years. This factor could probably explain the reason for highest mortality during this period. When crop is most vigorous, competition between stems will also be intense resulting in high mortality of the suppressed plants. Height Growth Height, being an independent character, generally is a good indicator of the age and site quality of the crop. The results of this study showed a good uniformity of height among the sites of the same age crops but with two exceptions, i.e. Sites 4 and 7. Except for sites 4 and 7 all the sites, showed no significant difference at 95% confidence level with each other within the same age crop. The reason site 7 had a higher value than the other sites in the 9-year old crops, could be due to the higher proportion of/?, mucronota. As for site 4, the higher value could be due to two possible reasons, viz: being a better site or it could be a slightly older crop. If planting had occured one year earlier than the other sites, that would make a significant difference in height growth. However, records on planting were not available. Figure 2 shows the trend of height growth of the three age classes. \X shows a typical sigmoidal curve. It is apparent that highest rate of growth occurs between 6 and 9 years. The height difference between 6 and 9 years was 4.63 m, as against 1.66 m between 9 and 12 years. In other words, the rate of growth was 1.54 m and 0.55 m per year during the respective periods. The growth during the period of 6 and 9 years is about 2.8 times faster than between 9 and 12 years. If it is related to the density and DBH, more or less some trends are obtained i.e. higher mortality and greater DBH in the 6-9 years old crop compared to 9-12 year crop. Weeds Two species of weeds that might affect the growth of Rhizophora spp. found in the present study were Acrostichum and Derris trifoliata. The former is a fern and latter a climber. The climber was seen to have girdled a large number of trees in the affected areas. However, further investigation should be conducted to determine exactly its role
PERTANIKA VOL. 11 NO. 3, 1988
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P.B.L. SRIVASTAVA, SAW LENG GUAN AND ASHARI MUKTAR
in suppressing tree growth. The present study revealed only 4 out of 10 sites affected by this climber. A. aureum (piai) is generally observed in the drier areas and frequently seen to be associated with peaty soils. On the whole in the present study, only three sites were found to be infested to any significant level (sites 1, 5 and 9). On site 5, piai was found in the large lasa form. This was recorded in the last sampling line. Stocking was found to be very low here. On the whole, however piai did not appear to be a major problem on most of the sites surveyed in the present study. CONCLUSIONS AND SUGGESTIONS
On the basis of this study, the following conclusions can be drawn :— (i) Based on the present findings, the optimum stocking of Rhizophora spp. at Thinning I is more realistic at 3363 stems per hectare, (ii) The studies further indicate that the period between 6 and 9 years shows intense activity in terms of growth and mortality, (iii) It is suggested that the Forest Department looks into the possibility of a silvicultural thinning between 6 and 9 years old crop which might improve the rate of growth and reduce mortality.
ACKNOWLEDGEMENT The authors take this opportunity to thank the Dean, Faculty of Forestry, Universiti Pertanian Malaysia and Forestry staff of Lamt - Matang Forest District for facilities. One of us (SLG) is specially thankful to Shu Chow for her support and patience during the study period. REFERENCES
HARON HJ. ABU HASSAN (1981): A working plan for Matang Mangrove Forest Reserve, Perak. Forest Dept. Pub. LIEW, T.C., MOHR NOR. DIAH and W.Y. CHUN. (1975): Mangrove Exploitation and Regeneration in Sabah.Mz/. For. 38: 260-270. MOHD. DARUS HAJI MAHMUD. (1969): Rancangan Kerja Bagi Hutan Simpan Paya Laut, Matang, Perak. Forest Dept. Pub. NOAKES, D.S.P. (1952): A working plan for the Matang Mangrove Forest Reserve, Perak. Forest Dept. Pub. SRIVASTAVA, P.B.L. and DAUD KHAMIS. (1978): Progress of natural regeneration after the final felling under the curren t silvicultural practise in the Matang Mangrove Reserve. Pertanika, 2: 126-135. SRIVASTAVA, P.B.L. and ABDULLAH SANI SHAFIE. (1979): Effects logging on natural regeneration of mangrove species in Matang Reserve. Pertanika, 2(1): 34-42. SRIVASTAVA, P.B.L. and H. SINGH. (1980): Composition and distribution pattern of natural regeneration after second thinning in Matang Mangrove Reserve, Perak. Proc. Asian Symp. on Mangrove Environment: Res. & Management. Kuala Lumpur. SAKHATAME and SUKHATAME. (1970): Sampling Theory of Surveys with Applications. Asia Publishing House, Bombay. TANG H.T., HARON HJ. ABU HASSAN and E.K. CHEAH (1980): Mangrove forest in Peninsular Malaysia: A review of management and research objectives and priorities. Asian Symposium on Mangrove Environment: Research and Management. Kuala Lumpur 25-29th August 1980. TAY, S.P. and P.B.L. SRIVASTAVA (1982): Crop composition and density after thinnings and before final felling in Matang Mangrove Reserve, Perak. Pertanika 5(1): 95-104. WATSON, J.G. (1982). Mangrove Forests of the Malay Peninsular, Malay. For. Rec. No. 6. Fraser and Neave, Singapore.
DIXON, R.G. (1959): A working pbn for the Matang Mangrove Forest Reserve, Perak. Forest Dept. Pub.
374
PERTANIKA VOL. 11 NO. 3, 1988
(Received 4 September 1987)
Pertanika 11(3), 375-384(1988)
Properties of Soils Derived from Some Metamorphic Rocks in Peninsular Malaysia S. ZAUYAH Department of Soil Science, Faculty of Agriculture, Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia. Key words: Soil properties; parent materials; pedofeatures ABSTRAK
•
Sifat-sifat morfologi, fizik, kimia, mineralogi dan mikromorfologi ke atas limajenis tanahyang terbentuk daripada batuan metamorf (skis kuarza-mika, filit, skis serisit bergrafit, skis amfibol dan serpentinitj telah dikaji. Sifat-sifat ini didapati terpengaruh oleh jenis bahan induk. Tanah-tanih yang ter~ terbentuk daripada batuan yang rendah kandungan mineral senang luluhawa dan jumlah ferum rendah (skis kuarza-mika, filit, skis serisit) adalah berwarna kuning kemerahan, bertekstur lempung berkelodak, berstruktur blok serta mempunyai jumlah keupayaan pertukaran kation, ketepuan bes dan ferum oksida bebas yang rendah. Horizon diagnostik pada tanah ini ialah horizon argilik. Tanah-tanih yang terbentuk daripada batuan yang tinggi kandungan mineral senang luluhawa (skis amfibol dan serpentinit) adalah berwarna merah hingga coklatf bertekstur lempung, berstruktur gerintil, serta mempunyai jumlah keupayaan pertukaran kation rendah, ketepuan bes sederhana, kandungan ferum oksida bebas tinggi. Horizon diagnostik ialah oksik. Ciri-ciri pedo yang banyak terdapat dalam kumpulan tanah yangpertama ialah pengisian lempung dan kotoran manakala dalam kumpulan tanah yang kedua pengisisan kotoranlah yang banyak sekali.
ABSTRACT The morphological physical, chemical, mineralogical and micromorphological properties of five soils developed over some metamorphic rocks (quartz-mica schist, phyllite, graphitic sericite schist, amphibole schist and serpentinite) were examined. These properties were found to be influenced by the parent materials. Soils developed over rocks with low amounts of weatherable minerals (ferromagnesian) and low total iron content (quartz-mica schist, sericite schist and phyllite) are reddish yellow, have silty clay textures and blocky structures, low CEC and base saturation and low free iron oxide content. The diagnostic horizon is argillic. Soils formed over rocks with high amounts of weatherable minerals (amphibole schist and serpentinite) are red to brown, have clayey textures, granular structures, low CEC, moderate base saturation and high free iron oxide content The diagnostic horizon is oxic. Pedofeatures in the first group of soils are dominated by clay and excremental infillings whilst in the second group, only excremental infillings are dominant. INTRODUCTION Metamorphic rocks such as phyllite, sericite schist and quartz-mica schist are some of the common parent materials of soils mapped in Peninsular Malaysia. Although there have been many studies related to the genesis of soils in Malaysia (Eswaran
and Wong, 1977; Lim, 1977; Paramananthan, 1977; Loh, 1981; Zainol, 1984) the weathering of metamorphic rocks in relation to soil formation has received less attention than that of igneous and sedimentary rocks. Properties of soils developed on igneous rocks and sedimentary
S. ZAUYAH
6 0 km
; v.
\
P E N INSULAR M ALAYSIA
1
Sampling site of profile I.
Quartz-mica schist
2.
Graphitic sericlte schist
3.
Phyllite
4.
Amphibole schist
5.
Ssrpentinite
Fig. 1: Map of Peninsular Malaysia showing the locations of the profiles studied.
376
PERTANIKA VOL. 11 NO. 3, 1988
PROPERTIES OF SOILS DERIVED FROM SOME METAMORPHIC ROCKS
rocks have been shown to be dependent on the lithology, and iron content of the rocks from which the soils are derived (Paramananthan, 1977, Zauyah, 1984). Studies on properties of soils derived from these parent rocks have provided more data for soil classification and soil survey interpretations. In this study, soils developed on five metamorphic rocks of different metamorphic grades and petrographic properties were examined. They are quartz-mica schist, phyllite, graphitic sericite schist, serpentinite and amphibole schist. The objective of this study is to examine the morphological, physical, chemical, mineralogical and micromorphological properties of these soils and relate these properties to the inherent properties of the parent materials. MATERIALS AND METHODS Materials
The five soil profiles sampled for the study are in different parts of the Peninsular Malaysia (Fig 1). The quartz-mica schist is situated in the north of Kuala Lumpur city and belongs to the Dinding Schist Formation of Lower Paleozoic. The graphitic sericite schist belongs to the Kajang Formation (predominantly of quartz sericite and graphitic sericite schists) which is of middle Upper Silurian age. It is located in the Universiti Pertanian Campus in Serdang. The phyllite outcrop is exposed 6 km from Karak town on the Kuala Lumpur-Karak Highway. It belongs to the Karak Formation which is of Lower Devonian age and consists predominantly of carbonaceous argillaceous rocks. Isolated bodies of amphibolites and serpentinites are found in several parts of Peninsular Malaysia. The serpentinite profile sampled is exposed on a roadcut of an ellipsoidal shaped intrusive body (about i.5 km long) with preCarboniferous quartz-mica schist on the Seremban -Kuala Pilah road. The amphibole schist profile sampled is a roadcut exposed along the Kuala Lumpur-Karak Highway near Bentong. It is a lenticular-shaped body (1 km long) occurring within a sequence of low—grade regionally metamorphosed schist of the Bentong Group.
Methods The soil profiles were examined and the morphological zones were first identified i.e. rock, saprock, saprolite, pedoplasmation zone and soil horizons. Field description of the soil horizons were done following the guidelines in the Soil Survey Manual (USDA, 1981) and FAO 'Guidelines for Soil Description' (1977). The horizons were then sampled by collecting about 2 kg for bulk analyses, clods for bulk density determinations and undisturbed samples in Kubiena boxes for thin sections. For the amphibole schist and the serpentinite profiles, samples were also collected close to and along the joint planes. The soil and weathered material were fractioned into clay (< 2 /xm), silt (2-50 jjm) and sand (50-200 jum) by successive sedimentation. The clays were deferrated following the method of Mehra and Jackson (1960). All the fractions were analysed for their mineralogical composition by X-ray diffraction. The routine chemical analyses carried out on the soil and some of the weathered samples are as follows: pH (1:2.5) in water, CEC and exchangeable bases (Chapman, 1965), extractable aluminium (Yuan, 1959) and extractable free iron (Mehra and Jackson, 1960), Thin sections (9 cm x 12 cm and 9 cm x 6 crn) were prepared by impregnating the samples with unsaturated polyester resin. The micromorphological features related to microstructure, coarse and fine fractions and pedofeatures were described according to the terminology of Bullock et al (1985). Fresh fracture surfaces of average size 4 mm were also examined with a Jeol 35C scanning electron microscope. Bulk density was determined by the method of Brasher (1966). The Atterberg Limits (plastic limit and liquid limit) were determined according to the BSI (1975) method of analyses. COLE values were calculated from the bulk densities of the moist soil and oven dried soil as outlined by Grossman et al (1968). RESULTS AND DISCUSSION
Morphological Properties The morphological properties of the soils studied are summarised in Table 1. From these properties
PERTANIKA VOL. 11 NO. 3, 1988
377,
S. ZAUYAH TABLE 1 Morphological properties of the B horizons of soils developed on some metamorphic rocks in Peninsular Malaysia. * ^ ^
Parent Phyllite
Graphitic Sericite Schist
Quartz-Mica Schist
Amphibole Schist
Serpentinite
Properties ^ ^ ^ ^ Subsoil Texture Diagnostic Horizon
Silty clay
Silty clay
Silty clay
Clay
Clay
Argillic
Argillic
Argillic
Oxic
Oxic
Reddish yellow 7.5 YR 6/8
Reddish yellow 7.5YR 6/6
Reddish yellow 7.5YR 6/8
Yellowish red 5YR 5/6
Dark reddish brown 5YR 3/3
Subangular blocky
Subangular blocky
Granular
Granular
Friable
Friable
Colour Pale brown 10YR 7/4 Structure
Consistency
Angular blocky Firm
Firm
Drainage
Moderately well drained
Well drained
Depth of Solum
Moderately deep
Moderately deep
two distinct groups of soils can be identified:those that developed over the first three parent rocks i.e. quartz-mica schist, phyllite and graphitic sericite and those over the amphibole schist and the serpentinite. The former group corresponds to rocks i.e. quartz-mica schist, phyllite and graphitic sericite schist and those over the amphibole schist and the serpentinite. The former gropp corresponds to rocks with low weatherable minerals (mainly quartz and mica) whilst the latter group has high weatherable minerals (serpentine, actinolite, epidote). In the field too, these two groups show differences in their profile morphology. The soils developed over rocks low in weatherable minerals or silica rich, have horizons which are easily distinguished in the field. They show characteristics of argillic horizons. On the other hand, soils developed over rocks high in weatherable minerals have almost indistinct horizonation and are oxic in characteristics. Other distinctintive properties of the first group of soils are : reddish yellow colours, silty clay textures, firm consistency, subangular to blocky structures and are well drained. 378
Firm Well drained
Deep
Well drained
Deep
Excessively drained Deep
The second group of soils (over serperitinite and amphibole schist) are deep, well drained to excessively drained profiles, are red to brown, have clayey textures and friable consistency. Paramananthan (1977) also noted that soils developed over parent materials high in weatherable minerals show an increase in sesquioxides which resulted in the formation of microaggregates thereby increasing the porosity. As such these soils are freely drained and lacking in features which can be used to differentiate the horizons. Chemical Properties The chemical properties of the soils are given in Table 2. The soils can be similarly grouped into those that formed on parent rocks low in weatherable minerals and total iron content (phyllite, graphitic sericite schist and quartz-mica schist) and those that formed over amphibole and serpentinite. The first group of soils show almost similar chemical properties: low pH, low CEC and base saturation, high aluminium saturation and low free iron content (by dithionite extraction).
PERTANIKA VOL. 11 NO. 3, 1988
PROPERTIES OF SOILS DERIVED FROM SOME METAMORPHIC ROCKS TABLE 2 Chemical and physical properties and mineralogy of the B horizons of soils developed on some metamorphic rocks in Peninsular Malaysia.
^•N.
Parent
SoU Nv Properties ^ " s .
CHEMICAL
pH (H2O) Charge Characteristics (c mol (+)/kg soil
Amphibole Schist
4.5
4.9
CEC < 12.5
CEC > 16
CEC < 1 6
ECEC
ECEC < 5
Al Saturation (ECEC basis)
>85%
<5
CEC < 12.5 ECEC < 16
Serpentinite
5.3 CEC < 1 0 ECEC < 1 0
<30% >70%
>70%
3.4-4.1%
3.4-4.9%
10.6-15.6%
16.8-19.6%
Quartz
Quartz
Quartz
Quartz
Quartz
Silt
Quartz Kaolinite Muscovite
Quartz Kaolinite Muscovite
Quartz, Kaolinite Mica-smectite
Gibbsite Goethite Kaolinite
Gibbsite Kaolinite Goethite
Clay
Kaolinite Quartz Muscovite
Kaolinite Quartz Muscovite
Kaolinite, MicaSmectite, Quartz
Kaolinite Goethite
Kaolinite Goethite
1.55-1.69
1.40-1.54
1.27-1.35
11-14
13
O
fi
MINE
4.3
Quartz-Mica Schist
ECEC < 12.5
< 10%
Sand
KYSIC
4.5
Base Saturation
Free iron oxide (dithionite)
<:
Graphitic Sericite Schist
Phyllite
Bulk Density (g/cm 3 ) Plasticity Index Cole
2.6%
1J3 14 0.015-0.020
8 0.011-0.012
The soils formed on the amphibole and the serpentinite have higher pH, high base saturation (especially on serpentinite), low aluminium saturation and high free iron oxide content. The slightly higher pH in these two soils may be due to the low or absence of extractable aluminium. Mineralogy The mineralogy of the sand, silt and clay fractions of the soils developed on the metamorphic rocks studied is given in Table 2. The sand fractions of the soils are dominated by quartz. The silt fractions of the soils developed over the phyllite, graphitic sericite schist and quartz-mica schist are composed essentially of kaolinite, quartz and muscovite. The silt fractions of the quartz-mica schist also contain mica-smectite, which is one of
0.010-0.018
<10*
0.015-0.020
0
1.25 8 nd
the alteration products of biotite. The muscovite in the graphitic sericite schist and phyllite shows some alteration as can be seen by the opening up of the 1.0 nm peaks. The silt fractions of the soils developed over amphibole schist and serpentinite are composed of kaolinite, gibbsite and goethite. In general, the clay fractions of all the soils are dominated by kaolinite. Traces of quartz and muscovite occur in the first three profiles whilst in the last two profiles, there are traces of goethite. Physical Properties The physical properties of the B-horizons of the soils studied are given in Table 2. The bulk density of the soils developed over phyllite, sericite schist and quartz-mica schist is slightly greater
PERTANIKA VOL. 11 NO. 3, 1988
379,
S. ZAUYAH TABLE 3 Some micromorphological properties of the B-horizons of soils developed over some metamofphic rocks in Peninsular Malaysia. Parent ^^Rock Properties^^^
Phyllite
Graphitic Sericite Schist
Quartz-Mica Schist
Amphibolite Schist
Serpentinite
Micro structure
Angular blocky
Subangular blocky
Subangular blockychannel
Granular
Granular
c/f related distribution
Open porphyric
Double spaced porphyric
Single spaced porphyric
Open porphyric
Open porphyric
b-fabric
Reticulate striated
Reticulate striated
Mosaic-speckled
Stipplespeckled and cross striated
Stipplespeckled and undifferentiated
Common Pedofeatures
Ferruginous nodules Clay coatings. Channel infillings a. Loose, continuous, excremental infillings. b. Dense, complete excremental infillings. c. Dense, complete, clay infillings.
than the bulk density of soils developed over the amphibole schist and the serpentinite. The soils in the second group have been so bioturbated that they have become very porous due to numerous channels. There is little variation in plasticity indices of these soils which range from 8 to 14. The COLE values too showed no difference between these two groups of soils. All the COLE values are low ranging from 0.01 to 0.02. Micromorphological Properties The micromorphological properties of the B horizons of the soils studied are given in Table 3. Blocky microstructures are the norm for the soils developed over phyllite, sericite schist and quartzmica schist. On the other hand, soils which developed over amphibole schist and the serpentinite show granular microstructures. The c/f (coarse to fine) related distribution at the limit of 5 fjm is porphyric for all the soils. The quartz-mica schist and sericite schist contain much more quartz grains which dominate the coarse fractions thus making the c/f related distribution single-spaced to double-spaced 380
Ferruginous nodules Clay coatings. Loose, continuous excremental infillings in channels.
Channel infillings: a.
b.
Dense, complete excremental infillings with bow-like internal fabric. Loose, continuous excremental infillings.
porphyric. Since the coarse grains make up a small percentage of the groundmass of the soils developed over amphibole schist and the serpentinite, the c/f related distribution is open porphyric. A range of b-fabrics (birefringence) was observed in the profiles studied. The profiles formed over sericite schist and phyllite show reticulate b-fabric. Porostriated and granostriated b-fabrics are present in some small areas. The quartz-mica schist shows mosaic speckled b-fabric. The striations in the first two profiles may be caused by the preferred orientation of the clay minerals due to pressure and also to wetting and drying. The profiles formed over serpentinite and amphibole schist show a dominance of stipple speckled b-fabric. ^ Pedofeatures found in the first three profiles are similar i.e. clay coatings with strong orientations in channels (Plate l)f ferruginous nodules disseminated in the groundmass and channel infillings. These infillings are of three types: a. loose, continuous and discontinuous excremental infillings (Plate II).
PERTANIKA VOL. 11 NO. 3, 1988
I 8
8 0 tn
I
8 2 Coating of speckled clay with strong orienta tions in channel
Plate II
Loose, discontinuous excremental in fillings in channel.
m
O
i o
I
Dense, complete clay infillings in channel.
Plate IV:
Dense, complete excremental infillings with bow-like internal fabric in channel
S. ZAUYAH
b. c.
dense, complete excremerftal infillings. dense, complete clay infillings (Plate
nil Pedofeatures in the soils over serpentinite and amphibole are only those formed by faunal activity i.e. channel excremental infillings which are of two types: a. loose and continuous b. dense, complete with bow-like internal fabric (Plate IV). Relationship between some Physico-chemical Properties and B-Fabric of the Soil Many workers (Stoops, 1967; Flach et at. 1968; Nettleton et at. 1969; Eswaran, 1972 and Zainol, 1984) have attemptd to relate the birefringence fabric (b-fabric) which was previously termed as the plasmic fabric (Brewer, 1964) to some soil properties such as COLE values, free iron oxide content and mineralogy. A comparison of the b-fabric with free iron oxide content (extractable by dithionite and oxalate) COLE values and plasticity indices are given in Table 4.
The dithionite-extractable free iron oxide content of the profiles developed over the phyllite, graphitic sericite schist and quartz-mica schist ranges from 2 to 5 %. In this range of values, the b—fabrics observed are reticulate striated and mosaic speckled. In the profiles developed over amphibole schist and serpentinite, the dithionite extracted free iron oxide content range from 10 to 20 %. In these soils, stipple speckled to undifferentiated b-fabrics are observed. There is thus some influence of the free iron oxide content over the b-fabrics of the soils. The amorphous iron (FeQ) content is also much higher for the serpentinite soils when compared to the other soils. Presence of a high amount of amorphous iron may also be the reason for the undifferentiated b-fabric. This amorphous iron coats the clay minerals (observed under SEM) and thus masked its orientation. A high free iron content « 18 %) was also found in soils with isotic plasmic fabric by Eswaran (1972). The author believes that the size of the clay particles may play a part in the b—fabric of the soils. As observed under SEM, the clay particles in the first three profiles are definitely larger (1 to 2
TABLE 4 Comparison between b-fabric and free iron oxide, COLE and PI of. soils formed over some metamorphic rocks in Peninsular Malaysia. Parent Rock
Horizon
B-Fabric
Free Iron Oxide % Fe
Quartz Mica Schist
Graphitic Sericite Schist Phyllite
Amphibole Schist Serpentinite
382
Btl Bt2 Bt3 Bt4
d
Fe
COLE
P.I.
o
Mosaiospekled
3.44 3.85 4.88 4.06
0.45 0.64 0.61 0.33
0.019 0.018 0.012 0.013
11 14 12 12
Best Btl B/C
Reticulate and granostriated Reticulate and mosaicspeckled
4,14 3.42 2.0
0.26 0.10 0.08
0.011 0.012 0.008
8 8 8
Btl Bt2 B/C
Reticulate and poro striated id to Btl Reticulate and cross striated
3.11 2.60 2.74
0.90 1.29 0.19
0.016 0.020 0.018
13 14 14
Bol Bo2 Bo 3
Stipple-speckled and cross striated
11.60 10.60 15.61
0.20 0.26 0.14
0.021 0.020 0.015
12 13 8
Bol Bocl Bo2
Undifferentiated and stipple-speckled
19.60 16.03 16.75
2.23 0.83 0.55
nd
12 nd 8
PERTAN1KA VOL. 11 NO. 3, 1988
PROPERTIES OF SOILS DERIVED FROM SOME METAMORPHIC ROCKS TABLE 5 Some properties of the major B soil horizons and the weathering stage of soils developed over five metamorphic rocks in Peninsular Malaysia. (Amt. = Amount) Properties Of Major B Soil Horizons Parent Rock
Weathering Stage Of Soil ^ 1 9 8 5 )
Fine Silt clay
Clay Coatings
>25
0.3-0.7
Present
Intermediary (Stage 3)
Quartz Mica Schist
<25
0.5 - 0.8
Present
Intermediary (Stage 3)
Phyllite
<25
0.3-0.5
Present
Intermediary (Stage 3)
Serpentinite
<25
0.2
Absent
Ultimate (Stage 4)
Amphibole Schist
<25
Absent
Ultimate (Stage 4)
ACEC Meq/lOOgclay
Graphitic Sericite Schist
Amt. of weatherable mineral in sand fraction
0.06-0.09
jL/m) whereas in the amphibole schist and the serpentiniite soils, they are finer (some even less than 0.5 /xm). Thus orientation of larger kaolinite and also muscovite grains in the micromass of the graphitic sericite schist and phyllite may acount for the reticulate b-fabric. Weathering Stage of Soil Many criteria have been used to indicate the stage of weathering of soils. Van Wambeke (1962) used the fine silt/clay ratio for tropical soils. Sys (1985) suggested the following set of criteria to indicate the stage of ferrallitic weathering: 1. Apparent CEC - more or less than 25 meq/lOOg clay. 2. Amount of weatherable minerals in the sand fractions - more or less than 5%. 3. Fine silt/clay ratios (intermediary stage : < 0.15 for metamorphic rocks) 4. Development of soil structure. 5. Presence or absence of clay coatings. Using these criteria, the weathering stage of each of the soil studied has been determined (Table 5). The soils developed over quartz-mica schist, phyllite and sericite schist are in the intermediary stage, whilst soils developed over the
serpentinite and the amphibole schist have the characteristics of the ultimate stage of weathering. CONCLUSIONS
This study shows that many of the properties of soils developed over metamorphic rocks in Peninsular Malaysia are dependent on the parent material from which the soils are formed. The soils studied can be divided into two broad groups: those forming over rocks low in weatherable minerals and those forming over rocks high in weatherable minerals. There are differences in profile morphology, colour, texture, structure, drainage, mineralogy, some micromorphological properties and some physico-chemical properties. The first group of soils are determined to be in the intermediary stage of weathering whilst the second group is in the ultimate stage of weathering. ACKNOWLEDGEMENT
The author wishes to thank Universiti Pertanian Malaysia for the research grant extended towards this study. REFERENCES BRASHER, B.R., D.P. FRANZMEIR, V.T. VALASSIS and S.E. DAVIDSON. (1966): Use of saran resin to coat natural clods for bulk density and waterretention measurements. 56/7 Sci. 101: 108.
PERTANIKA VOL. 11 NO. 3, 1988
386
S. ZAUYAH BREWER, R, (1976): Fabric and mineral analysis of soils. Robert E. Kreiger Publishing Co., New York 428 p.
LOH. K.F. (1981): A study of some soils derived form sedimentary rocks in the state of Pahang, Malaysia. M. Sc. Thesis, State Univ. Of Ghent. 167p.
BSI No. 1377. (1975): Methods of testing for soils for civil engineering purpose. 143 p.
MEHRA, P.O. and M.L, JACKSON. (1960): Iron oxide removal from soils and clays by dithionite-citrate system buffered with sodium bicarbonate. Clay and Clay Mineral, 7th Int. Conf: 317-342.
BULLOCK, P., N. FEDOROFF, A. JONGERIUS, G. STOOPS and T. TURSINA. (1985): Handbook for soil thin section description. Waine Res. Publ. England. 152 p. CHAPMAN, J.D. (1965): Cation exchange capacity. In: C.A, Black, D.D. Evans, J.L. White, L.E. Ensminger and F.E. Clark (Eds). Methods of soil analysis. Part 2. Serie Agron. no. 9. Amer. Soc. Agron. Madison, U.S.A.: 891-901. ESWARAN, H. (1972): Micromorphological indicators of pedogenesis in some tropical soils derived from basalts from Nicaragua. Geoderma. 7: 15-31, ESWARAN, H. and WONG CHOW BIN. (1977): A study of a deep weathering profile on granite in Peninsular Malaysia. I; Physico-chemical and micromorphological properties. Soils Set Soc. Am. J. 42: 144-149. FOA. (1977): Guidelines for soil profile description. Rome 66p. FLACH, K.W., J.G. CADY and W.D. NETTLETON. (1968): Pedogenetic alteration of highly weathered parent materials. Transactions of the 9th Int. Congr, Soils Sci. 4: 343-351. GROSSMAN, R.B., B.R. BRASHIER, D.F. FRANZMEIER and J.L. WALKER. (1968): Linear extensibility as calculated from natural-clod bulk density measurements. Soils Sci. Soc. Amer. Proc. 32: 570-573. LIM, J.S. (1977): A study of some highly weathered soils of the state of Pahang, Malaysia. M. Sc. Thesis. State Univ. Of Ghent. 174 p.
384
NETTLETON, W.D., K.W. FLACH and B.R. BRASHIER. (1969): Argillic horizons without clay skins. Soils Sci. Soc.Am. Proc. 33: 121-125. PARAMANANTHAN,S. (1977): Soil Genesis on igneus and metamorphic rocks in Malaysia. Doctorate Thesis, State Univ. of Ghent, 302p. STOOPS, G. (1967): Le profil d'alteration au BasCongo (Kinshasa). Sa description et sa genese. Pedologie, 17(1): 60-105. SYS, C. (1985): Pedogenesis. Introduction to the study of soil forming processes. ITC Lecture notes. State Univ. of Ghent. 41p. USDA SOIL SURVEY MANUAL. (1981): Revision of Chap. 4: Examination and description of soils in the field. 430-V-SSM, May, 1981. 170 p. VAN WAMBEKE, A, (1962): Criteria for classifying tropical soils. /. Soil Sci. 13: 124-132. YUAN, T.L. (1959): Determination of exchangeable hydrogen in clays by titration method. Soil Sci. 88: 164-167. ZAINOL, M.E. (1984): Contribution to the study of inland soils of Kedah, Peninsular Malaysia. Doctorate Thesis, State Univ. of Ghent. 308p. ZAUYAH, S., S. PARAMANANTHAN and A.B. ROSENANI. (1984). Properties of soils derived from sedimentary rocks of Malaysia. Proc. 5th ASEAN Soil Conf. Bangkok, 11;D6 1-9.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 9 April, 1988)
Pertanika ll(3>, 385-391 (1988)
Some Changes in Chemical Characteristics of a Paddy and Mangrove Soil Sample during Submergence in Water J. MARCUS, M. FARIDAWATI, and M.L. ZALMA Faculty of Science and Natural Resources UKM Campus Sabah, 88996 Kota Kinabalu Sabah, Malaysia. Keywords: Submergence; reduction; paddy, mangrove ABSTRAK Kajian ini dijalankan untuk menentukan perubahan beberapa ciri kimia bagi sampel tanah sawah padi dan tanah paya bakau yang kering apabila kedua-duanya ditenggelami air dalam jangka masa 0 hingga 10 minggu. Keupayaan redoks tanah dan pHy EC, kepekatan kation (Nay K, Mgt Ca, Fe dan Mn), dan kepekatan nitrogen, fosfat dan sulfat bagi larutan tanah berkenaan mengalami perubahan akibat penenggelaman. Perubahan tersebut paling ketara dalam masa dua minggu yangpertama penenggelaman. ABSTRACT This study was conducted to determine the changes of some chemical characteristics of a dry paddy and mangrove soil sample when submerged in water for a 0 - to 10 - weeks period. The redox potential of the soils and the pH, EC, concentrations of cations (Na, K, Mg, Ca, Fe and Mn) and concentrations of nitrogen, phosphate and sulphate of the soil solutions changed due to submergence. The changes were most significant during the first two weeks of submergence.
INTRODUCTION Under aerobic conditions, soil microorganisms such as aerobic bacteria use oxygen for respiration. Oxygen has a high affinity for electrons and thus acts as an electron acceptor during the respiration process. The oxygen is, however, reduced to water during the process (Rowell, 1981). When a soil is submerged in water, gas exchange between soil and air is drastically curtailed and therefore the oxygen supply to the soil is cut off. Within a few hours of submergence, microorganisms use up the oxygen present in the water or trapped in the soil and render a submerged soil practically devoid of oxygen (Evans and Scott, 1955; Scholander etaL, 1955; Takal etaL, 1956; Ponnamperuma, 1965; Turner and Patrick, 1968). Due to this anaerobic condition the aerobic organisms can no longer function and so either die or go into a resting stage. However, soils contain a wide variety of microorganisms and, in the absence of oxygen, anaerobic bacteria proliferate. These anaerobes
utilize oxidized soil components such as nitrate, manganic oxides, ferric oxides, sulphate, phosphate and dissimilation products of organic matter as electron acceptors in their respiration and thus reduce them (Ponnamperuma, 1965). The change from aerobic to anaerobic conditions causes changes in chemical properties of the soil and soil solution (Patrick, 1960; IRRI, 1964; Ponnamperuma, 1965, 1972, 1978; Patrick and Reddy, 1978; Yamane, 1978; Ponnamperuma, 1981). The objective of this study was to investigate the changes in some chemical characteristics of a Malaysian paddy and mangrove soil sample when both were changed from aerobic to anaerobic condition through submergence under water. MATERIALS AND METHODS Waterlogged samples of both paddy and mangrove soils were taken from their respective locations in Sabah Malaysia and, on reaching the lab, the soils
J. MARCUS, M. FARIDAWATI AND M.L. ZALMA
were filtered to get their respective soil solutions using filter paper No. 43 under vacuum conditions. The redox potential (Eh) of the wet soils and the pH as well as electrical conductivity (EC) of the soil solutions were later determined. The wet soils were then air dried and their < 2 mm fractions were collected through sieving. The pH, Eh, EC, amounts of exchangeable cations (Na, Mg, Ca, Al, H, Fe and Mn) and amounts of water - soluble phosphate and sulphate were determined. In order to investigate the effect of submergence, 400g of each soil sample were put in plastic containers and then filled with 300 ml of distilled water and mixed thoroughly to give a soil paste. Excess distilled water was then added until the water level was at least 4 cm above the soil surface. The soil was left in the submerged state for 0, 2, 4, 6, 8 and 10 weeks. Water level in the respective containers was maintained throughout the designated period without disturbing the soil. The above experiment was carried out in duplicate for both soil samples. When each period of submergence was achieved (the "zero" week sample was taken immediately after mixing the soil with water), excess water above the soil surface in each plastic container was removed and discarded. The Eh of the wet soil was then measured before the soil was filtered to give a soil solution. The pH, EC, concentrations of cations (Na, K, Mg, Ca, Fe and Mn) and concentrations of nitrogen, phosphate and sulphate in the soil solution were then determined. Throughout the above analyses, redox potential was measured directly using a Pt redox electrode. The electrode was initially standardised using a standard redox buffer which has an Eh of 430 mV (Light, 1972). The electorde was inserted into the respective wet soils up to a 6 cm depth and the Eh reading was recorded after 10 minutes. The pH and EC of the soil solutions were determined using a pH electrode and a conductivity electrode respectively. The pH and EC of the soils were, however, measured in a 1 : 2 suspension of distilled water. Exchangeable cations (Na, Mg, Ca, Al, H, Fe and Mn) of the soils were extracted using 1.0 M KC1 and then determined by atomic absorption spectrophotometry (AAS). Cations in the soil solutions were determined directly by AAS. Water-soluble phosphate and sulphate in 386
the soils and soil solutions were determined colourimetrically (Murphy and Riley, 1962) and by the turbidity method (using BaCl2) respectively. Total nitrogen in the soil solutions was determined using a micro-Kjeldhal method (Page et al, 1982), while ammonium was directly measured using a ammonium electrode. RESULTS AND DISCUSSION Some chemical properties of the paddy and mangrove soil samples studied are shown in Table 1. The wet (freshly taken from the field) and dry samples had negative and positive potentials, respectively. Soils with a negative Eh were considered to be in a reduced state, while those with positive Eh were considered to be in an oxidized state (Patrick and Mahapatra, 1968; Ponnamperuma, 1972). Therefore the wet and dry soils were in a reduced and oxidized state, respectively. The pH of the wet soils was always about pH 7. This value is common for soils under reduced conditions (Patrick and Reddy, 1978). The dry soils were, however, acidic. The EC of the mangrove soil was relatively high compared to the paddy soil, which might be due to the influence of sea water on the mangrove soil. Similarly organic matter content, water — soluble sulphate and exchangeable basic cations were significantly higher in the mangrove soil. Changes in Eh of the soil samples upon submergence are shown in Figure l(a). The Eh values of both soils decreased to a stable value of about — 200 mV after two weeks of submergence. Similar changes in Eh of submerged soils have been shown by IRRI (1964), Ponnamperuma (1965, 1972), and Yamane (1978). The rapid decrease in Eh was apparently due to a release of reducing substances accompanying oxygen depletion before Mn (IV) and Fe (III) oxide hydrates could mobilize their respective buffer capacities (Ponnamperuma, 1972, 1981). The decrease in Eh of most acid soils can be explained quantitatively in terms of the potential of the Fe(OH)3 - Fe 2 + redox system (Ponnamperuma et al, 1966; Yamane, 1978). The rapid decrease could also be related to the high organic matter contents of the soils (Yamane, 1978). The pH of soil solutions from both soils increased to a stable value after two weeks of submergence. The final pH of the paddy soil
PERTANIKA VOL. II NO. 3, 1988
CHANGES IN CHEMICAL CHARACTERISTICS OF A PADDY AND MANGROVE SOIL DURING SUBMERGENCE TABLE 1 Some chemical properties of the paddy and mangrove soil samples MANGROVE
PADDY wet
dry
wet
dry
pH (1 : 2)
6.7
4.5
7.2
5.9
Eh (mV)
-180
360
1.8
0.9
EC (mmhos cm" )
-200 52.5
420 19.5
Exchangeable -1 cations (meq 100 g Na
0.2
K
0,7
Ca
0.4
Mg
0.9
Al
1.0
H
1.1
62.5 2.6 10.8 38.9 0.0 0.1
Exchangeable (/Xg g~ ) Fe
178.0
Mn
74.0
278.0 6.4
Water soluble ()Ltg g" 1 ) Phosphate
25.0
0.6
Sulphate
1450
3125
was higher than for the mangrove soil, as shown in Figure l(b). Similar changes in the pH of submerged acid soils have been shown by Tomlinson (1957) and Ponnamperuma (1965, 1972). During reduction of the inorganic components of the soil under anaerobic condition, protons were used and therefore the pH increased. The increase in pH of most acid soils upon submergence is largely due to the reduction of Fe (III) to Fe(II) (Ponnamperuma et al, 1966). The fairly stable pH attained after two weeks of submergence was due to the consumption of initially - present electron acceptors and the sustained production of carbon dioxide. The significantly lower pH of the mangrove soil could reflect a small amount of acid-sulphate property for this soil with its high organic matter content. Changes in EC of the soil solutions of the two submerged soils are shown in Figure 2(a). Note the break in scale between the two soil curves. The EC of the submerged paddy soil attained a maximum during the second week, and then declined to a fairly stable value. Ponnamperuma (1965, 1972) obtained similar results.
The EC of the mangrove soil, however, reduced rapidly to a fairly stable value during submergence. The changes in conductance reflect the balance between reactions that produce ions and those which inactivate them, and it seems that inactivation of ions was prevalent in the mangrove soil. There was a close similarity between the changes in EC and the changes in total soluble cations (Na, K, Ca, Mg, Fe and Mn) in the soil solutions as shown in Figure 2(b). The cations were mainly Na and Mg for the mangrove soil, reflecting seawater influence. The EC increased as the total cation concentration increased, and vice versa. The increase in cations of the soil solution was due to the release of soluble Fe(II) and Mn(II) during the reduction of Fe(III) and Mn(IV), respectively, with some of these cations then being displaced from exchange sites on soil colloids into the soil solution (Ponnamperuma, 1972). Figure 3(a) and Figure 3(b), respectively, show the changes in Mn and Fe concentrations of the soil solutions during submergence. Concentrations of both Mn and Fe in the soil solution of
PERTANIKA VOL. 11 NO. 3, 1988
387
J. MARCUS, M. FARIDAWATI AND M.L. ZALMA
(a) 7.0
6.0
5.0
4.0
4
8
6
4
10
6
10
weeks
weeks
Fig. 1: Changes in Eh and pH of the soils and soil solutions, paddy (x) and mangrove (•), respectively during submergence.
the submerged paddy soil attained a maximum during the second week and then gradually declined to a fairly stable value. Similar changes were observed by Ponnamperuma (1965, 1972, 1981). The initial increase in Mn and Fe was due to the reduction of solid-phase Mn(IV) and Fe (HI) compounds, functioning as electron accep-
tors, during respiration of anaerobic bacteria. The reduction produced more-soluble Mn(II) and Fe (II) compounds and therefore the amounts of these ions in solution increased (IRRI, 1964; Ponnamperuma, 1972). The decrease in concentration might similarly be due to immobilization of Mn and Fe. The soluble Mn and Fe could pre4000
(a)
.
40
3000
30
2000
~
1.0
200
100
0.5
2
4
6
8
10
2
4
weeks Fig. 2: Changes in Ec and total cations concentration of the soil solutions, paddy (x) and mangrove (*), during submergence.
388
PERTANIKA VOL. 11 NO. 3, 1988
6 weeks
8
10
CHANGES IN CHEMICAL CHARACTERISTICS OF A PADDY AND MANGROVE SOIL DURING SUBMERGENCE
(b)
(8)
10
10
10
weeks
weeks
Fig. 3: Changes in Mn and Fe concentrations in the soil solutions, paddy (x) and mangrove (*)t during submergence.
cipitate as carbonate (Ponnamperuma et aL, 1969) and as sulphide (Ayotade, 1977) as well as being sorbed by Mn(IV) and Fe(III) oxide hydrates. Changes in Mn and Fe of the submerged mangrove soil were slightly different from those of the paddy soil. The concentration of Mn decreased to a fairly stable value during submergence, with the overall values being lower than those obtained for paddy soil. This might be due to the low level of active Mn in the mangrove soil, as indicated by the low amount of exchangeable Mn (Table 1). Soils with low contents of Mn normally show only slight changes of soluble Mn during submergence (Ponnamperuma, 1965). The total nitrogen in the soil solutions of the submerged soils is shown in Figure 4(a). Both soils showed an increase in total nitrogen during the first two weeks of submergence which then later decreased once more. In a submerged soil, nitrateN that is initially present should be reduced to gaseous nitrogen, while the mineralization of organic nitrogen should not proceed to the nitrification stage producing nitrate-N due to the absence of oxygen. The process stops instead at the ammonification stage, thus producing ammoniumN (Patrick and Mahapatra, 1968; Patrick and Reddy, 1978).
Comparison between the total nitrogen and ammonium-N in the soil solutions indicates that ammonium-N contributed significantly to the total nitrogen. Higher amounts of ammonium-N were detected in the mangrove soil, which could be related to its high content of organic matter. One of the main factors affecting the production of ammonium-N is the organic matter content of a submerged soil (Ponnamperuma, 1965). The subsequent decrease in ammonium-N concentration could be due to fixation by soil colloids. Phosphate and sulphate contents of soil solutions from submerged soils also changes during submergence, as shown in Figure 4(b) and Figure 4(c). Phosphate concentration decreased to a minimum during the first four weeks of submergence and then later increased slightly once more. These changes were not similar to the results obtained by Ponnamperuma (1965, 1972), where phosphate concentration increased initially and then decreased once more. The principal effect of anaerobic conditions on phosphorus in soils is a change in the solubility of phosphate (Ponnamperuma, 1972). The increase in pH during submergence will normally increase the concentration of phosphate in the soil solution, since iron and aluminium phosphates liberate
PERTAN1KA VOL. 11 NO. 3, 1988
389
J. MARCUS, M. FARIDAWATI AND M.L. ZALMA
(a)
i
a
(b)
70
•g 3000
60
3 2000
I 50
£
1000
S §
S 30
I" Z
a
10
60 60 40 20
4
6
8
10
2
weeks
4
6
10
weeks Fig. 4: Changes in nitrogen phosphate and sulphate of the soil solutions, paddy (x) and mangrove (a), during submergence.
phosphate ions as the pH rises (Larsen, 1967). The reduction of Fe(III) to Fe(II) will also cause a release into solution of adsorbed, chemicallybound and occluded phosphate. These might be the likely reasons why the phosphate concentrations in the soil solution of the submerged paddy and mangrove soils increased after decreasing initially. The initial decrease in concentration might be due to a precipitation of phosphate. The concentrations of Mn and Fe were highest during the initial stage of submergence. Therefore a proportion of the Mn and Fe released into the soil solution might form lesssolube precipitates with the phosphate. The concentration of sulphate in the soil solution of the paddy soil increased during the initial stage of submergence and then decreased. In contrast, it decreased first and then increased for the mangrove soil. Values were also much higher for this soil, further suggesting an acidsulphate nature for it as discussed previously for pH effects. The changes in water soluble-sulphate varies widely with soil properties (IRRI, 1965; Ponnamperuma, 1981). Reduction of sulphate to sulphide would cause a decrease in sulphate concentration of the soil solution, while 390
release of sulphate from anion-exchange sites as pH increased would increase its concentration instead. CONCLUSION Submergence of dry paddy and mangove soils changed them from an oxidized to a reduced state, as shown by their negative redox potentials. This resulted in changes in the chemical characteristics of the soil solutions. Therefore, soils under oxidized conditions have different chemical characteristics compared to soils under reduced conditions. REFERENCES AYOTADE, K.A. (1977): Kinetics and reactions of hydrogen sulphide in solutions from flooded rice soils. PL Soil. 46: 381-389. EVANS, D.D. and A.D. SCOTT (1955): Soil Sci. Soc. Amer.Proc. 19: 12-16. IRRI (1964, 1965). International Rice Research Institute, Annual Reports. IRRI, Los Banos, The Philippines. LARSEN, S. (1967): Soil phosphorus. Adv. Agron. 19: 151-210.
PERTANIKA VOL. 11 NO. 3, 1988
CHANGES IN CHEMICAL CHARACTERISTICS OF A PADDY AND MANGROVE SOIL DURING SUBMERGENCE LIGHT, T.S. (1972): Standard solution Tor redox measurement. Anal Chem., 44: 1038-1039. PATRICK, W.H. (1960): Nitrate reduction in a submerged soil as affected by redox potential. Trans. 7th Int. Congr. Soil Sd., Madison. 2: 494-500. PATRICK, W.H. and I.C. MAHAPATRA (1968): Transformation and availability of nitrogen and phosphorus in waterlogged soils. Adv. Agron. 20: 323-358. MURPHY, J. and J.P. RILEY (1962): A modified single solution method for determination of phosphate in natural waters. Anal Chim. Acta. 27: 31-36. PAGE, A.L., R.H, MILLER, and D.R. KEENEY (1982): Methods of Soil Analysis, Part 2. Am. Soc. Agron. Inc. Madison, Wisconsin. PATRICK, W.H. and C.N. REDDY (1978): Chemical changes in rice soils, pp. 361-379. In: Soils and Rice. IRRI, Los Banos, The Philippines. PONNAMPERUMA, F.N. (1965): Dynamic aspects of flooded soils and the nutrition of the rice plant, pp. 295-298. In: Mineral Nutrition of the Rice Plant. Johns Hopkins Press, Baltimore, Maryland, USA. PONNAMPERUMA, F.N. (1972): The chemistry of submerged soils. Adv. Agron. 24: 29-96. PONNAMPERUMA, F.N. (1978): Electrical changes in submerged soils and the growth of rice, pp. 421-444. In: Soils and Rice. IRRI, Los Banos, The Philippines.
PONNAMPERUMA, F.N. (1981): Some aspects of the physical chemistry of paddy soils, pp. 59-94 in Institute of Soil Science, Academia Sinica (ed.) 1981. Proceedings of Symposium on Paddy Soil, Nanjing, China. Science Press, Beijing. 864 pp. PONNAMPERUMA, F.N., E. MARTINEZ and T. LOY (1966): Influence of redox potential and partial pressure of carbon dioxide on pH values and the suspension effect of flooded soils. Soil Sci. 108: 48-57. ROWELL, D.L. (1981): Oxidation and reduction, pp. 401-461. In: The Chemistry of Soil Processes. Greenland, D.G. and Hayes, M.H.B. (eds.). John Wiley and sons, New York. SCHOLANDER, P.F., L. VAN DAM and S.I. SCHOLANDER (1955): Amer. J. Bot 42: 92-98. TAKAI, Y., T. KOYOMA and T. KAMURA (1956): Soil Plant Food (Tokyo), 2: 63-66. TOMLINSON, T.E. (1957): Seasonal variation of the surface pH value of some rice soils of Sierra Leone. Trop.Agr. 34(4): 287-296. TURNER, F.T. and W.H. PATRICK (1968): Chemical changes in waterlogged soils as a result of oxygen depletion. 9th Int. Congr. Soil Sci. IV: 53-65. YAMANE, I. (1978): Electrochemical changes in rice soils, pp. 381-398. In: Soils and Rice. IRRI, Los Banos, The Philippines.
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(Received 3 April 1987)
391
Pertanika 11(3), 393-398(1988)
Evaluation of Potato Crisps Fried in Market Samples of Palm Olein, Corn Oil and Soya Oil M.A. AUGUSTIN1, L.K. HENG1 and NOR AINIIDRIS2 1
Department of Food Science, Faculty of Food Science and Biotechnology, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia 2
Palm Oil Research Institute of Malaysia, P.O.Box 10620, 50720 Kuala Lumpur, Malaysia. Key words: frying, potato crisps, shelf-life, palm olein, corn oil, soya oil. ABSTRAK Hanya penstoran kerepek ubi kentang yang digoreng di dalam minyak kelapa sawit olein, minyak jagung dan minyak kacang soya yang dibeli di Malaysia dinilai oleh panel penilaian deria. Ahli panel terdiri daripada orang tempatan Malaysia. Hayat penstoran kerepek yang digoreng dalam minyak kelapa sawit olein adalah lebih lama (p < 0.05 ) di bandingkan dengan kerepek yang digoreng dalam minyak kacang soya. Didapati bahawa tidak ada perbezaan bererti (p < 0.05) di antara hayat penstoran kerepek yang di goreng dalam minyak kelapa sawit olein dan minyak jagung. Dalam tiap-tiap sistem minyak, hayat penstoran kerepak yang digoreng berkurang dengan penambahan nombor penggorengan. Hayat penstoran kerepek yang digoreng dalam tiap-tiap minyak dipengaruhi oleh paras % komponen polar dan nilai asid minyak sewaktu penggorengan dibuat. ABSTRACT The shelf life of potato crisps fried in market samples of palm olein, corn oil and soya oil was evaluated by a sensory panel comprising Malaysian panelists. The shelf-life of crisps fried in palm olein was significantly longer than those fried in soya oil (p<0.05). There were no significant differences (p < 0.05) between the shelf-life of crisps fried in palm olein and those fried in corn oil In each oil system, the shelf-life of the crisps fried in the oils decreased with an increase in fry number. The shelf life of the crisps fried in each oil was infuenced by the level of % polar components and the acid value of the oil INTRODUCTION The potato crisp industry uses large amounts of oil for frying. The choice of oils for the frying process becomes very important from a view point of quality of the fried product as well as the economy of the production process (Berger, 1984). The quality changes of market samples of palm olein, corn and soya oils during frying have been studied (Augustin et alf 1987). It was found that the extent of oxidation and polymer formation were less in palm olein than in corn and soya oils and that the change in acid value was highest
in palm olein (Augustin at al, 1987). It is also important to assess the keeping quality of the fried product. This paper reports the shelf-life of potato crisps fried in palm olein, corn oil and soya oil. These oils were chosen as they are the more readily available frying oils in the Malaysia market. In the USA, soyabean and other vegetable oils intended for frying are generally hydrogenated to reduce most or all of the linolenic acid but in other countries and in the Middle East, many of the cooking and frying oils are not hydrogenated (Frankel etal> 1984).
M.A. AUGUSTIN, I.K. HENG AND NOR AINI IDRIS
Oil is picked up by the food during frying and forms an integral part of the deep-fried product. The type and quality of the oil used for frying have an influence on the quality and shelflife of the fried product as the oxidative deterioration of lipids is one of the major limiting deteriorative reactions in fried food proucts (Quast and Karel, 1972). The aim of this study was to evaluate the keeping quality of potato crisps fried in market samples of palm olein, corn oil and soya oil in Malaysia. MATERIALS AND METHODS
The experimental procedures except that of the sensory evaluation of crisps have been described before (Augustin etaL, 1987). Materials Random samples of refined, bleached and deodorized palm olein (RBD olein) (Batches 1, 2 and 3), corn oil (Batches 1 and 2) and soya oil (Batches 1 and 2) were obtained from local retail outlets. Batch 3 of soya oil was obtained directly from a local factory. Potatoes were bought from local shops and thus the same variety of potatoes could not be assured every time. The potatoes were cleaned and sliced to a thickness of 2 mm. The slices were blotted slightly with tissue before frying. Frying The frying of potato slices was carried out in Valentine fryers which initially contained 4 kg of oil. The temperature of the oil was raised to 180°C and frying was started half an hour after the temperature had reached 18O°C. Each time, 100 g of slices were fried for 3.0 min. Fryings were carried out at half hour intervals. A total of 10 fryings were done per day for four consecutive days. At the end of each day, the fryer was switched off and 100 g samples of oil were removed from each fryer. The lid of the fryer was replaced and the fryings were continued on the next day. Fresh oil was not added to the frying vessel. Crisps from selected fryings were kept for sensory assessment. Analyses of the Oils The analyses of peroxide value (AOCS, 1974) and p-anisidine value (IUPAC, 1979) were carried out before the start of the frying trials. Other 394
analyses on the frying oils were carried out within three weeks. The percentage polar components of the oils were determined by column chromatography (Billek el al, 1978). The IUPAC methods were used for the determination of acid and iodine values (IUPAC, 1979). The oils were screened for the antioxidants BHA, BHT and TBHQ (Kirleis and Stine, 1978; Page, 1982). Sensory Evaluation For each trial, crisps from selected fryings (1st, 10th, 20th, 30th, and 40th) in palm olein, corn oil and soya oil were taken for sensory evaluation. Crisps from each of these fry numbers were placed in 10 glass sample bottles with screw-capped lids, with each containing 2 potato crisps. The samples were stored at room temperature (21—23°C) and presented at weekly intervals to a sensory panel comprising 10 judges with previous experience on sensory panels for fat and fat-based products. The panelists were instructed to take short sniffs of the samples on removing the lids from the sample bottles. The time for the onset of rancidity was taken when five or more of the panelists judged that the product had an unacceptable odour. This period is taken as the shelf-life of the product. RESULTS
The characteristics of the fresh oils are given in Table 1. The quality changes of market samples of corn, soya and palm olein during frying have been described in detail elsewhere (Augustin et al.f 1987). BHA, BHT and TBHQ were not found in any of the samples of palm olein, BHA was found in all samples of corn oil while TBHQ was found in batch 1 and 2 of soya oil. TBHQ was not detected in soya oil (Batch3) which was obtained from the factory. Although there were differences in the initial oxidation values of the oils and antioxidants were found in market samples of corn and soya oil but not in palm olein, the samples were nevertheless used as the aim was to compare market samples of these oils. Soya oil (Batch 3), which was obtained directly from the factory was also used for comparison. Quality of Potato Crisps Crisps obtained from all fryings in all types of oils were of acceptable quality immediately after frying. Off-odours, resulting from deterioration of oil in the crisps developed during storage.
Prrlanika 11(3), 399-406(1988)
EVALUATION OF POTATO CRISPS FRIED IN MARKET SAMPLES OF PALM OLEIN, CORN OIL & SOYA OIL TABLE 1 Characteristics of the fresh oils Type of oil
Batch
Acid
Iodine
value (meg/kg)
value
value (mgKOH/g oil)
value
Peroxide
p-Anisidine
Palm olein
1 2 3
0.83 1.48 0.84
1.89 1.61 1.05
0.15 0.20 0.34
58.3 58.2 58.5
Corn oil
1 2
3.55 3.74
6.80 7.57
0.16 0.16
130.0 128.1
Soya oil
1 2 3
14.16 2.14
2.70
0.21
136.5
2.23 0.74
0.33 0.22
141.5
0.87
135.2
Each value is the average result of duplicate analyses. The average standard errors of the mean for duplicate analysis were 0.25 meq/kg for peroxide value, 0.5 for p-anisidine value, 0.02 for acid value and 0.2 for iodine value. All oil samples except Batch 3 (soya oil), were random samples obtained from retail outlets. Batch 3 (soya oil) was obtained from a local factory.
The results in Table 2 showed that the shelf-life of crisps fried in market samples of palm olein was significantly longer than that of crisps fried in market samples of soya oil (p <0.05). A separate trial on the comparison of shelf-life of crisps fried in soya oil with low oxidation values (Batch 3) and that of crisps fried in palm olein confirmed that crisps fried in soya oil had a shorter shelf-life. In the latter trial, the shelf-lives of crisps fried in soya oil were 17 weeks, 15 weeks, and 11 weeks respectively for crisps obtained from the 1st, 10th and 20th fryings while the corresponding shelf-lives for those fried in palm olein (Batch 3) were > 17 weeks > 17 weeks and 14 weeks respectively. Although the shorter shelf-life of crisps fried in soya oil (Batch 1) compared to palm olein may in part be attributed to the high initial oxidation state of the soya oil, the results of trials with Batch 2 and Batch 3 of soya oil showed that the type of oil (i.e. soya oil or palm olein) also influenced the keeping quality of the fried product. There were no significant differences (p < 0.05) between the shelf-life of crisps fried in palm olein and corn oil. The shelf-life of crisps fried in soya oil was generally shorter than that of crisps fried in corn oil. Significant differences (p <0.05) were found between the shelf-life of crisps fried in corn and soya oils from Batch 1 only and this may in part be due to the higher initial peroxide
velue of soya oil from Batch 1. A separate trial on the comparison of shelf-life of crisps fried in market samples of soya oil (peroxide value 11.9 meg/kg, p-anisidine value 2.46) and corn oil (peroxide value 3.65 meg/kg,p-anisidine value 6.62) also showed that the shelf-life of crisps fried in market samples of soya oil was shorter than that of crisps fried in corn oil. The shelf-lives of crisps fried in soya oil were 6 weeks, 5 weeks and 3 weeks for the 1st, 20th and 40th fryings, whereas the corresponding shelf-lives for crisps fried in vorn oil were 9 weeks, 9 weeks and 6 weeks respectively. In each frying system, there was a decrease in the shelf-life of the crisps with increase in fry number (Table 2). The decreasing shelf-life of crisps of later fryings might be expected as the quality of the frying oil deteriorates with an increase in frying time and this has an influence on the shelf-life of the fried product (Asap and Augustin, 1986). The effect of frying oil quality on shelf-life in each of the systems can be demonstrated by plots of shelf-life against % polar components (Figure 1) and shelf-life against acid value (Figure 2). The % polar components is a reliable criterion for determining the quality of frying oils (Billek et at., 1987). It is generally believed that acidity of the frying medium also affects the storage stability of a fried-product. The plots of
PERTANIKA VOL. 11 NO. 3, 1988
395
M.A. AUGUSTIN, IX HENG AND NOR AINI IDRIS TABLE 2 Shelf-life of crisps fried in market samples of palm olein, corn and soya oils Batch
Fry Number
Palm Olein
Corn Oil
Soya Oil
1
15
15
12
10
15
15
9
20
13
14
8
30
11
12
3
40
10
9
1 6.6b
12.8a
13.0a
16
18
11
15
15
9
20
13
13
8
30
12
10
6
40
8
6
4
Mean (n = 5) 1 10
12.8a
Mean (n = 5)
12.4ab
7.6b
The shelf-life is taken as the time when 50% or more of the panelists judged that the product had an unacceptable odour Different letters within a row indicate significant differences at P < 0 . 0 5 . The L.S.D.'s at the 5% level are 4.50 and 4.92 for Batches 1 and 2 respectively.
shelf-life against acid value indicate a decrease in shelf-life with increasing acidity. However, these results do not allow us to differentiate between the relative importance of each of these quality parameters, % polar components and acidity, on product stability. DISCUSSION Sensory evaluation of crisps by Malaysian panelists showed that crisps fried in market samples of palm olein without antioxidants were more resistant to the formation of off-odours and off-flavours during storage when compared to those fried in market samples of soya oil containing TBHQ and a factory sample of soya oil without TBHQ. The evaluation also showed that there were no significant differences (p < 0.05) between the shelf-life of crisps fried in market samples of corn oil containing BHA and palm olein without antioxidants. Furthermore, the study showed that the shelf-life of crisps fried in market samples of corn oil containing BHA was generally longer than those fried in market samples of soya oil containing TBHQ. 396
Although this study may be of limited applicability because of the nature of choice of starting oils and the use of Malaysian panelists who were most familiar with products fried in palm olein, it is nevertheless of potential practical significance in Malaysia. The study showed that the type of oil bought from the market in Malaysia and used for frying had an influence on the shelf-life of the fried product. It showed that, despite the presence of antioxidants in market samples of corn and soya oil and the absence of antioxidants in palm olein, the keeping quality of the product fried in palm olein was comparable to those fried in corn oil and better than those fried in soya oil. The susceptibility of crisps fried in soya oil to rancid offodour development may be due to the resence of linolenic acid in the oil. About 7.5% linolenic acid is present in market samples of soya oil (Augustin et ah, 1987). It was interesting to note that there were differences in shelf-life of crisps fried in different types of oil with the same level of % polar components (Figure I). Apart from differences in the level and presence of antioxidants in the different oils at the time of frying which influence keeping
PERTAN1KA VOL. M NO. 3, 1988
EVALUATION OF POTATO CRISPS FRIED IN MARKET SAMPLES OF PALM OLEIN, CORN OIL & SOYA OIL
BATCH 1
20
BATCH 2
20 16
-.16 i
i 12
Ji2
i
'' 8
i 0
8 16 24 32 °/c polar components
0
8 16 24 32 % polar components
Fig. 1: Shelf-life of crisps versus % polar components of the oil at the time of frying. ( o -o palm olein; • • corn oil; a -« soya oil).
20
BATCH 1
BATCH 2
20
16
J12 H »*—
J- 8
I 8 5
4
0
01
0.4 0.8 1.2 IIT acid value (mg KOH/g)
0
0.4 0.8 1.2 17b acid value (mg KOH/g)
Fig. 2: Shelf life of crisps verus acid value of the oil at the time of frying ( o o palm olein; • -# corn oil; &* • soya oil).
quality of the fried product, it may be expected that oils with the same level of % polar components contain differing levels and types of individual breakdown products and different types of unchanged triglycerides which exert differing influences on product stability. Thus although % polar components is regarded as a reliable indicator of frying oil deterioration, prediction of the comparative shelf-life of products fried in
different oils cannot be based on % polar components. However, within each oil system, there is a general trend of decreasing shelf-life with increase in % polar components in the oil.
CONCLUSION These results indicate that market samples of palm olein are comparable to or better than mar-
PERTANIKA VOL. 11 NO. 3, 1988
397
M.A. AUGUSTIN, I.K. HENG AND NOR AINI IDRIS ket samples of corn oil and soya oil respectively for the preparation of fried potato crisps. The intentional use of random samples (and thus the non-avoidance of the differing initial oxidation levels and varying presence of antioxidants in different oils) has limited the applicability of the results but it nevertheless has its usefulness in Malaysia at the present time. Finally, these studies have been based on model experiments in which repeated fryings were carried out without constant topping up of oils and thus differs from commercial deep-frying operations during which oil lost from the fryer is replenished. ACKNOWLEDGEMENTS The authors would like to thank the DirectorGeneral of PORIM for permission to publish the paper. Helpful discussions with Prof. A.S.H. Ong are appreciated. The research was funded by PORIM. REFERENCES AOCS (1974): Official and tentative methods of the American Oil Chemists* Society, Champaign, Illinois, USA, 3rd edn. AUGUSTIN, M.A., L.K. HENG and Y.K. TEAH. (1987): Comparison of the frying performance of market samples of palm olein, corn oil and soya oil in Malaysia. Pertanika, 10(3), 295-304.
398
ASAP, T. and M.A. AUGUSTIN, (1986): Effect of frying oil quality and TBHQ on the shelf-life of potato crisps. / Sci. Food Agric, 37; 1045-1051. BERGER, K.G. (1984): The practice of frying. PORIM Technology. No. 9, May 1984. BILLEK, G., G. GUHR and J. WAIBEL. (1978): Quality assessment of used frying fats: A comparison of four methods./. Amer. Oil Chem. Soc, 55: 728-733. FRANKEL, E.N., L.M. SMITH, CL. HAMBLIN, R.K. CREVELING and A.J. CLIFFORD. (1984): Occurrence of cyclic fatty acid monomers in frying oils used for fast foods. J. Amer. Oil Chem. Soc, 61: 87-89. IUPAC (1979): Standard methods for the analysis of oils, fats and derivatives (Paquot, C. eds.), IUPAC, App. Chem. Div. Commission on oils, fats and derivatives, 6th edn. KIRLEIS, A.W. and CM. STINE. (1978): Retention of synthetic phenolic antioxidants in model freezedried food systems. / Food Set, 43: 1457-1466. PAGE, B.D. (1979): High performance liquid chromatographic determination of nine phenolic antioxidants in oils, lard and shortening. /. Assoc. Off. Anal. Chem. 62: 1239-1246. QUAST, D. and M. KAREL. (1972): Effects of environmental factors on the oxidation of potato chips. J.Food Set, 47:584-587.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 23 June, 1987)
Pertanika 11(3), 399-406 (1988)
The Digestion, Absorption and Utilization of Refined Palm Oil, Palm Olein and Palm Stearin in the Rat T.K.W. NG, H.T. KHOR and Y.H. Chong Division of Human Nutrition, Institute for Medical Research, 50588 Kuala Lumpur Key words : Palm oil; palm olein; palm stearin; soybean oil, digestibility; rate of absorption; utilization; food efficiency
ABSTRAK Satu penilaian pemakanan yang dibuat pada minyak sawit, minyak sawit olein dan minyak sawit stearin dengan menggunakan tikus menunjukkan bahawa pencernaan, kadar resapan dan efisiensi pemakanan minyak sawit olein adalah lebih baik dibandingkan dengan minyak sawit dan minyak sawit stearin. Namun demikian, indeks-indeks pemakanan tersebut.bagi minyak sawit dan minyak sawit stearin tidak berbeza dengan kebanyakan nilai yang biasanya dilapor untuk minyak masak. Pencernaan dan efisiensi pemakanan minyak sawit olein hampir sama dengan indeks-indeks pemakanan tersebut untuk minyak kacang soya, iaitu 97% dan 98% bagi setiap satunya, walaupun kadar resapan minyak sawit olein adalah 10% kurang dari yang boleh didapati dengan minyak kacang soya.
ABSTRACT A nutritional evaluation of refined, bleached and deodorised (RBD) palm oil and its fractionation products, RBD palm olein and RBD palm stearin in terms of their digestibility, rate of absorption and food efficiency in rats, shows that RBD palm olein is a better oil by these criteria than RBD palm stearin and the unfractionated RBD palm oil. Nevertheless, the nutritional indices reported for RBD palm oil and RBD palm stearin are well within the range reported for most cooking oils and fats. The digestibility and food efficiency of RBD palm olein are comparable to that of soybean oil, being 97% and 98% respectively of the values found for the latter oil, although the absorption rate of RBD palm olein is 10% less than that obtained for soybean oil. INTRODUCTION Edible fats and oils are generally 93-98% digestible (Langworthy, 1923; Crockett and Deuel, 1947). The digestibility of palm oil, which was reported to be 97% (Calloway et al, 1956), falls within this range. On the other hand, fats with melting points above 50°C (Crockett and Deuel,
1947), those that contain higher amounts of stearic acid (Mattil and Higgins, 1945) or triglycerides that contain only saturated, long-chain fatty acids (Mattson, 1959) are less readily digested or absorbed. Subsequent studies showed that pancreatic lipase specifically hydrolyse the fatty acids esteri-
*The information reported in this paper is contained in the Ph. D thesis of T.K.W. Ng, University of Malaya, 1987. 1
Department of Biochemistry, University of Malaya, 59200 Kuala Lumpur.
2
Patm Oil Research Institute of Malaysia, Bandar Bant Bangi 43000 Kajang.
T.K.W. NG, H.T. KHOR AND Y.H. CHONG
fled in the I- and 3-positions of a triglyceride (Mattson et al, 1952) and that a high content of palmitic acid esterified at the 2-position of a fat favours its absorption (Tomarelli et al, 1968; Filer et al.t 1969). Generally, only a small percentage of the total palmitic acid of vegetable fats and oils is present in the 2-position of the triglyceride molecule; saturated fatty acids having a chain length of more than 18 carbons are found predominantly in the 1—and 3—positions (Bracco and Bauer, 1978). In an earlier report (Ng, Chong and Khor, 1987), it was shown that the energy availability of RBD palm oil and RBD palm olein are comparable to that of soybean oil. Other than this communication and the very few early fat digestion and absorption studies which included unfractionated refined palm oil reported in the literature, there is practically no information available on common nutritional criteria such as digestibility, absorbability and food efficiency for Malaysian RBD palm olein and RBD palm stearin, although these relatively new processed palm oil fractions are now widely used in cooking and food processing, Thus, there is a fundamental need to establish these nutritional indices mentioned for
Malaysian RBD palm oil, RBD palm olein and RBD palm stearin. The information obtained should be of value to local nutritionists and food technologists involved with product development or marketing in the palm oil industry. MATERIALS AND METHODS
Fat Samples The fat samples investigated (RBD palm oil, RBD palm olein and RBD palm stearin) were obtained from a production plant in Petaling Jaya. Refined soybean oil was used as the reference fat in this study and was purchased from a local supermarket. The nature and composition of these fat samples were ascertained by fatty acid analysis. Determination of Fat Digestibility Ninety 21—25 days old Albino-Swiss strain male rats were divided into 9 uniform groups on the basis of body weight. One group was assigned a fat-free diet (to correct for metabolic fat excreted in the faeces) while the remaining 8 groups were each provided with an experimental diet that
TABLE 1 Composition of the experimental diets Fat level Component (g/ 100 g diet)
0%
10%w/w*
20% w/w*
(21 energy %)
(39 energy %)
0.0
10.0
20.0
Casein
20.0
20.8
23.1
Sucrose
20.0
20.0
20.0
Corn starch
Test fat
(fat-free)
51.0
39.9
26.8
Alphacel
4.0
4.0
4.4
AIN mineral mixture 76
3.5
3.7
4.1
AIN vitamin mixture 76
1.0
1.06
1.18
D L - methionine
0.3
0.3
0.3
Choline bitartrate
0.2
0.2
0.2
370
420
467
kcal per 100 g diet
*Diets containing equal amounts of protein, minerals, vitamins and fibre per kcal 400
PERTANIKA VOL. 11 NO. 3, 1988
THE DIGESTION, ABSORPTION & UTILIZATION OF PALM OIL & ITS FRACTIONATION PRODUCTS IN THE RAT
contained either 10% w/w (21 energy %) or 20% w/w (39 energy %) of each of the four fats studied (see Table 1). Two animals were housed in each stainless steel cage equipped with a wire-mesh floor and removable aluminium tray at the bottom to facilitate the collection and record of spilled food. Food and drinking water were provided ad libitum while food consumption was recorded throughout the 6-week study. The animals were weighed once a week and the faeces for each dietery group were collected for 8 consecutive days at the beginning of the second, fourth and sixth weeks. Faecal lipids were determined using the extraction procedure described by Bligh and Dyer (1959) with a slight modification: IN HC1 was used in place of water in the chloroform-methanol -water mixture ( 1 : 2 : 0.8). The coefficient of digestibility values of the fats studied were calculated from the following formula (Alfin-Slater and Aftergood, 1980): Coefficient
Amount of f a t digestibility consumed
of
=
Amount
Metabolic "Total faecal -fat excreted
Jipid of
x
100
in faeces
tat
by Deuel at al (1940). A correction was made for the quantity of diethyl ether-soluble material which was removable from the GIT of rats fasted for 48 hours but fed no fat. As a criterion for comparing the rates of absorption of the fats investigated, the Absorption Time (AT) 50 was employed i.e. the time after which 50% of the administered fat has disappeared from the GIT (Thomasson, 1956). Statistical Techniques The standard deviation (SD) was used to measure the variation of a particular determination or parameter. Where appropriate, the one-way analysis of variance (ANOVA) was used to compute for differences in weight gain, food consumption, food efficiency, digestibility and rate of fat absorption among the various, experimental groups, using &• = 0.05 as the level of significance (Welkowitze/dt/., 1976). In order to investigate whether two independent factors eg. type of fat and fat level, operate jointly, the F value for interaction was determined using the two-way ANOVA.
consumed
RESULTS AND DISCUSSION Determination of Rate of Absorption of Fats At the end of the digestibility studies, the adult rats were maintained on their respective diets and used later in the absorption tests. Animals weighing about 240 g were fasted for 48 hours at the end of which their body weights were recorded. Each test animal was then very lightly anesthesized with diethyl ether and a known weight of 1.1 ml of test fat was administered via a 2 misyringe connected to a stomach tube. This dosage was equivalent to about 325 mg per 100 cm2 body surface area according to the formula of Lee (1929). For RBD palm oil and RBD palm stearin, there was a need to warm the fats slightly above their melting points and then allowed to cool as much as possible before 1.1 ml of the liquid samples were taken and administered orally. At the end of the two absorption periods investigated i.e. 3 hours and 6 hours, the rats were anesthesized and the intact gastrointestinal tract (GIT) from oesophagus to rectum was removed. Fat remaining in the GIT was extracted and quantified according to the procedure described
Growth and Food Efficiency The amount of food consumed, growth of the experimental animals and food efficiency of the diets used after 6 weeks are summarised in Table 2. On the whole, the experimental diets were well accepted and animals on the same level of dietary fat consumed approximately equal amounts of food irrespective of the type of fat. However, significantly more of a 10%-fat diet (21 energy %) was consumed compared to a 20% fat dht (39 energy %) for each dietary fat except for the case of RBD palm stearin. Growth of the experimental animals, as reflected by weight gain, was comparable in all the dietary groups provided with 21 energy % fat. When the dietary fat level was approximately doubled to 39 energy %, the animals fed the three palm fats still exhibited comparable growth. However, at this higher dietary fat level, only the palm olein-fed animals showed growth that was similar to that of the soybean oil-fed animals, while growth of the animals fed the other two
PERTANIKA VOL. 11 NO. 3, 1988
401
T.K.W. NG, H.T. KHOR AND Y.H. CHONG TABLE 2 Weight gain, food consumption and food efficiency in male rats fed the experimental diets for 6 weeks (Mean ±SD)
Food consumed (g/rat)
Food efficiency4
154.1 ±10.26 169.1 ± 8.97
387 ±17.4 351 ± 7.0
0.398 10.016 0.482 ±0.011
94.813.81 103.2 12.27
37.9 37.8
145.9 ±13.23 165.0 ±13.01
383 ±18.5 352 ±11.6
0.381 10.018 0.468 10.008
90.7 ±4.24 100.2 ±1.81
10 20
37.8 37.7
146.2 ±10.05 159.5 ± 5.40
387 ±21.5 350 ±10.1
0.378 ±0.006 0.453 ±0.011
90.0 ±1.56 97.0 12.34
10 20
37.7 37.8
146.1 ±18.75 157.9 ± 9.88
384 ±22.5 377 ±10.5
0.380^0.011 0.419±0.008
90.6 ±2.73 89.7 ±1.76
8.75
20.5
Dietary level
Initial weight
<%)
(f)
Soybean oil
10a 20 b
37.7 37.8
Palm olein
10 20
Palm oil
Palm stearin
Fat
LDS
12 9
(5%)5
n.s.6
Weight gain2 (g/rat)
Wt gain per
1,000 kcal (g)
0.0158
3.38
3 Mean of 10 rats per experimental group Calculated from 5 pairs of rats per group Food efficiency • Weight gain in grams per gram of food consumed; F value for interaction of fat type and fat level significant by two-way ANOVA for °c= 0.05
5 a
Least significant difference by one-way ANOVA foi «= 0.05 Equivalent to 21 energy %
Equivalent to 39 energy %
palm fats was now significantly lower than that obtained with the reference fat mentioned. The food efficiency, defined as weight gained in grams per gram of food consumed, found for the 39 energy %-fat diets was significantly higher than that obtained for the diets containing the same fat at 21 energy %. While this result was not unexpected, the higher food efficiency obtained with a 39 energy %-fat diet was apparently due to a more efficient utilization of food at this higher level of dietary fat rather than to the greater energy intake attributed to 39 energy %-fat diets per se. This effect of dietary fat level on efficiency of utilization of food can be ascertained by the provision of isocaloric feeding. Alternatively, as was done in the present analysis, 402
Not significant
the influence on weight again by differences in energy intake between animal groups can be excluded to a large extent by expressing the efficiency of utilization of food as weight gain* per 1,000 kcal intake. When this was done, the efficiency of utilization of food generally remained significantly higher with the 39 energy %-fat diets than with the 21 energy %-fat diets. As shown in Table 2, the F value for interaction of fat type and fat level in the 4 x 2 factorial design was found to be significant (
PERTANIKA VOL. 11 NO. 3, 1988
THE DIGESTION, ABSORPTION & UTILIZATION OF PALM OIL & ITS FRACTIONATION PRODUCTS IN THE RAT TABLE 3 Digestibility of the various fats studied
Fat
Dietary level
(%) Soybean oil
Coefficient of Digestibility (%)* Analysis by Analysis by One-way Anova
10 a
98.8 ± 0.13 C
20b
98.8 ± 0.21
10
96.0 ± 0.24
20
96.8 ± 0.50
10
95.5 ± 0.46
20
96.1 ±0.21
10
93.7 ± 0.51
20
94.7 ± 0.36
Two-way Anova 3
98.8
RBD palm olein
96.4
RBD palm oil
95.8
PBD palm stearin
94.2
LSD (5%)
0.61
0.43
1 Data obtained from 5 pairs of male rats per experimental group Mean for three periods F value for interaction of fat type and fat level is not significant at <*- 0.05 a
Equivalent to 21 energy % b Equivalent to 39 energy %
C
SD
similar food efficiency values but on doubling the dietary fat level to 39 energy %, the food efficiency of the RBD palm olein diet, while not different from either the soybean oil or RBD palm oil diet, was significantly higher than for the RBD palm stearin diet. These findings agree with the report of Barki (1950) that different fats may have different optimum levels of intake and that comparing the- nutritional value of fats based on only one dietary fat level may be inappropriate.
Fat Digestibility The coefficient of digestibility values for each of the fats used were closely similar for the three periods (scond, fourth and sixth weeks) investigated. No apparent trend was observed for the digestibility of a fat in relation to the age of the
animals after an initial orientation period of a week. The mean digestibility values found for these three periods were calculated for each fat and the results are presented in Table 3. Table 3 shows that on the whole, slightly higher digestibility values were obtained with the diets containing the higher level (39 energy %) of fat except for the case of soybean oil. Similar observations were recorded by Hoagland and Snider (1943) in feeding tests with rats using purified diets containing 5% w/w and 15% w/w of fat. Although the coefficient of digestibility values obtained for the individual refined palm oil fractions differed significantly ( a = 0.05), these values nevertheless fall within the range reported for most edible fats and oils as shown in Table 4.
PERTANIKA VOL. 11 NO. 3, 1988
403
T.K.W. NG, H.T. KHOR AND Y.H. CHONG TABLE 4 Digestibility of Malaysian refined palm oil compared with some common edible oils a Coefficient of Fat/oil
digestibility (%)
Soybean oil
98.8
Corn oil
98.3 b
Cottonseed oil
97.0
Coconut oil
96.5 b
RBD palm olein
96.4
RBD palm oil
95.8
RBD palm stearin
94.2
Hydrogenated soybean oil
91.7
Butterfat
90.7 b
a
Values presented are those obtained by the authors unless otherwise stated Data of Hoagland and Snider (1943), using diets containing 15% w/w of fat
TABLE 5 Absorption of the test fats following the administration of 1.1 ml to fasting male rats Wtof Fat
Animal
Body surface Area2
1
(cm2)
(g)
Fat Absorbed 3 % of fat fed Mgper 100 cm 2 Absorbed per hour
Soybean oil
226.0 ± 6.59
324.1 ± 5.27
41.5 ± 5.85
42.7 ± 5.96 )
o K. Palm olein Palm oil t m JO Palm stearin <
227.5 ± 4.17 225.6 ± 2.38 226.0 ± 3.62
325.4 ± 3.37 323.9 ± 2.10 324.1 ± 3.04
38.0 ± 5.24 36.5 ± 4.57 30.2 ± 3.89
38.6 ± 5.37 ) 36.8 ± 4.45 ) 30.5 ± 3.73 )
225.5 227.2 225.5 224.5
4.13 3.85
323.6 ± 3.92 325.4 ± 3.54 323.8 ± 3.24
1.41
322.8 ± 1.03
66.6 61.5 60.0 57.9
34.4 34.4 30.3 29.3
sor
c o
Soybean oil .2 a,
Palm olein
8
Palm oil
X) <
Palm stearin LSD
± ± ± ±
4.50
± ± ± ±
6.88 7.20 8.76 5.89
3,17
(5%)
' Mean ± S D for 8 animals per experimental group Corrected for the quantity of ether-soluble material removable from the GIT of rats after a 48-hour fast but fed no oil Significantly different by two-way ANOVA for
404
oc
= 0.05
PERTANIKA VOL. 11 NO. 3, 1988
± ± ± ±
3.87 3.87 4.57 3.03
) ) ) )
37 1*
31 1*
THE DIGESTION, ABSORPTION & UTILIZATION OF PALM OIL & ITS FRACTIONATION PRODUCTS IN THE RAT
Rate of Fat Absorption The absorption of the test fats following the administration of a known weight of 1.1 ml sample to fasting male rats are shown in Table 5. The absorption rates of the fats over the first 3 hours (mean of 37. 1 mg per 100 cm2 per hour) were consistently higher than the rates for the 6-hour periods (mean of 31.3 mg per 100 cm2 per hour). This was expected and might be explained by the relatively faster rate of absorption of the shorter-chain fatty acids or unsaturated fatty acid components intially and later, the remaining longer-chain saturated fatty acids were not as readily absorbed or utilised (Deuelef al, 1940. From Table 5, it can be seen that the rate of absorption of soybean oil was significantly higher than for RBD palm olein, although their growthpromoting effects were comparable at either 21 energy % or 39 energy % dietary fat level. Thus, the rate of absorption of a fat need not be correlated with its growth-promoting effect or food efficiency, as was earlier suggested by other investigators (Thomasson, 1956; Gottenbos and Vies, 1983). Nevertheless, absorbability of a fat reflects its value as an energy substrate and a source of the fat-soluble vitamins, and is therefore a useful nutritional criterion. Using a fat dosage of about 325 mg per 100 cm2 body surface area, the percentage of fat administered remaining in the GIT after 3 hours and 6 hours, and the corresponding AT SO values are shown in Table 6.
For a particular fat dosage employed, a lower AT 5 0 value represents a faster rate of absorption. Under the present experimental conditions, the AT 5 0 values of the fats studied ranged from 241 minutes for soybean oil to 309 minutes for RBD palm stearin. Using a higher fat dosage of 400 mg per 100 cm2 body surface area, Thomasson (1956) reported AT S 0 values of 359 minutes and 416 minutes for soybean oil and unfractionated palm fat respectively. CONCLUSION
Refined palm oil and its fractionation products, palm olein and palm stearin are easily digested, well absorbed and efficiently utilised for growth. The coefficient of digestibility values obtained for RBD palm oil, RBD palm olein, RBD palm stearin and soybean oil were 95.8%, 96.4%, 94.2% and 98.8%, respectively which all fall within the range reported for most edible fats and oils. The relative rates of absorption found for the three refined palm oil fractions, as measured by the amount of fat absorbed after 3 hours or 6 hours and their Absorption Time so values, were in the order RBD palm olein > RBD palm oil > RBD palm stearin. ACKNOWLEDGEMENTS
This study was supported by a research grant provided by the Palm Oil Research Institute of Malaysia (PORIM).
TABLE 6 values derived from the percentage of fat administered remaining in the GIT after 3 hours and 6 hours
Fat
Percentage of fat administered remaining in the GIT
AT
50
3 hours
6 hours
(minutes)
Soybean oil
58.5
33.4
241
RBD palm olein
62.0
38.5
272
RBD palm oil
63.5
40.0
283
RBD palm stearin
69.8
42.1
309
Time after which 50% of the administered fat had disappeared from the GIT. PERTANIKA VOL. 11 NO. 3, 1988
405
T.K.W. NG, H.T. KHOR AND Y.H. CHONG REFERENCES
LANGWORTHY, CF. (1923): The digestibility of fats. /. Ind. Eng. Chem. 15: 276.
ALFIN-SLATER, R.B. and L AFTERGOOD (1980): Lipids In "Modern Nutrition in Health and Disease", Sixth Edition. Goodhart, R.S. and M.E. Shils(eds.)> Lea and Febiger, Philadelphia.
LEE, M.O. (1929): Determination of the surface area of the white rat with the application to the expression of metabolic results. Am. J. Physiol 89: 24-33.
BARKI, V.H., R.A. COLLINS, CA. ELVEHJEM and E.B. HART (1950): The importance of the dietary level of fats on their nutritional evaluation. / Nutr. 40: 383-392. BLIGH, E.G. and W.J. DYER (1959): A rapid method of total lipid extraction and purification. Canadian J. Biochem. Physiol 37: 911-917. BRACCO, U. and H. BAUER (1978): Human millc lipids and problems related to their replacement In "Lipids and Nutrition*1, Annales Nestle 40, p. 55-81. COLLAWAY, D.H., G.W. KURTZ, J.J. MCMULLEN and L.V. THOMAS (1956): The absorbability of natural and modified oils. Food Research. 21: 621. CROCKETT, M.E. and H.J. Jr. DEUEL (1947): A comparison of the coefficient of digestibility and the rate of absorption of several natural and artificial fats as influenced by melting point. / Nutr. 33: 187-194. DEUEL, H.J. Jr., L. HALLMAN and A. LEONARD (1940): The comparative rate of absorption of some natural fats,/. Nutr. 20: 215-226. FILER, L.J., Jr., F.H. MATTSON and S.J. FOMAN (1969) Triglyceride configuration and fat absorption by the human infant. / Nutr. 99: 293-298. GOTTENBOS, JJ. and R.O. VLES (1983): The nutritive value of palm oil In "Nutrition**, Berger, K.G. (ed.). PORIM Occasional Paper No. 8.
MATTIL, K.F. and J.W. HIGGINS (1945): The relationship of glyceride structure to fat digestibility, I. Synthetic glycerides of stearic and oleic acids. / Nutr. 29: 255-260. MATTSON, F.H. (1959): The absorbability of stearic acid when fed as a simple or mixed triglyceride. J.Nutr. 69: 338-342. MATTSON, F.H., J.H. BENEDICT, J.B. MARTIN and L.W. BECK (1952): Intermediates formed during digestion of triglycerides. / Nutr. 48: 335-344. NG, T.K.W., Y.H. CHONG and H.T. KHOR (1987): Food energy from palm oil and its fractionation products, palm olein and palm stearin. Pertanika. 10(3): 305-309. THOMASSON, H.H. (1956): The biological value of oils and fats. IV. The rate of intestinal absorption. J.Nutr. 59:343-352. TOMARELLI, R.M., B.J. F.W. BERNHART distribution on the of human milk and 583-590.
WELKOWITZ, J., R.B. EWEN and J. COHEN (eds.) Introductory Statistics for the Behavioral Sciences, 2nd Edition. Academic Press, New York, pp 140146, 1976.
HOAGLAND, R. andG.G. SNIDER (1943): Digestibility of some animal and vegetable fats. /. Nutr. 25: 295-302.
406
MEYER, J.R. WEABER and (1968): Effect of positional absorption of the fatty acids infant formulas. /. Nutr, 95:
PERTANIKA VOL. 11 NO. 3, 1988
(Received 10 March, 1988)
Pertanika 11(3), 407-417 (1988)
Effect of Chemical Treatments on the Shelf Life of Rambutans (Nephelium lappaceum )—II SUHAILA MOHAMED, ESHAH OTHMAN and FAUZIAH ABDULLAH Department of Food Science Faculty of Food Science and Biotechnology Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia. Key words: Rambutans; calcium chloride; sodium metabisulphite; benomyl dips and sulfur dioxide treatment. ABSTRAK Mencelup rambutan selama 5 minit dalam larutan sodium metabisulfit, calcium chloride dan metabisulfit atau calcium chloride dan asid askorbik dapat mengurangkan % buah-buah rosak sehingga 30%. Perlakuan sulfur dioxide didapati kurang berkesan untuk mengekalkan kesegaran rambutan bila dibandingkan dengan penyimpanan di bawah udara terubah suai Teknik-teknik yang dapat menukar kadar pernafasan dan kadar kehilangan kelembapan adalah lebih berkesan dalam mengekalkan kesegaran rambutan dari cara-cara yang menghalang keperangan enzim. Buah-buah yang telah dicelup dengan sodium metabisulphite + calcium chloride selama 5-10 min, dalam bungkusan PE pada suhu simpanan 8 C, masih manis dan mantap pada hari yang ke 17. Buah-buah yang disimpan pada suhu bilik tidak menunjukkan perbezaan bererti antara yang telah dicelup dan yang tidak melalui perlakuan kimia. ABSTRACT Dipping rambutans for 5 minutes in sodium metabisulphite, calcium chloride and metabisulphite or calcium chloride and ascorbic acid solutions helped reduce % unacceptable fruits by up to 30% when fruits were stored packaged at 8 C. Chemical dipping of rambutans and sulfur dioxide treatments were found to be less effective in prolonging the shelf life compared to low temperature storage. Techniques which slow down respiration rafe and dehydration rate were found to be more effective in prolonging the shelf life of rambutans than methods to inhibit enzymic browning. Fruits treated with sodium metabisulphite + calcium chloride in PE at 8 C with soaking time of 5-10 minutes, remained sweet and firm on the 17th day. Shelf life of treated fruits stored at room temperature is not significantly different from untreated fruits. INTRODUCTION Chemical control of post harvest wastage has become an integral part of the handling and successful marketing of fruit. A wide range of chemicals has been used for the control of postharvest wastage of fruits. Lieberman and Mapson (1962), Lieberman et alf (1964) and Ayres et al, (1964) reported that treatment on tomatoes with ethylene oxide showed a definite delay in ripening, developed attractive skin colours with firm texture. Whereas usage of potassium permanganate as ethylene absorbent (Scott et aL, 1970) in film bags package
of bananas showed firmer texture than fruits sealed with calcium hydroxide to removed carbon dioxide. Usage of mold fumigant, methyl bromide, gave the maximum inhibition of mango ripening throughout the storage period of 20 days at ambient temperature (Subramanyam etaL, 1969). Benomyl, a fungicide used to control spoilage during storage was observed to delay ripening of Alphonso mango (Shanta Krishnamurthy and Rao, 1983). Sulfur dioxide as a gas or in solution as the acid or salt has been in use for centuries especially in the preservation of dried fruit and
SUHAILA MOHAMED, ESHAH OTHMAN AND FAUZIAH ABDULLAH
wine to maintain their light, natural colour during extended storage. It is also being used for the extension of shelf-life of raw grape and preservation of raw crushed grapes or wine (USDA, 1977). The objective of this research is to determine whether chemical treatments can be used to prolong the shelf life of fresh rambutans. MATERIALS AND METHODS
Rambutans of the red variety (R7) were harvested in bulk from the University Orchards (Ladang 5). They were sorted, weighed and subjected to various treatment and storage conditions on the same day. Each result is an average reading from analysis of 6 fruits. Chemical Treatments The rambutans were dipped for 5, 10, 15 and 20 mins. in the following solutions: i) 1000 ppm sodium metabisulphite. ii) 1000 ppm sodium metabisulphite+300 ppm calcium chloride, iii) 500 ppm benomyl at 50 C. iv) 1000 ppm calcium chloride + 1% ascorbic acid. Results were compared to that of untreated fruits. The chemical treatments were also studied in combination with the following storage conditions: a. Stored unwrapped at ambient temperature.
b. c. d.
Sealed in low density polyethylene bags at ambient temperature. Sealed in low density polyethylene bags at 8 ± 2°C. Packed in nitrogen gas, sealed in low density polyethylene bags at ambient temperature.
The effect of leaving the fruit stalks attached to the fruits, on shelf life of rambutans were observed. Fruit stalks of different lengths (0 cm, 15cm and 30cm) were left attached to the fruits. The colour and texture of the fruits were compared to that of the control (fruits with no stalks). Treatments with Sulfur Dioxide Fruits were exposed to dense sulfur dioxide fumes from burning sulphur in a closed dessicator for 1 min, 5 min, 10 min, 15 min, 120 min and 240 min respectively. Care was taken to ensure that neither the flame nor the hot crucible touched any part of the fruit. Fruits were weighed, packed and sealed in LDPE bags after exposure-. RESULTS AND DISCUSSION
Chemical Treatments The effect of chemical dips on rambutan stored unwrapped at ambient temperatures and in LDPE bags at ambient and 8 ± 2 C are shown in Figs. 1, 2 and J respectively. For fruits kept unwrapped at ambeint temperatures, all the chemical treatments (except with benomyl) reduced the percentage unacceptable fruits by 20-30%. However, the storage LEGEND
Figure 1.
408
Effect of chemical treatments on % unacceptable fruits (unwrapped ambient temp) PERTANJKA VOL. 11 NO. 3, 1988
*
CONTROL
o
NAMET
o
NAMETCACL
*
BENOMYL
EFFECT OF CHEMICAL TREATMENTS ON THE SHELF LIFE OF RAMBUTANS % unacceptable fruits (ambient in LDPE) 100
a benomyl
+
control
0
Figure 2.
doyi NoMetCaCI
A
NoMet
X
CoClAacA
Effect of chemical treatments on % unacceptable fruits (ambient in LDPE).
life could not be extended beyond four days. By the 14th day all the samples have turned black accompanied by an unfavourable fermented smell and softening of the skin and flesh. At this stage discolouration of the flesh had occurred and the fruit was inedible. By the fifth day fungal growth
was present in all except those treated with sodium metabisulphite. The skins was beginning to rot. Rambutans sealed in LDPE, both at 8 C and ambient temperature, have reduced % unacceptable fruits only with sodium metabisulphite,
on % unacceptable fruits (LDPE at 8°C)
1 +
3
NoMetCcC!
19 • Control
Figure 3.
o
days NaMat
A
CaCL AscAcid
21
* benomyl
Effect of chemical treatments on % unacceptable fruits (LDPE at BC). PERTANIKA VOL. II NO 3, M9W
40V
SUHAILA MOHAMED, ESHAH OTHMAN AND FAUZIAH ABDULLAH
calcium chloride + sodium metabisulphite and calcium chloride + ascorbic acid treatments compared to the control (chemically untreated fruits under the same storage condition). Benomyl treatments increased the percentage of unacceptable fruits showing that the degradation is physiological, i.e. not due to microbes or fungus. Hot water treatment at 50°C increased the rate of browning, causing drying of the skin tissue and a faster drying rate. The best chemical treatments appeared to be sodium metabisulphite + calcium chloride which reduced the % unacceptable fruits by 10-20% of the control. Metabisulphite acts both as an enzymic browning inhibitor and preservative. Sulphites acts as a reducing agent and also gradually reduces the effectiveness of the enzyme (Embs &Markakisl965). Calcium acts synergistically with either sulphites or ascorbic acid to prevent browning in apple slices (Ponting etaly 1972). Faust & Klein (1973) and Faust & Shear (1972) have shown an inverse relation between respiration and the Ca concentration in the flesh of fruit. In this experiment the Ca may have done the same to the pericarp. Calcium has been shown to be able to preserve cellular organisation and at deficiency levels, to induce disintegration of cytoplasmic membranes. One function of Ca may be to main-
tain protein synthesis which may supplement its effect in preserving cell membranes as a factor in the preservation of cellular organisation. (Faust & Klein 1973, Faust & Shear, 1972). The chloride present can also inhibit the activity of phenol .oxidase enzymes and will retard development of brown color for a limited time. Poix Ade et alM (1980) showed an exponential relation existing between reaction speed and amount of added chloride. Ascorbic acid is also known to delay browning in fruits, possibly by reducing the oxidized intemediate of phenolic compounds. In addition, it is believed to act on some functional group in the enzyme (Baruah and Swain, 1952). For fruits sealed in LDPE, the differences between treatments with sodium metabisulphite and sodium metabisulphite + calcium chloride is not much at ambient temperature but is quite significant at 8°C. The effect of varying the soaking time for the chemical treatments are seen in Fig. 4. The optimum time appears to be 5-10 minutes and soaking at 20 minutes adversely affects the storage quality of the fruits. The fruits became too moist and decayed faster. A similar trend is seen for all the chemical dipping treatments. By the 17th day, fruits treated with sodium metabisulphite + calcium chloride in LDPE at 8 C, remained sweet and firm at soaking time of 5 to 10 minutes.
on shelf life of rambutans (8°C in LDPE) 100
CD
a
control
+
5 mln Figure 4.
410
$
10 min
A
15 mm
Effect of NaMetCaO soaking time on shelf life of rambutans BC in LDPE). PERTANIKA VOL. 11 NO. 3, 1988
EFFECT OF CHEMICAL TREATMENTS ON THE SHELF LIFE OF RAMBUTANS
Brown patches were observed in the inner layer of the skin for samples soaked at 15 and 20 minutes. Mendoza et al (1972) study on Seematjan and Maharlika rambutans showed that storage decay could be controlled effectively by treatment with 1000 ppm benlate (l-butylcarbonyl-2-benzimidazol carbonic acid) but wax coating reduced moisture loss for 2 days only.
day compared to that of the control at 8 C (wrapped in LDPE), (Fig. 5). For ambeint temperature storage, sulfur dioxide treatment was detrimental to the fruits (Fig. 6). Although sulfur dioxide inhibits enzymic browning it does not slow down the respiration rate, hence the physiological deterioration that occurs in the fruits during storage. Fruits treated with excess sulfur dioxide for 120 minutes and 240 minutes have the lowest shelf life. The fruits
Sulfur Dioxide Treatments Since sodium metabisulphite dips in the presence or absence of calcium ions were able to slightly extend the shelf life of wrapped rambutans at 8°C, a further experiment was carried out using sulfur dioxide gas exposure. Sulfur dioxide gives better penetration into the skin, and like metabisulphite, it behaves both as a preservative, and anti browning agent. When fruits were exposed for different durations, 15 minutes sulfur dioxide exposure extended the shelf by 2 - 3 days but had a higher % unacceptable fruit at anytime until the 16th
unacceptable level
100.
unacceptable level
Days 0
Figure 6. 60
SO -
D.ys 3
Figures.
T
11
15
19
% unacceptable fruits with time stored at l(Pc (SO2 treatments).
1
3
3
7
% unacceptable fruits with time stored at ambient temperature (SO2 treatments).
suffered from excessive bleaching and injury, with the skin turning bright pink and browning occurring on the inner side of the skin. The flesh also suffered injury - had turned slightly brown and gave off a strong acidic, sulfurous odour thus making it inedible. Treatment with sulfur dioxide gas can provide satisfactory quality maintenance for other fruits. However, the limiting parameter is usually the acid-sulfurous odour and taste, and also its corrosivity. It is necessary to promptly and vigorously sweep out excess sulfur dioxide to avoid a sulfurous off-flavour. A recent comprehensive
PERTANIKA VOL. 11 NO. 3, 1988
411
SUHAILA MOHAMED, ESHAH OTHMAN AND FAUZIAH ABDULLAH
review of sulfur dioxide application to food revealed no deleterious effects with the exception of a reduction in thiamine (Post, 1979a). Any condition that accelerates sulfur dioxide loss will in turn accelerate the darkening of the product (Schrader and Thompson, 1947; Bolin and Boyle, 1972). Weight Loss Sulfur dioxide treated fruits showed a slight increase in dehydration compared to untreated fruits sealed in LDPE. Maximum weight loss for sulfur dioxide treated fruits was 0.8% at 8 C and 1.3% at ambient (Fig. 7). Sulfur dioxide may have caused some changes in the cells and decreased the ability of the cells to retain moisture. Ambient temperature
8-c Control
in fruits which underwent longer exposure (120240 mins).
Texture Fruits stored at 8 C, and which had undergone 15 mins of exposure to sulfur dioxide gas, showed firmer skin texture than those of the control fruits at the end of its shelf life (Fig. 8). Fruits exposed to sulfur dioxide for 10 and 15 mins showed significantly firmer skin texture, compered to the control up till the 9th day of storage. Exposure of fruits to sulfur dioxide for 1 min, 120 mins and 240 mins decreased the skin firmness. Too long an exposure to sulfur dioxide may have caused injury to the fruits. Storage at ambient temperature showed a similar trend, but with a shorter storage life of 4 days, hence a steeper slope of decline is obtained. Sulfur dioxide treatments for any duration adversely affect the pulp firmness (Fig. 9).
lmin
X
1.4S mins
•
1.3-
10 mins N •*•
1.2 1.1 -
IS mins
V
i'
120 mins
bw
240 mins
1
$i
1.00.9 _
it'iu »'!' ni ii i
0.8-
nJt
S: mil
0.70.6 H
m
0.5 . 0.40.3 ,
A h ill
0.2 _ o.i -
w^
"9
t><>
1
0
Figure 7.
1
3
5
7
9
11
r
.
12
15
.
•
17
19
V Days •>
% weight loss with time (SO2 treatments).
Colour Sulfur dioxide treatments helped retain the highest L values (lightness) and b values (yellowness) but gave the lowest a values (redness) (Table 1). This showed that sulfur dioxide preserved the colour by a bleaching effect i.e. sulfur dioxide inhibits browning, but is detrimental to the natural red pigment present in the fruit skin. Discolouration to yellowish brown could be seen in fruits which were exposed for short durations (1-15 mins) while a pink discolouration was seen 412
Days 0
Figure 8.
1
3
5
Pulp firmness with time (SO2 treatments).
PERTANIKA VOL. 11 NO. 3, 1988
TABLE 1 Effect of various SO2 treatment on colour of the skin for rambutans stored in LDPE BAGS (0.05 mm thickness)
O
o 1 min
Control Day
5 mins
10 mins
15 mins
120 mins
ffl
240 mins
L
a
b
L
a
b
L
a
b
L
a
b
L
a
b
L
a
b
L
a
b
19.9 18.2 18.5 17.7 16.7 15.9 15.0 13.5
13.0 10.9 10.4 9.0 8.5 7.8 7.0 6.2
6.9 6.4 6.0 5.7 5.4
27.4' 26.9 24.7 21.3 20.0 16.1 16.4
1.6 4.9 6.3 7.5 8.0 7.6 3.9
3.4 5.6 6.6 7.8 7.9 5.9 5.8
27.3 20.7 25.0 20.5 19.9 16.4 15.1 14.3
2.3 3.7 4.6 5.9 6.7 4.6 5.8 4.9
4.1 4.6 5.9 6.4 6.9 5.8 4.9 4.5
25.7 24.5 23.6 22.3 21.7 17.4 16.6 14.5
4.3 5.9 7.7 8.4 3.6 7.4 4.0 4.0
5.4 5.6 6.9 7.5 9.5 6.3 5.7 5.3
26.4 25.7 24.3 23.9 22.6 19.9 17.8 16.7 15.3
1.2 3.9 4.5 5.2 5.3 5.6 6.2 5.4 4.5
2.5 5.2 6.0 6.8 6.9 7.0 6.7 6.3 5,2
28.5 26.1 24.3 21.0 19.8
6.3 S.4 5.3 5.2 4.9
10.0 9.7 9.1 8.0 6.8
29.0 25.6 23.8 24.6 20.4
4.9 3.6 2.1 5.1 4.4
11.8 10.4 10.1 8.0 7.6
28.3 25.0 21.5
1.9 4.8 3.4
3.8 5.6 4.0
24.0 19.3 16.9
S n
(8°C)
fj
1 3 5 7 9 11 15 17 19
(Ambient) 1 20.1 3 18.6 15.5 5
11.9 11.3 9.7
5.2 4.1 3.9
7.0 5.8 3.9
25.4 20.1 18.2
3.7 4.5 3.5
4.1 6.3 5.4
26.1 21.8 17.4
1.7 4.0 3.0
4.0 5.6 3.7
26.2 21.8 18.7
2.1 4.9 2.9
1.5 5.9 4.0
> H
m O
z H
m X
rfl *1
4,0 5.4 3.5
8.6 7.9 7.5
22.9 20.9 18.7
3.7 2.8 2.0
9.8 8.1 6.2
m O
> DO
a *SD values ranges from 0.1 - 0.3
SUHAILA MOHAMED, ESHAH OTHMAN AND FAUZIAH ABDULLAH
11
13
IS
17 18
Ambient temperature
k
8-c Control 1 min 5 mliw 10 mini
v. \\\
ItnfM UOminr 240 mini
V
i 0
Figure 9.
1 1
3
r~> Days 5
Skin firmness with time (SO2 treatments).
Chemical Treatment together with Storage in Nitrogen at Ambient Temperature The result of treating fruits with various dips and then storing them under nitrogen in sealed LDPE bags are shown in Table 2. Storage in nitrogen show some potential in prolonging the shelf life of rambutans treated with calcium and a reducing agent which reduced the respiration and oxidation rate of the fruit. The effect of storing rambutans treated with calcium chloride and Na met under nitrogen at 8 C could very well prolong the shelf life of the fruit further but this was not investigated because it would mean a costly postharvest handling of the fruit. 4J4
Norman and Craft (1971) found that oranges held in nitrogen produced increasing amounts of ethanol, acetaldehyde and methanol as the duration of the anaerobic treatment was prolonged. Volatile emission fell upon transfer of fruits to normal air. Rind injury, however developed in air preceeding an exposure period of 3 or more days in N. The high oxygen uptake or carbon dioxide production as a result of exposure to nitrogen may be reversed if low oxygen levels are used. Craft et al (1968) observed that the anaerobic carbon dioxide output initiated by lemon fruits in the nitrogen atmosphere was suppressed by adding 2 or 5% oxygen. Oranges and lemons were compared to rambutans because they all fall under the category of non-climateric fruits (Mendoza et al., 1972). Effect of Leaving Fruit Stalks Attached to the Fruit Leaving the fruits stalks on the fruit, did not significantly prolong the shelf life of the fruits when compared to the control. (Table 2) This is contrary to the general belief that the presence of fruit stalk can prolong the shelf life of the fresh fruits but is in agreement with the fact that the major moisture loss occured through the soft spines (hairs) of the pericarp (Mendoza et al 1972). Correlations between Weight Loss, Texture and Colour There appears to be a good correlation between skin firmness and L value for all the treatments both at 8 C and ambient temperature (Table 3). However, the correlation of a and b values to texture is poor with sulfur dioxide treatments. There is a good correlation between firmness of the skin and pulp with weight loss in sulfur dioxide treated fruits.
ACKNOWLEDGEMENT The authors thank the Universiti farm personnel for making available the fruits for research (especially Wan Jaharah Wan Ngah and Haji Mahmood Jaafar), Encik Abdullah Hassan (BTM, MARDI) for useful suggestions at the start of the research, and Norhayati Salamuddin for typing the manuscript.
PERTANIKA VOL. 11 NO. 3, 1988
EFFECT OF CHEMICAL TREATMENTS ON THE SHELF LIFE OF RAMBUTANS
TABLE 2 Effect of varying stalk length and storage in Nitrogen on % unacceptable fruits
% Unacceptable trulti Stalk length/days
0
2
10
12
100
U n wrapped/ KT 0
0
40
IS
0
50
90 90
100 100
30
0
50
90
100
LDPL/RT 0
0
20
65
95
100
15
0
20
65
95
100
30
0
20
65
95
100
LDPE/RT Namct + Cacl2 0
0
20
60
90
100
15
0
20
60
90
100
30
0
20
60
90
100
LDPE/RT (Namet + Asc. Acid) 0
0
30
75
90
100
15
0
30
75
90
100
30
0
30
75
90
100
Unwrapped/8 C 0
0
15
30
60
80
90
15
0
15
30
60
80
90
100
30
0
15
30
60
80
90
100
0
0
5
15
20
35
45
50
15
0
5
15
20
35
45
50
0
5
15
20
35
45
50
LDPE/8 C
30 LDPE/8 C (Namct + Cacl2) 0
0
5
15
20
30
40
50
15
0
5
15
20
30
40
55
30
0
5
15
20
30
40
55
5
20
30
35
40
50
5
20
30
35
40
45
5
20
30
35
40
45
90
100
90
100
LDPE/8 C (Namet + Asc. Acid) 0 15 30
0 0 0
Chemically treated fruits stored in
nitr OQ«n/
LDPE
0
33
Na Benz/sorbate
0
37
57 93
100
Hot benomyl
0
63
93
100
NaMet + CaC12
0
27
40
50
73
CaQ2 + Asc. Acid
0
23
60
73
100
PERTANIKA VOL. 11 NO. 3, 1988
415
TABLE 3 Correlation between skin and pulp firmness with weight loss and between skin firmness with colour Correlation Coefficient Skin Texture vs W|:. Loss
TRFATMFNTS N 1.
Best Fitted Line Pulp Texr-ire vs Wt. Loss Sicin Texture vs Wt. Loss
Pulp Texture vs Wt. Loss
CO
SO 2
C
X > >
(8°C) a) Control b) 1 min c) 5 mins d) lOmins c) 15 mins 0 120 mins g) 240 mins
-0.903 -0.969 -0.976 -0.931 -0.972 -0.942 -0.965
16 14 16 16 18 10 10
-0.918 -0.907 -0.967 -0.895 -0.981 -0.961 -0.896
y y y y y y y
= = = = = = =
-23.261 -20.740 -10.310 -20,537 -20.041 -51.517 -60.567
x x x x x x x
+36.502 +38.257 +28.022 +42.746 +41.304 +40.455 +43.261
y y y y y y y
=
= =
= s
-15.137 X - -.968 X - *\5O7 X -15.536 X -11.338 X -;o.O96 X -40.155 X
+30.149 +23.428 +23.986 +30.609 +25.437 +32.156 +34.126
5 > m f—i
m
2. a) b)
c) d)
e)
0
o o
g)
(Ambient) Control I min 5 mins lOmins 15 mins 120 mins 240 mins
TREATMENTS 1. S0 2 (Ambient) a) Control b) 1 min c) 5 mins d) lOmins e) 15 mins 0 120 mins 8) 240 mins
6 6 6 6 6 6 6
-0.943 -0.983 -0.945 -0.955 -0.981 -0.909 -0.948
-0.880 -0.998 -0.918 -9.995 -0.932 -0.945 -0.988
y y y y y y y
Correlation Coefficient Texture vs Colour N
L
a
- 6.097 -17.777 - 6.723 -12.735 -10.890 -11.334 -11.501
x x x x x x x
+32.433 +41.135 +27.594 +39.370 +38.227 +35.436 +33.358
y s y y* ~ y = y = y = y *
- >J96 X +32.070 - T . 2 7 6 X +36.693 - 3.224 X + 19.068 -10.919 X +31.271 - 7.057 X +25.474 -5.972 X +26.839 -10.543 X +31.538
X
o X >
5 * >z
Best Fitted Line Texture vs Colour a
L
b
= = = = = =
m on X >
i
b
a
Tl
••
6
6 6
6 6 6 6
0.919 0.992 0.940 0.997 0.990 0.995 0.997
0.894 0.051 -0.243 -0.439 -0.500 -0.027 0.989
0.944 -0.694 0.480 -0.700 -0.081 0.992 0.976
y y y y y y y
* * = = *
0.997 1.876 0.638 1.513 1.335 1.771 2.708
x x x x x x x
+11.502 -9.189 + 9.666 -3.089 -1.602 -8.558 -31.310
y y y y y y y
2.001 0.686 = -0.622 = - 1.740 * -1.564 - -0.173 6.765
x x x x x x x
+7.568 +27.972 +25.363 +36.283 +36.960 +27.718 +5.934
y y y y y y
* K
* = = *
y =
1.536 4.428 1388 -1.814 -0.373 11.448 3.155
X X X X X X X
+20.956 +53.966 + 17.407 + 37.432 +33.361 -64.614 -0.248
*
>
N >
X >
s r >
X 2.
(8°C) a) Control b) 1 min c) 5 mins d) lOmins e) 15 mins 0 120 mins g) 240 mins
16 12 16 16 18 10 10
0.893 -0.938 0.962 0.970 0.928 0.984 0.985
0.948 0.930 -0.817 0.402 -0.750 0.893 -0.059
0.922 0.994 -0.587 0.075 -0.614 0.943 0.987
y y y y y y y
« * * * * «
1.815 0.850 0.441 1.275 1.017 1.264 1.691
x x x x x x x
-0.422 + 10.642 + 13.749 + 6.445 9.621 -1.449 12.447
1.778 x +13.925 y y = -1,525 x +39.097 y - -1.225 x +29.128 1.117 x +26.617 y y - -2.302 x +42.078 7.808 x -13.505 y y = -0.271 x +29.694
3.716 = 5.710 = * -1.299 0.283 • y • • -1.928 3.219 y • 3*. 137 y =
y y y y
X X X X X X X
+ 10.025 -3.821 +30.412 +31.093 +42.653 + 0.872 -1.445
EFFECT OF CHEMICAL TREATMENTS ON THE SHELF LIFE OF RAMBUTANS puree, Lebensmittel-Wissenschaft und-Technologie 13(2), 105-110.
REFERENCES AYRES. J.C, A.A. KRAFT, and L.C PIERCE. (1964): Delaying spoilage of tomatoes. Food Tech. 18(8): 1210. BARUAH, P. and T. SWAIN. (1925): The effect of 1-ascorbic acid on the in vitro of polyphenoloxidase from potato. Biochem. J. 55: 392-399. BOLIN, H.P and F.P. BOYLE. (1972): Effect of storage and processing on sulfur dioxide in preserved fruits. Food Prod. Dev. 6(7): 82. EMBS, R.J. and P. MARKIKIS. (1965): The mechanism of sulfite inhibition of browning caused by polyphenol oxidase. J. Food Sci 30: 753-758. FAUST, M. and J.D. KLEIN. (1973): Levels and sites of metabolically active Ca in apple fruit. / Am. Soc. Hort Sci. 99: 93-94. FAUST, M. and C.B. SHEAR. (1972): The effect of calcium on respiration of apples. /. Am. Soc. Hort. Sci., 97: 437-439. LIEBERMAN, H. and L.W. MAPSON. (1962): Inhibition of the evolution of ethylene and the ripening of fruit by ethylene oxide. Nature 196 (4855): 660.
PONTING, J., D.R. JACKSON and G. WATERS. (1972): Refrigerated apple slices. Preservative effects of ascorbic acid, calcium and sulfite. /. Food Sci 37: 434-436. POST, R. (1979): Sulfur dioxide kinetics. In quarterly report from University of Maryland. DOE, contract EC77-S-01-5073:13. SCHRADER, A.C. and A.H. THOMPSON. (1947): Factors influencing the keeping quality of dehydrated apples. Proc. Am. Soc. Hortic. Sci 49: 125. SCOTT, K.J., W.B. MCGLASSON and E.A. ROBERTS. (1970): Potassium permanganate as an ethylene absorbent in PE bags to delay ripening of bananas during storage. Austr. J. Exp. Agric. Anim. Husb. 10: 237-240. SHANTAKRISHNA MURTHY G. and K.P. GOPALAKRISHNA RAO. (1983): Post harvest control of spoilage in Mango (Mangifera Indica L.) with hot water and fungicides. J. Fd. Sc. & Tech. India. 20(2): 74-77.
LIEBERMAN, M., S. ASEN, and L.W. MAPSON. (1964): Ethylene oxide an antagonist of ethylene in metabolism. Nature 204 (4960): 756.
SUBRAMANYAM, H., N.V.N. MOORTHY, N.V. SUBHADRA and M. MATHU. (1969): Control of spoilage and inhibition of ripening in 'alfonso' mangoes by fumigation. Trop. Sci 11(2): 120.
MENDOZA, D. BJ. E.B. PANTASTICO, and F.B. JAVIER. (1972): Storage and Handling of rambutan. Phillipines agriculturist 55: 322.
USDA (1977): Freezing grapes for wine. Press release. Western Utiliz. Res. Lab., US Dept. Agric. Albany, Calif.
POIX ADE, M.A. ROVET MAYER and J. PHILIPPON. (1980): Combined action of chlorides and ascorbic acid on inhibition of enzymic browning in apple
(Received 19 June 1987)
PERTANIKA VOL. 11 NO. 3, 1988
417
Pertanika 11(3), 419-426 (1988)
Topographic Anatomy of the Abdomen of the Lesser Mousedeer (Tragulus javanicus) K.C. RICHARDSON, M.K. VIDYADARAN*, N.H. FUZINA** and T.L AZMI* School of Veterinary Studies, Murdoch University, Murdoch, Perth, W.A., 6150 Australia. ^Department of Animal Sciences, Faculty of Veterinary Medicine and Animal Sciences, Unwersiti Pertanian Malaysia, 43400 Serdang, Selangor DE, Malaysia. **Institute of Medical Research, Kuala Lumpur, Malaysia. Key words: Mousedeer; Tragulus Javanicus; anatomy; gastrointestinal tract; stomach; rumen; intestine; kidney. ABSTRAK Suatu huraian, disokong dengan kajian radiografU diberikan ke atas anatomi kasar struktur-struktur dalam abdomen pelanduk /Tragulus javanicus/ Perut yang besarf kompleks dan berkantung adalah sifat yang paling ketara dalam rongga abdomen haiwan inl Perut menduduki sebahagian besar sebelah kiri rongga abdomen dan meluas hingga mengisi sebelah kanan bahagian ventral rongga abdomen. Dengan hal yang demikian usus tertolak ke kuadran kauda dorsum, krania kepada apertur pelvis krania. Hati yang multilobus terletak di sebalah kanan rongga abdomen. Ginjal tidak mempunyai lobus; khutub ginjal kanan menusuk lobus kaudat hati sementara kutub ginjal kiri berletak berhampiran, tetapi kauda ginjal kanan. Limpayang berbentuk segitiga terletak di aspek krania dorsum kantung dorsum rumen.
ABSTRACT A description is given of the gross anatomy supplemented by radiographic studies of the abdominal structures of (1. javanicus/ The large sacculated stomach complex is the dominant feature of the abdomen, it occupies most of the left side and extends across to fill the ventral right side. The intestine is primarily relegated to the dorsal caudal quadrant immediately cranial to the cranial pelvic aperture. The multilobed liver lies entirely on the right. The kidneys are not lobed, with the cranial pole of the right kidney abutting the caudate lobe of the liver and the left kidney lying adjacent but immediately caudal to its fellow. The small triangular spleen lies on the dorsal cranial aspect of the dorsal sac of the rumen.
INTRODUCTION The phylogenetic relationships of the Lesser Mousedeer (Tragulus javanicus) have long been the subject of speculation in the scientific literature (Simpson, 1945; Vaughan, 1978). It has a complex stomach as do most of the Artiodactyla but is characterised by having a rumen, reticulum
and abomasum but no omasum (Bolk et aly 1937; Vidyadaran et al} 1982; Langer, 1988). It is this reported unusual anatomy which has stimulated this detailed morphological study of the abdomen of T. javanicus.
K.C. RICHARDSON, M.K. VIDYADARAN, N.H. FUZINA AND T.I. AZM1
MATERIALS AND METHODS
Animals Five adult T. javanicus born in captivity were donated by the Institute of Medical Research, Kuala Lumpur. These were housed in individual cages T x 2' x T at the Animal House Facilities, Faculty of Veterinary Medicine and Animal Sciences, UPM. They were fed on a dietary mix of Ipomoea leaves, long beans, sweet potato, bananas and commercial rabbit pellets (Gold coin) and provided water ad libitum. The weights of ^aerified animals ranged from 1.63 kg to 1.76 kg. Limited data on intestinal lenghts and weights of organs were available from an additional 10 animals used in an earlier study (Vidyadaran etal, 1982).
recumbency with the femora pulled caudally to prevent their superimposition upon the abdomen. No chemical tranquillizers were used. At each session, left lateral and dorsoventral recumbent radiographs were taken at 5 minutes, 2, 7, 24 and 48 hours following the administration of barium sulphate. Between radiographs, the animals were held in their cages.
Dissection Animals were euthanased by an intraperitoneal barbiturate. Three animals were dissected fresh and two following embalming. One embalmed and one freshly killed mousedeer were dissected from a left abdominal flank approach, the other three were dissected from a right abdominal flank approach. Black and white photographs were taken progressively throughout the dissections. Line drawings of the organ positions and their orientation to each other were noted. Dimensions of organs were measured; see Table 1 for measurements actually recorded. The relationships and the varying lengths of the mesenteries were noted.
B
Radiography Another four animals were radiographed using a Mobicon III condenser discharge,, mobile x-ray apparatus, model MB 101S. Exposure was 4mAs and 44kV for lateral views and 6mAs and 45kV for dorsoventral views. These were all taken at a focal distance of 90 cm. High-speed intensifying screens (Kyokko, 250) were used with Curix film (Agfa Gevaert) which were automatically processed (Protec M45) to record the image. A dose of 20 mis of ultrafine suspension of barium sulphate (Barytgen, Fushimi Pharmaceuticals) was given by stomach tube into the distal oesophagus. Animals were positioned by gentle physical restraint on the x-ray table in both left lateral recumbency and in dorsal 420
Fig. 1: A, diagram of left flank of a mousedeer dissected with its thoracic cage intact: B, diagram of left flank of a mousedeer dissected with a thoracic cage removed where; a, lumbar hypaxial musculature; b, left kidney; c, spleen; d, reticulum; e, abomasum; f, dorsal sac of rumen; g, ventral sac of rumen; h, caudoventral blind sac of rumen; i, apex of caecum; k, loops of small intestine, single arrow is left ruminal longitudinal groove; double arrow is ventral coronary groove. Scale line is 30 mm.
PERTANIKA VOL. II NO. 3, 1988
TOPOGRAPHIC ANATOMY OF THE ABDOMEN OF THE LESSER MOUSEDEER
RESULTS
Anatomy: Left Flank The stomach occupied most of the abdominal cavity immediately beneath the left flank (Fig. 1). From this side the three principal compartments of the stomach could be identified with the large rumen dominating most of the abdominal cavity whilst the smaller reticulum and abomasum lay more cranially adjacent the diaphragm (Fig. lb). The reticulum occupied the most cranial position of the stomach compartments with its curved parietal surface abutting the diaphragm over three intercostal spaces, i.e. ventrally in the 6th and dorsally in the 9th space. The reticulum was roughly the size of a golfball (35 mm) and triangular in shajpe with its ventral border being horizontal (Fig. lb). Its caudoventral extremity extended into the 9th intercostal space. A ruminoreticular groove distinctly delineated the caudal extremity of the reticulum. Ventral to the reticulum and slightly more caudal to it lay the fundic extremity of the abomasum. This was firmly adherent to the adjacent cranial protion of the ventral sac of the rumen. The rumen consisted of a distinct dorsal sac directly overlying the ventral sac which led further caudally to a large caudoventral blind sac (Figs, la and b). No caudodorsal blind sac could be found. The rumen ran from the 9th intercostal space caudally to the level of the last lumbar vertebra (L7) or the first sacral vertebra. Its dorsal sac while sometimes extending towards the last lumbar vertebral levels more often terminated in the midlumbar region. The dorsal sac lay primarily on the left side with a little extending past the midline onto the right side. A smal! triangular spleen lay dorsally at the junction of the reticulum and dorsal sac of the rumen. It was held in place by a short gastrosplenic ligament. The spleen had a central hilus. The most cranial portion of the ventral sac usually lay in the 9th intercostal space. Its ventral border lay on the floor of the abdomen and ran caudally to virtually enter the pelvic cavity. Surprisingly little fat occupied the left longitudinal groove between the dorsal and ventral ruminal sacs or in the ventral coronary groove between the caudoventral blind sac and the
ventral rumen sac. Depending on the level of stomach fill and the extent of gas in the-stomach, the ventral sac and the caudoventral blind sac usually lay contiguously in a straight line. However, sometimes, the caudoventral blind sac lay above the ventral sac. Taking origin from the left longitudinal groove and extending ventrally and caudally was the greater omentum which covered the ventral sac and caudoventral blind sac before running over
Fig. 2: A, diagram of right flank of a mousedeer with its thoracic cage intact. B, diagram of right flank of a mousedeer dissected with the thoracic cage removed where a, lumbar hypaxial musculature; b, right kidney; c, liver; d; ventral sac of rumen; e, caudoventral blind sac; f, descending duodenum; g, ascending duodenum; h, descending colon, single arrow is ventral coronary groove. Scale line is 30 mm.
PERTANIKA VOL. 11 NO. 3, 1988
421
K.C RICHARDSON, M.K. ViDYADARAN, N.H. FUZINA AND T.I. AZMI TABLE 1 Measurement taken of the mouse deer T. javanicus Mean
S.E.
15
1585.13
53.04
Length (mm) Height (mm) Width (mm) Weight (g)
8 8 8 4
34.38 11.00 12.88 4.00
0.42 0.33 0.30 0.41
Reticulum Length (mm) Height (mm)
4 4
35.50 42.75
2.47 3.30
Rumen Dorsal Sac Length (mm) Height (mm)
4 4
96.25 44.00
6.88 2.94
Rumen Ventral Sac Length (mm) Height (mm)
4 4
83.75 52.00
2.39 4.32
Rumen Caudoventral Blind Sac Length (mm) Height
4 4
60.00 53.75
0.82 4.27
4 4
66.25 23.75
1.75 2.39
13
176.38 5.75
7.48 0.25
65.00
2.04
47.50
1.44
77.62 20.00
3.75 1.70
69.69 8.75
3.73 1.11
9.25
0.48
Weight (g) Kidney
Abomasum Length (mm) Height (mm) Small Intestine Length (cm) Diameter (mm) Duodenum Descending Length (mm)
4
Duodenum Ascending Length (mm)
4
Caecum
4
Lenth (mm) Diameter (mm) Large Intestine Length (cm) Diameter (mm)
13 5 13 4
Ascending Colon (Proximal) Length (cm) Spiral Colon Length (cm) Diameter (mm)
4 4
27.50 5.50
0.9574 0.29
Descending Colon Length (cm) Diameter (mm)
4 4
16.75 4.50
0.75 0.29
422
PERTANIKA VOL. 11 NO. 3, 1988
TOPOGRAPHIC ANATOMY OF THE ABDOMEN OF THE LESSER MOUSEDEER
the ventral abdominal floor and up the right flank separating the duodenum from the remainder of the intestine (Figs. 1b and 2b). The caecum was elongate, cylindrical and without any longitudinal muscle bands (taenia) or sacculations (haustra). Its base was on the right side but the body extended across to the left side so the bulk of the caecum could be seen usually from the left {Figs, la, b). Also occupying the dorsal left were many coils of small intestine. Abutting the hypaxial lumbar muscles in the same quadrant was the left kidney. This was oval shape with no indication of lobulation. It measured about 35mm long, 10.5 mm high and 13 mm wide. See Table 1. Little or no perirenal fat was visible.
a loose mesoduodenum in which lay the distinctly lobulated pancreas. Jejunal coils could be seen through the greater omental sheet lying haphazardly medial to the duodenum (Fig. 2a). This right lateral sheet of greater omentum ran dorsally to insert into the hypaxial muscle tendon origins medial to the right kidney. Once the greater omentum had been removed, the long tangle of jejunal coils could be seen (Fig, 2b). These were highly mobile and were held by an elongate mesojejunoileum. The terminal element of these coils, the ileum, was conspicuous where it entered the colon. A short triangular mesentery, the ileocaecal mesentery, joined the lesser curvature of the caecum to the ileum.
Anatomy: Right Flank From this approach the ventral rumen and the liver were the most obvious features. The liver lay completely on the right side and could be seen lying with it§ parietal surface closely following the concavity of the diaphragm. The visceral surface was distinctly fissured thus delineating its various lobes. The caudodorsal portion of the caudate lobe had a definite concavity in which lay the cranial pole of the right kidney. As in other mammals, the bile duct, hepatic portal vein and hepatic artery all traversed the hepatic porta. The ventral half of the abdomen was occupied mostly by the extensions of the ventral sac and caudoventral blind sacs of the rumen (Fig. 2). The abomasum lay across the axis of the body with its body lying against the ventral diaphragm and below the right lateral lobes of the liver. The pylorus was directed caudodorsally to the descending duodenum. A short hepatoduodenal ligament anchored this portion to the hepatic porta. Running from the porta was a short bile duct which passed into the descending duodenum 3—4 cm from the pylorus. The descending duodenum ran dorsally and obliquely to the midlumbar region where it turned at the cranial duodenal flexure (Fig. 2) to become the ascending duodenum. This ran forwards adjacent to the hypaxial muscles then ventral to and obliquely across the right kidney to become the jejunum. The ascending duodenum was tightly bound to the proximal descending colon by a 1-2 mm long duodenocolic ligament. Both arms of the duodenum were linked by
Anatomy: Mid Abdomen The proximal portion of the large intestine was not readily observable from either flank. The caecum, which ran from right to left, had most of its bulk on the left, and was contiguous with the ascending colon. The proximal portion of the ascending colon was simple, non-taeniated, nonhaustrated, short and wide. After 4 - 5 cm it was twisted into a loose spiral which had 2.5 centrifugal and 2.5 centripetal coils. These were constant in diameter. The caecum, ascending colon and spiral colon were all loosely held by mesenteries which allowed them much freedom of movement. These elements normally lay in the dorsal caudal quadrant of the abdomen but occasionally strayed forwards to lie beneath the thoracic cage. The last centrifugal coil of the spiral colon ran into a few simple loops then became the transverse colon and ultimately the proximal descending colon. The descending colon was held in place by mesocolon which was 3 cm long proximally and tapered down to 3 mm within the pelvic canal. The descending colon could be seen from the right side, medial to the cranial duodenal flexure (Fig. 2b). Radiography The radiographic studies confirmed and clarified the general topographic anatomy of the gastrointestinal tract. In the normal mousedeer, without the administration of radiographic contrast media, the stomach and intestine could not be distinguished readily (Fig. 3). Occasionally gas in the
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Fig. 3: A, plain radiograph of the mousedeer abdomen, left lateral view; B, line drawing of 'A' where; a, diaphramatic silhouette; b, lumbar vertebra; c, gas in rumen; d, sacrum; e, pelvis; f, femur; g, abdominal floor; h, craniodorsal quadrant; it cranioven tral quadran t; kf caudodorsal quadrant; 1, caudoventral quadrant.
Fig. 4: A, radiograph of the mousedeer abdomen, left lateral view, 5 minutes after the administration of the contrast agent; B, line drawing of fA' where; a, reticulum; b, dorsal sac of the rumen; c, left kidney; d, ventral sac of the rumen; e, caudoventral blind sac.
stomach allowed the ruminal pillars to be seen but overall this was of little use to interpreting structure. However the contrast media studies allowed an overall picture of the form of the alimentary tract to be gradually built up from the timed radiographic series (Figs. 4, 5 and 6). The reticulum appeared triangular on lateral view and rhomboid or oval in the dorsoventral view. A generalised mottling with islands of contrast media up to 2 mm in diameter, was seen in some very early radiographs of some series. Usually the reticulum was flooded with contrast agent which precluded a detailed analysis of its internal architecture. The rumen occupied most of the abdominal cavity, it being responsible for the displacement of the liver to the right and of the intestine to the right craniodorsal and to both the left and the right caudodorsal portions of the abdominal
cavity. The dorsal sac of the rumen was principally on the left side (Fig. 5) and ran from the 8th— 10th intercostal spaces, caudally to the level of the 5th or 6th lumbar vertebrae (Fig. 4). The dorsal sac lay above the much larger ventral compartment which ran, in its extreme cases, from the 11th intercostal space through to the 3rd sacral vertebral level. The ventral sac usually extended from T13 to L6 and the caudoventral blind sac from L5 to S3 (Figs. 4, 5). The distinct outline of the longitudinal and caudoventral pillars mirroring the ruminal and caudoventral coronary grooves respectively were noted (Figs. 4, 5). These varied a little in their position and were at times difficult to identify. The contrast agent in the rumen usually prevented a clear view of the pylorus. However occasionally it could be seen immediately caudal and medial to the right lateral lobe of the liver.
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Small intestine loops were seen lying usually in the caudodorsal quadrant of the abdomen. These were highly coiled and often extended caudally into the entrance to the pelvis. The caecum was readily seen having its base to the right of the abdomen and its apex across to the left side. Its cylindrical nature was obvious (Fig. 6). The spiral colon was usually difficult to see because it was masked by contrast agent elsewhere in the intestine. The descending colon was an obvious dorsal feature to the right midline of the abdomen. Masses of faecal pellets could be seen in the distal descending colon and even more in the rectum. DISCUSSION
A, radiograph of the mousedeer abdomen, dorsoventral view, 5 minutes after the administration of the contrast agent; B, line drawing of 'A' where; a, reticulum; b, dorsal sac of the rumen; c, ventral sac of the rumen; d, caudoventral blind sac.
Fig. 6;
A, radiograph of the mousedeer abdomen, left lateral view, 21 hours after the administration of the contrast agent; B, line drawing of 'A' where; a, caecum; b, spiral colon; c, descending colon; d, rectum.
In the true ruminant both the reticulum and omasum are responsible for particle separation (Hungate 1966; Becker et alt 1963). Yet the absence of a functional omasum in the mousedeer, as reported by Vidyadaran (1982), was confirmed by this study thus posing the question of which structures are critical for the sorting of digesta particulate matter. If, as has been suggested by Simpson (1945) and Langer (1988) the Tragulidae are truly a primitive Artiodactyl, then what is the significance of an omasum? It is well developed in the Giraffidae and Bovinae but not so in the Cervidae and Antilopinae (Moir, 1968). The importance of the presence and absence of the omasum in the order Artiodactyla is still a puzzle. The absence of an omasum results in the cardia and reticuloabomasal orifices being close to one another. This in turn has resulted in the abomasum being permanently held close to the reticulum with the abomasal fundus lying to the left of the midline. The reticulum is constant in its position, it being held against the left dorsal region of the diaphragm. The abomasal fundus lies fixed ventral to the reticulum and the abomasal body is twisted to the right so that ultimately the pylorus lies medial and caudal to the right lateral lobes of the liver. The topographic relationships and mobility of the small intestine segments are similar to those found in domesticated small ruminants (Nickel et aly 1973). The caecum is large without taenia or haustra. Together with the distal small intestine, ascending colon and spiral colon, the caecum is extremely mobile. This complex, which is held closely together by many mesenteric attachments, usually lies in the caudodorsal abdomen but may
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occasionally swing forward to lie close to the diaphragm. Of the other abdominal organs, the kidneys and liver are noteworthy. The liver is displaced to the right presumably by the large stomach complex on the left. However it is still multilobed and has not been simplified in form as is seen in the domestic ruminants (Nickel et al, 1973). On the other hand the kidneys are simple bean-shaped and not lobed as stated by Simpson (1984) which is the normal situation for small ruminants (Nickel et al> 1973). The left kidney has long loose mesenteric attachments which allow it to be displaced probably to allow extensive distension of the rumen. However, of all the differing and unusual morphological features possessed by the Tragulidae, it is the absence of the oamsum which most readily leads to important ongoing studies. The use of either radioisotopes or radiopaque makers are needed to determine the fluid and particulate matter dynamics of the mousedeer's alimentary tract. These techniques could determine where and how particulate matter is sorted in these animals. It is possible that the absence of an omasum may simplify digesta transport and could lead to the efficient conversion of food into body tissue. Consequently nutritional studies of the semi-domesticated Lesser Mousedeer would be extremely important in assessing whether or not these may be a potential source for meat production.
Department of Clinical Sciences, Faculty of Veterinary Medicine and Animal Sciences, Universiti Pertanian Malaysia for their helpful cooperation. REFERENCES BECKER, R.B., S.P. MARSHALL and P.T. ARNOLD. (1963): Anatomy development and functions of the bovine omasum. / Dairy Set 46: 836-839. BOLK, L., E., GOPPERT, E. KALLIUS, and W. LUBOSCH. (1937): Handbuch der Vergleichenden Anatomime der Wirbeltiere. Berlin and Wien: Urban and Schwarzenberg. HUNGATE, R.E. (1966): The rumen and rts microbe*. New York: Academic Press. LANGER, P. (1988): The Mammalian Stomach, Stuttgart: Gustav Fisher.
Herbivore
MOIR, R.J. (1968): Handbook of Physiology. Am Phyt. Soc. 5: 2673-2694. NICKEL, R., A. SCHUMMER and E. SEIFERLE. (1973): Viscera of the Domestic Mammals. Berlin: Verlag Paul Parey. SIMPSON, CD. (1984): Artiodactyls in "Orders and Families of Recent Mammals of the World*', eds, S. Anderson and J. Knox Kames. New York: J. Wiley and Sons. SIMPSON, G.G. (1945): The principles of classification of mammals. Bull Amer, Mus. Nat Hist. 85: 1-350. VAUGHAN, T.A. (1978): Mammalogy. 2nd. ed. Philadelphia: W.B. SaundersCo. VIDYADARAN, M.K., M. HILMI and R.A. SIRIMANF (1982): The gross morphology of the stomach of the Malaysian Lesser Mousedeer {Traguhis javanicus). Pertanika 5(1): 34-38.
ACKNOWLEDGEMENTS
The authors wish to thank Encik K. Palaniandy, Encik H. Hamdy and Encik R. Sivasoorian for their technical assistance. We wish to thank the
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(Received 2 June, 1988)
Pertanika 11(3), 427-435 (1988)
The Radiographic Anatomy of the Gastrointestinal Tract of the Lesser Mousedeer (Tragulus javanicus) K.C. RICHARDSON, M.K. VIDYADARAN*, T.L AZMI* and N.H. FUZINA** School of Veterinary Studies, Murdoch University, Murdoch, Perth, W.A., 6150, Australia. ^Department of Animal Sciences, Faculty of Veterinary Medicine and Animal Sciences, Universiti Pertanian Malaysia, 43400 Serdang, SelangorDE, Malaysia. **In$titute of Medical Research, Kuala Lumpur, Malaysia. Key words: Mousedeer; Tragulus javanicus; anatomy; gastrointestinal tract; rumen; reticulum; intestine; passage rate; barium sulphate; radiopaque markers; contrast agent. ABSTRAK Anatomi radiografi pelanduk (Tragulus javanicus) adalah dihuraikan. Pelanduk mempunyai perut yang sangat besar dan kompleks yang mengisi sebelah kiridan ventral kanan abdomen. Omasum dan pundi tuli kaudodorsum tidak wufud. Retikulum biasanya terletak kranio-ventral di sebelah kiri menyusuk diafragma. Fundus abomasum terletak kaudal dan ventral kepada retikulum. Kadar pergerakan bahan radio legap di sepanfang salur alimentari juga dihuraikan. Tinja mula diketahui membentuk dalam kolon ber~ lingkar. Partikel kecil dan cecair mengalir dengan cepat melalui salur alimentari (x » 44.5 jam). Partikel yang mempunyai densiti hampir sama dengan jerami tertahan dalam rumen sementara logam berat tertahan dalam retikulum.
ABSTRACT «
The radiographic anatomy of the Lesser Mousedeer (Tragulus javanicus) is described. The mousedeer has an extremely large stomach complex which fills the left and the ventral right of the abdomen. No omasum or caudodorsal blind sac were found. The reticulum usually lay cranioventrally on the left side abutting the diaphragm. The abomasal fundus lay caudal and ventral to the reticulum. The rate of passage of radiopaque substances along the alimentary tract is given. Faeces was first noted to form in the spiral colon. Small particles and fluid passed rapidly through the alimentary tract /x = 44.5 hr). Particles with densities approaching that of straw remained in the rumen whilst heavy metals were retained in the reticulum. INTRODUCTION Little is known of the form and function of the digestive system of the Lesser Mousedeer {Tragulus javanicus). It is known that they may eat a variety of foods whilst browsing near or rooting in the forest floor. Leaves and particularly fruit are
important in their diet (Medway 1977, 1978). In captivity, they eat a wide variety of food such as bananas, carrots, sweet potato, mixed grain hulled oats, commercial monkey pellets and alfalfa hay (Medway 1977, 1978) as well as Ipomoea leaves, long beans and peanuts (Morat
K.C. RICHARDSON, M.K. VIDYADARAN, T.I. AZMI AND N.H. FUZINA
and Nordin 1978, Nordin 1978). However details of their actual natural diet are lacking. It may be surmised that this animal, the smallest ruminant and smallest ungulate, need a high quality diet if microbial fermentation is to adequately supply its nutritional needs. Vidyadaran et al (1982) in a general anatomical study of the mousedeer stomach reports the absence of an omasum. Morat and Nordin (1978) using marker studies determined that 95% excretion time were 107.5 ± 2.6h (mean ± SE) for sorghum grain and 54.9 ± 5.72h (mean ± SE) for chromium sesqui-oxide. This study is to determine the radiographic anatomy of the alimentary tract of the T. javanicus, to ascertain the rate of passage of different radiopaque markers as well as to identify anatomical sites where markers may be retained. MATERIALS AND METHODS
Animals Eleven adult female Tragulus javanicus, born in captivity, were borrowed from the Institute of Medical Research, Kuala Lumpur. They were housed either singly or in groups of three in small cages 2' x 2' x 2' at the Animal House Facilities, Faculty of Veterinary Medicine and Animal Sciences, Universiti Pertanian Malaysia. The diet consisted of Ipomoea leaves, long beans, sweet potato, bananas and commercial rabbit pellets (Gold Coin) with ad libitum access to water. The weight of the animals ranged from 1.4 kg to 1.76 kg.
the distal oesophagus. The animals were positioned by gentle physical restraint on the x-ray table in both left lateral and donsoventral recumbency with the femora pulled caudally to prevent their superimposition upon the abdomen. No chemical tranquillizers were used. Each radiograph series consisted of left lateral and dorsoventral recumbent radiographs taken at 5, 15, 30 minutes then 1, 2, 8, 24, 30 and 48 hours following the administration of the contrast agent. When needed, additional radiographs were taken at intermediate times. Between radiographs, the animals were held in their cages. RESULTS
In the absence of barium sulphate or similar radiopaque agenf the alimentary tract structure could not be determined in the live animal. Occasionally gas within the viscus allowed a few intestinal loops or a segment of rumen to be seen. Overall the abdomen of the normal mousedeer had an amorphous granular appearance (Figs. 1 and 2).
Radiography Radiographs were taken using a Mobicon HI condenser discharge, mobile x-ray apparatus, model MB101S. The exposure used was 4mAs and 44kV for lateral views and 6mAs and 45 kV for dorsoventral views at a focal distance of 90 cm. The image was recorded on Curix film (Agfa Gevaert) in combination with High-speed intensifying screens (Kyokko, 250). The radiographs were automatically processed (Protech M45). Plain radiographs were taken. Then either a dose of 20 mis of ultrafine suspension of barium sulphate (Barygen Fishimi Pharmaceuticals) or fifty l-2mm radiopaque pellets either light weight as in straw or heavy metal were given by a stomach tube into 428
'A' photograph and 'B' line drawing of a radiograph of a mousedeer in lateral recumbency where a, diaphragmatic silhouette; bf ribs; c, lumbar vertebra; d, ilium; e, sacrum; f, femur; gf gas in rumen; h, sternum; i, floor of abdomen.
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extending caudally for up to 4 vertebral body lengths, well beyond the last rib (Fig, 3). In the early lateral radiographs contrast agent was absent from the caudodorsal quadrant (Fig. 4) in which the loops of intestine usually lay (Richardson etal, in press).
Fig. 3:
Fig. 2:
'A' photograph and 'B' line drawing of a radiograph of a mousedeer in dorsoventral recumbency where a, diaphragmatic silhoutte; b, thoracic vertebra; c, ribs; d, lumbar vertebra; e, region occupied by liver and right kidney; f, gas in rumen; g, sacrum; h, ilium; i, femur; k, tibia.
Contrast Studies - Anatomy Barium sulphate delineated the form of the stomach and intestine of the healthy mousedeer. In its normal state the stomach complex occupied virtually the entire left of the abdomen and much of the right (Fig. 3). Early dorsoventral views show the liver to be lying entirely on the right side abutting the concavity of the diaphragm and
'A' photograph and B* line drawing of a radiograph of a mousedeer in lateral recumbency 10 minutes following a barium sulphate drench. Here a, reticulum; b, fundus of abomasum; c, dorsal sac of the rumen; d, ventral sac of the rumen; e, caudoventral blind sac; f left kidney.
When most clearly seen, the reticulum appeared to have a regular mottled appearance, probably where the barium sulphate had highlighted the depths and walls of the reticular cells. The reticulum appeared to have a pentagonal or rhomboidal shape in dorsoventral views (Fig. 5) and oval or triangular in lateral views (Fig. 4). Often the reticulum was found abutting the diaphragm dorsally, running between the seventh and eleventh intercostal spaces. In some animals, it was more ventral and cranial in position and ran between the sixth and seventh intercostal spaces. It was only in the earlier radiographs that the vertically or near vertically oriented reticuloruminal orifice could be seen. In later radiographs the reticulum and the dorsal sac of the rumen appear as one.
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Fig. 4: 'A' photograph and 'B' line drawing of a radiograph of a mousedeer in lateral recumbency 3 hours following a barium sulphate drench. Here a, reticulum; b, fundus of abomasum; c, dorsal sac of rumen; d, ventral sac of rumen; e, caudoventral blind sac; f, small intestine coils.
The rumen lay on both sides of the abdomen. A dorsal sac, ventral sac and caudoventral blind sac of the rumen could be identified but no caudodorsal blind sac was present (Figs. 3, 4, 5). The dorsal sac was confined to the left side where it lay with its dorsal wall against the lumbar and thoracic hypaxial musculature from T10 to L5. At least one third and more likely half of its bulk lay below the thoracic cage. The dorsal sac extended cranioventrally across the caudal aspect of the diaphragm at intercostal spaces 10 to 8 where it became confluent with the reticulum. The reticular groove joining the cardia to the reticuloabomasal orifice was never visible. Caudally the dorsal sac ended as a cranially directed concavity level with lumbar vertebra 5-6. The orifice between the dorsal sac and ventral sac could not be seen easily, nor could the horizontal pillars which delineated the orifice. The ventral sac of the rumen lay on the left floor of the abdomen and extended across to the right side. It usually extended between T13 and .430
Fig. 5: 'A' photograph and 'B' line drawing of a radiograph of a mousedeer in dorsoventral recumbency 10 minutes after a barium sulphate drench. Here a, reticulum; b, dorsal sac of rumen; c, caudoventral blind sac.
L6 but occasionally it stretched from T i l to S3 (Fig. 3). Both its cranial and caudal borders curved obliquely upwards. A distinct outline of the pillar associated with the caudoventral coronary groove was visible. This pillar was oriented obliquely upwards from the ventral abdominal floor in the vicinity of L5 dorsally and slightly caudally to L7. Sometimes the pillar lay vertically.
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approximately at L4 and the right was at L5. Parts of the abomasum were visible on both sides of the abdomen. From a left lateral radiograph, its fundic portion could be seen lying caudoventrally to the reticulum and cranial to the ventral sac of the rumen (Figs. 3 and 4). From dorsoventral views the pylorus could be seen, occasionally, on the right side adjacent the hepatic porta (Fig. 6). Only the initial one to two centimeters of the duodenum could be seen regularly. The bulk of the jejunum usually lay as a coiled mass in the caudodorsal quadrant of the abdomen (Fig. 6). Often a few of these coils extended caudally to the pelvic entrance. Occasionally the jejunal coils were seen sandwiched between the abomasum and reticulum or were seen lying adjacent the hepatic porta.
Fig. 7: Fig. 6:
'A' photograph and 'B' line drawing of a radiograph of a mousedeer in dorsoventral recumbency 2.5 hours after a barium sulphate drench. Here a, dorsal sac of rumen; b, pylorus; c, caudoventral blind sac overlain by small intestine coils.
The caudoventral blind sac extended from where the ventral sac ended and ran along the abdominal floor to virtually enter the pelvic cavity. This sac ran from L5 to S3. It was cone-shaped with its greatest circumference being at its junction with ventral sac. The left side of this junction was
'A' photograph and *B' line drawing of a radiograph of a mousedeer in lateral recumbency 21 hrs following a barium sulphate drench. Here a, caecum; b, spiral colon; c, descending colon with faecal pellets; d, rectum with faecal pellets.
The large intestine was radiographically more complex than the small intestine. The caecum usually lay with its base to the right mid dorsal flank, its body in the mid abdomen and its apex to the left flank. Sometimes the caecum ran caudally, remaining on the right side whilst on other occasions the body of the caecum lay in the cranioventral sector of the abdomen.
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The caecum was continuous with the short straight segment of ascending colon which after about 90 cm became coiled upon itself as the spiral colon (Fig. 7). The spiral colon lay in the dorsal right of the abdomen between L2 and L6. Details of the spiral and subsequent irregularly arranged colonic loops could not be determined. However, it was in the spiral colon that elongate faecal pellets of low radiopacity were seen first. Further along the colon especially in the descending colon, the pellets were shorter and had a greater radiodensity. The descending colon was a dorsal midline structure which had a wide diameter and regularly accomodated 100-200 faecah pellets (Fig. 7). These pellets were clumped as large masses not as individual units. One animal was found to be in an advanced state of pregnancy. In fact she gave birth to a healthy offspring during the radiographic series. In this animal the foetus lay with its spine lying against its mother's abdominal floor. The uterus with foetus, occupied the caudal half of the abdomen. The stomach complex plus intestine were pushed forward (Fig. 8). The caudoventral blind
Fig, 9:
'A* photograph and 'B' line drawing of a radiograph of a pregnant mousedeer in lateral recumbency 24 hours after a barium sulphate drench. Here a, caecum; b, descending colon; c, stomach and liver; d, skull of foetus; e, spinal cord of foetus; f long bones of foetus.
sac still extended along the right side as far as L6 but the dorsal sac and ventral sac ran as far as L3 and L4 respectively. The intestine was crammed forward, with the caecum and spiral colon lying vertically in the centre of the abdomen. The descending colon was not pushed out of position {Fig. 9). Contrast Studies - digesta transit
VT photograph and 'B' line drawing of a pregnant mousedeer in lateral recumbency 5 minutes after a barium sulphate drench. Here a, reticulum; b, dorsal sac of rumen; c, ventral sac; d, caudoventral blind sac; e, skull of foetus; f long bones of foetus. 432
Barium sulphate administered by a gastric lavage tube into the distal oesophagus gave an estimate of the movement of the fluid/small particle phase of digesta along the alimentary tract. Radiopaque pellets administered similarily, dispersed through the rumen and reticulum. The light weight pellets remained spread throughout the rumen for about 48 hours (Fig. 10). During this time, surprisingly, the heavy metal impregnated plastic gradually dissolved. By 72 hours the pellets in their original form did not exist, rather they had collapsed to tiny fragments (0.1 mm) which initially collected in the reticulum (Fig. 12) and were then passed through the abomasum and ultimately eliminated from the body.
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The heavy pellets were swallowed and dispersed throughout the rumen and the reticulum (Fig. 11) but within 3 hours these were all collected into the reticulum (Fig. 12) where most remained in excess of 10 days. After 4 - 5 days, 1-2 of these pellets passed through the reticuloabomasal orifice and then along the alimentary tract and were eliminated in the faeces. It is estimated that in excess of 30 days would be necessary for the complete elimination of the 50 heavy pellets. The times of first entry of barium sulphate or radiopaque pellets into the small and large intestine is given in Table 1. The time it took for the stomach, small intestine and the entire alimentary tract to be devoid of barium sulphate or radiopaque pellets is given in Table 1, as an estimate of the time at which faecal pellets, incorporating contrast agent, were first formed. DISCUSSION
This study reports the normal radiographic anatomy of the mousedeers gastrointestinal tract. As in other ruminants the stomach complex is the dominant feature of the abdomen. In the mousedeer, it occupies virtually the entire left side of the abdomen as well as fills the ventral right side. The ventral sac of the rumen with its continuation,
the caudoventral blind sac, is the largest chamber, occupying most of the floor of the abdomen. In fact the caudoventral blind sac occasionally extends into the pelvic cavity. The caudoventral coronary grooves with their associated pillars are constant features being routinely visible in the caudoventral abdominal region. No caudodorsal blind sac was seen. The dorsal sac appears to be restricted to the left side and has no special features. The longitudinal pillars delineating the orifice between the dorsal and ventral sacs were seen occasionally but were not seen clearly or routinely. The reticulum was a little more mobile than initially expected. It was found lying most often cranioventrally adjacent to the left concavity of the diaphragm but occasionally it lay against the diaphragm but in the craniodorsal quadrant. No reticular groove was seen radiographically at any stage yet upon dissection it was present (Vidyadaran et al, 1982, Langer, 1988). Heavy particles were held for a long period in the reticulum. How and why these particles were ultimately passed into the abomasum is not known. The gross radiographic anatomy of the mousedeer's small intestine is unremarkable. The passage of digesta with enclosed radiopaque markers is rapid and would need fluoroscopic
TABLE 1 Passage time of barium sulphate and radiopaque pellets through the alimentary tract.
Mean Contrast agent first through pylorus (h)
1.3
Barium Sulphate Suspension (n=6) Range
0.3-
2.0
Radiopaque Pellets (Heavy)
4 days
Stomach empty of contrast agent (h)
16.5
1 2 . 0 - 22.0
30 + days
Small intestine empty of contrast agent (h)
20.8
18.0 - 24.0
30 + days
Contrast agent first into large intestine (h)
7.0
6.0-
9.0
4.5 days
Faecal pellets first formed (h)
7.2
5.5-
9.3
4.6 days
3 6 . 0 - 48.0
30 + days
Alimentary tract devoid of contrast agent (h)
44.5
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Fig. 10: A photography of a radiograph of a mousedeer in lateral recumbency 4 hours after the administration of light weight radiopaque pellets. Here the pellets are dispersed throughout the rumen and reticulum.
Fig. 11: A photograph of a radiograph of a mousedeer in a lateral recumbency 5 minutes after the administration of heavy radiopaque pellets. The pellets lie in the rumen and reticulum.
Fig. 12: A photograph of a radiograph of a mousedeer in lateral recumbency 6 hours after the administration of heavy radiopaque pellets. The pellets all lie in the reticulum.
techniques to clarify. Like the Tammar wallaby (Richardson and Wyburn, 1980) the mousedeer has a simple small caecum which is probably only of secondary importance in digestion. Again, as in studies on kangaroos the caecum probably does not retain preferentially, either the liquid or the 434
particulate phase of the diet (Hume, 1982). Whilst the present study does show that the final clearance of barium sulphate from the caecum takes a long time, the final clearance of this agent from the alimentary tract is difficult to determine. Radioisotope studies would certainly clarify such issues. The observation of faecal pellet formation being initiated in the spiral colon agrees with the observations by Macrae et al, (1973) of similar events in the spiral colon of the sheep. Pregnancy has a major effect on the abdominal topography where the enlarged uterus pushes most of the rumen complex and the entire intestine forwards into the cranial half of the abdomen. In these cases the caecum and spiral colon become located towards the cranial midline of the abdomen immediately in front of the uterus. Parra (1978), utilizing information on fermentation rates in many species of Artiodactyla of quite different body weight as well as data on the Langur monkey, Quokka, rabbit, porcupine and pig, calculates that in animals of less than 10 kg body weight, foregut fermentation is a less viable strategy than hindgut fermentation to meet the energy requirements of these smaller herbivores. Yet there a few species of herbivores and folivores below this arbitrary limit and the Lesser mousedeer is well below. For a forestomach fermenter, the size of the stomach complex relative to its body weight is critical. With a drop in body weight the metabolic rate per unit weight increases but the stomach size is limited, so much so that there is a disproportionately small amount of food available for fermentation (Parra, 1978). Fermentation rate cannot increase sufficiently, a possible compensatory mechanism, is the animal must improve the quality of its diet, increase the rate of passage of digesta or do both (Mertens, 1973). Generally an increase in the rate of passage of digesta drops the apparent digestability of the food. This drop is greater for poor quality high fibre foods than it is for food of higher quality (Blaxter, 1963). It is possible that the absence of an omasum in the mousedeer allows an increase in the passage rate of its digesta. This coupled with a selection of more nutritious, higher quality portions of plants such as fruits, nuts, tubers and fresh shoots would
PERTANIKA VOL. 11 NO. 3, 1988
THE RADIOGRAPHIC ANATOMY OF THE GASTROINTESTINAL TRACT OF THE LESSER MOUSEDEER probably allow the retention of forestomach fermentation in such a small ruminant. Detailed documentation of food actually ingested by mousedeer in their natural situation is necessary to clarify the issue. Similarly nutritional analysis of their actual diet, not supposed diet, is needed. ACKNOWLEDGEMENTS We wish to thank Encik Ganesamurthy, Encik Hamdy and Encik Sivasoorian for their able technical assistance. We thank Encik Palaniandy for his radiographic assistance and the Department of Clinical Sciences for allowing us the use of their facilities. REFERENCES
BLAXTER, K.L. (1963): The Energy Metabolism of Ruminants* London: Hutchinson Scientific and Technical. HUME, I.D. (1982): Digestive Physiology and Nutrition of Marsupials. Cambridge: Cambridge University Press. LANGER, P. (1988): The Mammalian Stomach. Stuttgart: Gustav Fisher.
Herbivore
MACRAE, J.C, REID, C.S.W., DELLOW, D.W. and WYBURN, R.S. (1973): Caecal cannulation in sheep. Res. Vet. Sci 14: 78-85.
MEDWAY, L. (1978): The wild mammals of Malaya (Peninsular Malaysia) and Singapore. 2nd Edn. Kuala Lumpur: Oxford University Press. MERTENS, D.R. (1973): Application of theoretical mathematical models to cell wall digestion and forage intake in ruminants. Ph.D. Thesis, Cornell University, Ithacca, New York. MORAT, P. and NORDIN, M. (1978): Maximum food intake and passage of markers in the alimentary tract of the Lesser Mousedeer. Mai Appl. Biol 7(1): 11-17. NORDIN, M. (1978): Voluntary food intake and digestion by the Lesser Mousedeer. /. Wild!. Man. 42(1): 185-187. PARRA, R. (1978): Comparison of the foregut and hindgut fermentation in herbivores. Pages 205-299 in G.G. Montgomery editor, "The Ecology of Arboreal Folivores: Washington D.C. : Smithsonian Institution Press. RICHARDSON, K.C. and WYBURN, R.S. (1980): The structure and radiographic analysis of the alimentary tract of the Tammai Wallaby, Macropus eugenii (Marsupialia) II The Intestines: Aust. J, Zool 28: 499-509. VIDYADARAN, M.K., HILMI, M. and SIRIMANE, R.A. (1982): The gross morphology of the stomach of the Malaysian Lesser Mousedeer (Tragulus javanicus). Pertanika 5(1): 34-38.
MESWAY, L. (1977): Mammals of Borneo: Monographs of the Malaysian Branch of the Royal Asiatic Society. (7). Kuala Lumpur.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 23 May, 1988)
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Pertanika 11(3), 437-439 (1988)
Preparation of Arsenobetaine hydrobromide •
HAZAR ISMAIL1 and ROBERT F. TOIA2 Department of Organic Chemistry University of New South Wales P.O. Box 1 Kensington, N.S. W. 2033, Australia.
Key words: Arsenobetaine hydrobromide; arsenobetaine: marine organoarsenicals. ABSTRAK Habluran arsenobetaina hidrobromida telah mendap terus apabila trimetiiarsina ditambahkan kepada satu larutan asid bromoasetik dalam benzena. Hasilnya adalah stabil, tak-lembab air dan boleh ditukarkan dengan mudah kepada betaina bebas dengan membenarkan ia melalui resin pertukaran ion berbes.
ABSTRACT Crystalline arsenobetaine hydrobromide is deposited directly on addition of trimethylarsine to a benzene solution of bromoacetic acid. The product is stable, non-deliquescent and may be readily converted into the free betaine by passage through basic ion exchange resin. INTRODUCTION Since the first report of arsenobetaine (Edmonds et al 1977) as a naturally occurring marine organoarsenical isolated initially from the flesh of the western rock lobster, Panulirus cygnus, numerous workers have reported this compound from a plethora of marine animals. (Ismail 1986) In addition, evidence has been obtained for the presence of arsenocholine in some marine animals (Norin & Christakopoulos, 1982) and a series of arsenosugars have been isolated from various marine algae. (Edmonds & Francesconi, 1981) Given its occurrence in human foodstuffs, arsenobetaine has also been the subject of metabolic studies. In a preliminary report (Cannon et ah 1983) the compound was shown, in mice. to be non-toxic and rapidly excreted in the urine and faeces. In addition, a negative result was obtained in the Ames' test for chemical mutagens (Cannon et al, 1983). Subsequent metabolic
studies have utilised radiotracer techniques, in particular 74 As labelled arsenocompounds, but in some cases are of questionable value since the work was carried with radiochemically impure material (Goetz & Norin, 1983). In this note, we report a modified approach to the synthesis of arsenobetaine which is suitable for adaptation to the preparation of labelled material of high chemical and radiochemical purity. MATERIALS AND METHODS General Melting points were determined on a Kofler hot stage and are uncorrected. Infrared spectra were recorded on an Hitachi Model EPI-G2 instrument for KBr discs and nuclear magnetic resonance spectra were recorded on either a Bruker CXP300 (300 MHz for ! H) or a JEOL FZ100 (25 MHz for 13 C) instrument. Chemical shifts are reported
Present address: J Pusat Asasi Sains, Universiti Malaya, 59100 Kuala Lumpur, Malaysia. 2 Pesticide Chemistry and Toxicology Laboratory, 201 Wellman Hall, Univeristy of California, Berkeley CA 94720 USA.
HAZAR ISMAIL AND ROBERT F. TOIA
relative to TMS, for CDCI3 solution of 3-(trimethylsilyl)propane sulphonic acid for D 2 0 solution, Dowex 2 x 8 (Cl~) ion exchange resin was cycled with 5% aqueous HC1, water and 5% aqueous NaOH until the titre was constant. The resin was washed thoroughly before use. Arsenic Analysis Arsenic analysia of aqueous samples utilised an inductively coupled plasma-atomic emission spectrometer (Labtam Plasmalab, Melbourne, Australia) monitored at 193.7 nm. The instrument was calibrated against As standards of 0 and 100 ppm sodium arsenate. Synthesis Iododimethylarsine and trimethyiarsine were prepared as described in the literature. (Burrows & Turner, 1920; Challenger & Ellis, 1935) Thus, sodium cacodylate (77.6 g) and Kl (160 g) were dissolved in water (300 ml). The solution was continuously saturated with SO2 whilst HC1 (10 M, 142 ml) was added dropwise. The product separated as it was formed, as a dark yellow oil. Distillation of the dried product gave iododimethylarsine as a pale yellow oil (73.7 g, 56% b.p. 80-84°C/35 mm (Burrows & Turner, 1920: 154-157°C/760 mm). Iododimethylarsine (68.4 g) in di-rt-butyl ether (100 ml) was added dropwise to methyl magnesium iodide (prepared from methyl iodide (40.8 g) and Mg (7 g)) in di-H-butyl ether (100 ml) under an atmosphere of nitrogen. The mixture was stirred continuously whilst maintaining the temperature at 4°C. On completion of the addition, the solution was allowed to warm to room temperature and then heated to distil trimethyiarsine (24.4 g, 69%, b.p. 54°C (Burrows & Turner, 1920: 51.9-52°C) directly. The product was stored in ampoules under nitrogen for use as required. Arsenobetaine hydrobromide: Bromoacetic acid (0.41 g) was dissolved in benzene (2.5 ml) and trimethyiarsine (0.5 ml) added dropwise under nitrogen. The solution initially turned green, then blue and finally colourless at which point a crystalline solid began to deposit. The reaction mixture was left overnight. The solvent was removed and the residue twice re crystallised from methanol/acetone to give arsenobetaine hydrobromide as prisms, 0.62 g (81%), m.p. 248°C decomp., 438
dependent on the rate of heating). Found C, 23.16%; H, 4.60%. C 5 H 15 AsBr0 2 requires C, 23.10%; H, 4.90%. I.R: V m a x 3450, 3050, 2950 2900, 2850, 1715, 1640, 1400, 1365, 1275, 1260, 1170, 940, 935, 910, 900, 850 cm" 1 . ^ N M R : 5 1.95 (s, 9H), 3.58 (s, 2H). 13 CNMR 5 8.6 (q), 31.3 (t), 170.5 (s). Arsenobetaine picrate: Arsenobetaine hydrobromide (0.1 g) was dissolved in hot ethanol and an ethanolic solution of picric acid (0.1 g) added. On cooling, crystals of arsenobetaine picrate were deposited and were recrystallised from hot ethanol as yellow needles, 0.15 g (94%), m.p. 177-178°C. Found C, 32.37%; H, 3.68%; N, 10.14%. C n H 1 4 AsOgN 3 requires C, 32.43%; H, 3.44%; N. 10.14%. Arsenobetaine reineckate: Arsenobetaine reineckate was prepared from arsenobetaine hydrobromide as described in the literature (Edmonds et ai, 1977) for arsenobetaine. The product was found to be identical in all respects with an authentic sample. Arsenobetaine: Arsenobetaine hydrobromide (1.0 g) was dissolved in distilled water (2 ml) and passed through a column of Dowex 2 (OH") (100 g). The column was eluted with distilled water and the fractions analysed for arsenic. The arsenic-rich fractions were combined and lyophillised to yield arsenobetaine as a white solid. The product was dissolved in dry methanol and crystallised, on the slow addition of acetone, as white prisms, 0.6 g (87%), identical in all respects with an authentic specimen. 13 C NMR: 5 8.3 (q), 34.4 (t), 172.7 (s). RESULTS AND DISCUSSION Arsenobetaine (1) is an extremely deliquescent, white crystalline compound which has previously been prepared (Edmond et al, 1977) as shown in scheme 1 path A. In the present work we have modified this preparation, scheme 1 path B, by reacting trimethyiarsine with bromoacetic acid. This leads directly to arsonebetaine hydrobromide which crystallises from the reaction mixture as it forms. The hydrobromide is a non-deliquescent, white, crystalline and stable solid. As observed for arsenobetaine (Edmond et ah 1977) the melting point of the hydrobromide varies with the rate of heating and decomposition accompanies melting.
PERTANIKA VOL. 11 NO. 3, 1988
PREPARATION OF ARSENOBETAINE HYDROBROMIDE ACKNOWLEDGEMENT
(CH3) 3 As + CH 2 CO 2 ei Dr"
Dowex 2 OH
One of us (HI) acknowledges the Public Service Department of Malaysia and Universiti Malaya for a scholarship. REFERENCES
BURROWS, G.J. and E.E. TURNER. (1920): A new type of compound containing arsenic. /. Chem, Soc, 117: 1372. (CH 3 )3AsCH 2 CO2H Br Scheme 1
The C NMR spectra of arsenobetaine and the hydrobromide in D2O solution show only minor differences in chemcial shift values as would be anticipated and these differences may be ascribed to solution pH differences. The picrate and reineckate derivatives prepared from both arsenobetaine and its hydrobromide were identical in all respects. With respect to the preparation of radiolabelled compounds, the ease of preparation of the hyrobromide, and more importantly its stability and the ease with which it may be recrystallised, make it an ideal compound to work with. Moreover 14C-labelled bromoacetic acid is commercially available although a label in the other portion of the molecule is also feasible. In particular, whilst it is possible to use the earlier strategies (Goetz & Norin, 1983) with arsenic isotopes, 14 C-or 3H-labelled trimethylarsine is also feasible. Possible tritium labelling of trimethylarsine is particularly attractive since this may be effected, in principle, via direct exchange and the product may be purified by distillation. Since this work is directed in part to a study of the metabolism of arsenobetaine in model marine ecosystems, (Ismail Hazar, 1986) the purified hydrobromide may be used directly. Howver, in studies requiring the free betaine. passage of the salt through a basic ion exchange resin readily produces arsenobetaine.
CANNON, J.R., J.B. SAUNDERS and R.F. TOIA, (1983) : Isolation and preliminary toxicological evaluation of arsenobetaine-the water-soluble arsenical constituent from the hepatopancreas of the Western Rock Lobster. Set Total Environ,, 31 : 181. CHALLENGER, F. and L. ELLIS. (1935) : The formation of organo-metalloidal compounds by micro-organisms part III methylated alkyl- and dialkyl-arsines. /. Chem. Soc. : 396. EDMONDS, J.S., K.A. FRANCESCONI, J.R. CANNON, C.L. RASTON, B.W. SKELTON and A.H. WHITE. (1977): Isolation, Crystal structure and synthesis of arsenobetaine, the arsenical constituent of the Western
Rock
Lobster Panulirus logipes cygnus
George. Tetrahedron Lett : 1543; see also Cannon, J.R., Edmonds, J.S., Francesconi, K.A., Raston, C.L., Saunders, J.B., Skelton, B.W. and White, A.H., (1981) : .Isolation, crystal structure and synthesis of arsenobetaine, a constituent of the Western Rock Lobster, the Dusky Shark and some samples of human urine. Aust J. Chem., 34 : 787. EDMONDS, J.S and K.A. Francesconi. (1981) : Arsenosugars from brown kelp {Ecklonia radiata) as intermediates in cycling of arsenic in a marine ecosystem. Nature, 289 : 602. GOETZ, L. and H. NORIN. (1983) : Synthesis of 73 Asradiolabelled arsenobetaine and arsenocholine for metabolic studies and their fate and distribution in laboratory animals. Int. J. Appl Radiat. hot, 34: 1509. ISMAIL, H. (1986) : Ph.D. Thesis, University of New South Wales. NORIN, H. and A. CHRISTAKOPOULOS. '1982) : Evidence for the presence of arsenobetaine and another organoarsenical in shrimps. Chemosphere, 11 :287.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 7 May, 1988)
439
Pertanika 11(3), 441-449 (1988)
Malaysian Natural Rubber Market Model MOHAMMED BIN YUSOF
Department of Economics, Faculty of Economics and Management Universiti Pertanian Malaysia 43400 Serdang, Selangor Darul Ehsan, Malaysia. Key words: Natural rubber; model; simulations. ABSTRAK
Tujuan kajian ini ialah untuk membentuk satu model persamaan serentak pasaran getah asli Malaysia. Keputusan regresi menunjukkan bahawa model yang dibentuk ini adalah memuaskan di segi tandatanda yang betul, R2 yang tinggi, dan pttnbolehubah yang bermakna. Simulasi juga dicuba untuk melihat kebolehan model ini mengesan nilai sebenar. Pada umumnya, model ini berupaya mengesan pergerakan pembolehubah endogen terpilih. Model ini boleh digunakan untuk membuat ramalan kesan perubahan sesuatu pembolehubah eksogen, seperti cukai eksport, kadar pertukaran, atau kemelesetan ekonomi, ke atas pembolehubah endogen.
ABSTRACT
This study is an attempt to formulate a simultaneous equations model of the Malaysian natural rubber market. The regression results suggest that the model is quite satisfactory in terms of correct signs, high R2, and significance of variables concerned. A simulation exercise was done to ascertain the adequacy of the model in tracking the actual values. It was found that, in general, the model has the ability to trace, at least, the directions of the movements of certain selected endogenous variables. The model developed here could be used to forecast the effect of a change in an endogenous variable, such as export duty, exchange rates, or recession on endogenous variables. INTRODUCTION
The importance of foreign trade to the Malaysian economy is fairly well accepted by policy makers and academics. Substantial portions of the Malaysian GNP is derived from the foreign sector. For example, in 1980 and 1985 the ratios of the value of exports to GNP were 54.5 percent and 49.5 respectively. One of the major commodity exports of Malaysia is natural rubber. Although the share of natural rubber in the total exports has been declining in recent years, its contribution to Malaysia's foreign exchange earnings is still substantial. For example, the share of rubber export in the total exports in 1980 was 16.4 percent and by 1985 it declined to 7.9 percent. Due to recent world recession and competition with synthetic
rubber, the world demand for natural rubber seems to be stagnant. Thus, steps should be taken to maintain its competitiveness in the world market, for example, by lowering the unit cost of production. In order to analyse the effects of policy actions, a Malaysian natural rubber market model is formulated in this study. The model specifications presented here represent a significant departure from previous studies such as Yusoff (1978). The model could be used to analyse the effects of, for example, world recession, exchange rate adjustment, and export tax on production acreage, prices, and export demand for natural rubber. Historical simulation exercise was carried out by using this model to ascertain its adequacy
MOHAMMED BIN YUSOF
for forecasting. The results suggest that the model has the ability to trace the historical data reasonably well. THE MODEL In this section, we shall describe briefly a simple model of the Malaysian natural rubber market. The model consists of six behavioral equations and five identities. The behavioral equations are the equations for export demand, the world price of natural rubber, acreages for estate and smallholding sectors, and the yields of estate and smallholdings sectors; while the identities define the production of smallholding and estate sectors and the price of. rubber in Malaysian ringgit, the price received by the producers, and the total production. In the discussion that follows the disturbance term in each of the behavioral equations will be suppressed for simplicity but without loss of generality.
mand for rubber to increase, other things unchanged. The exchange rate, e, was computed by finding the average exchange rate of ringgit in terms of Malaysian major trading partners' currencies, namely the U.S. $, Singapore $, U.K. pounds, Japanese yen, and German mark, weighted by their respective trade shares. For more detail, see Yusoff (1987). The variable IPIW represents the economic activities of the world. Since natural rubber is a raw material used by industries, it is expected that if industrial activities are higher, the demand for natural rubber would increase. The trend variable t is supposed to indicate the change in the tastes and preferences of the importing nations. World Price The world price of natural rubber, PRW is specified as log PRWt = j30 + ft log PSRt + ft log IPIWt + ft
Export Demand Following Yusoff (1978), the export demand is specified as Log XR t = OQ + <*! log (FR*e/PSR)t + a2 l o g t + a 3 logIPIW t
(1)
where XR = foreign demand of Malaysian rubber in metric tons PRW = world price of rubber in foreign currency per metric ton in Kuala Lumpur market PSR = world price of synthetic rubber in foreign currency per metric ton in the U.S. market e • effective exchange rate (foreign currency per ringgit) t = year IPIW • world industrial production index. Equation (1) postulates that the foreign demand of natural rubber from Malaysia would be negatively related to the relative price of natural rubber to synthetic rubber and positively related to the world industrial production index. If the price of natural rubber is higher in relation to the synthetic rubber, the quantities demanded would fall and vice-versa; if the value of Malaysian currency depreciates, we would expect the de442
logRSRWt-1 (2) where RSRW = ratio of stock of natural rubber between producing and consuming countries. Since natural rubber and synthetic rubber are substitutes, these prices should be positively correlated; an increase in the price of synthetic rubber would encourage consumers to switch to natural rubber, thereby raising the price of natural rubber. And a fall in the industrial production index would reduce the demand for natural rubber and consequently, its price falls. The presence of the stock variable suggests that if the stock in the consuming countries are lower in relation to stock in producing countries, the price will rise as many consuming countries purchase rubber in the world market to replenish their stocks. Acreage Rubber growing in Malaysia is composed of two distinct sectors, namely the estates and the smallholdings. The acreage equations are specified as Log AERt = Log (AER) t
PERTANIKA VOL. 11 NO. 3, 1988
0 + 0! log (PR/CPI)t 62 log t
(3)
log (AER) t _j m ^ [log AER* -logAERt_1]
(4)
MALAYSIAN NATURAL RUBBER MARKET MODEL
where AER* = optimal harvested * acreage of rubber in estate PR = price of rubber in ringgit per metric ton CPI = consumer price index in Malaysia PR/CPI = deflated rubber price AER • actual harvested acreage in estate Since AER* is an unknown quantity, it is then approximated by an adjustment process (4). Substituting (3) in (4) for AER*, we obtain log AERt =
X?
0O
+
*2 01 log (PR/CPI)t + A* e2
logt + U - X ^ l o g A E R ^
(5)
Equation (3) suggests that the harvested acreage is determined by the price of rubber deflated by the consumers price index. As the price of rubber increases in relation to the price of the producers for other crops in relation to price index, we would expect the harvested acreage to increase. The trend variable t is to capture the change in taste and the preferences of the producers for other crops in relation to natural rubber. Thus, if the returns from rubber is low in relation to other crops and economic activities, a profit maximizing grower should divert his resources to the more lucrative business opportunities. In a similar manner, the harvested acreage for the smallholding sector, is specified as logASH t = 50 + 5! log(PR/CPI)t + (6) log(ASH)t • X3 [log(ASH)*-log(ASH) t -l]
(7)
where ASH* = desired acreage of smallholding harvested ASH = actual acreage of smallholding harvested Substituting (6) in (7) for log ASH* , we obtain
Equation (8) is the estimating equation for acreage. Since the actual harvested acreage for smallholders is unknown, it is replaced by the actual matured acreage for estimation purposes. This equation is used to forecast smallholdings production through equation (16). Simulation results suggest that equation (16) could corecast smallholdings production reasonably well. Therefore, we could proxy matured acreage for harvested acreage for our purpose here. YIELD The yield of any crop is expected to depend on the technological breakthrough in the industry. As new high yielding varieties are discovered, we would expect the yield to rise accordingly. The yield equation is a variant of Fisher and Temin (1978) model and is written as log (YRE)* = So + Silog
(9)
log(YRE) t -log(YRE) t _ 1 = *4 [log(YRE)J -
(10)
where YRE = optimal yield YRE = actual yield Equation (9) says that the optimal yield would depend on technological progress which is represented by time trend t. Substituting (9) into (10) for log (YRE*), we obtain log(YRE)t = X4 50 + *4 5X log t + (l-X4)log(YRE)t_1
(11)
Similarly, the yield in the smallholders sector could be written as log t lo
(12)
S (YRSH)t - log (YRSH)t_j (13)
log ASHt = X3 50 + \3 §! log (PR/CPI)t + X3 52 log t + (1 - X3) log ( A S H ) t l
0
(8)
0
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MOHAMMED BIN YUSOF
log (YRSH)t -
4>i log t + (1-X,) (14)
The actual harvested acreage for smallholders is not available. Therefore YSRH was approximated by dividing its production with actual matured acreage. Since actual matured acreage is expected to be larger than harvested acreage, then YSRH calculated in this study tends to underestimate the actual YRSH. The lagged yield as a regressor could be interpreted in the following manner. Write equation (10), for example, as (10.1)
Y t -Y t _,
Production In this model, quantity produced or simply production is identically equal to the yield multiplied by harvested acreage. Thus, for the estates, it is written as QRE t = YRESt * ARE t (15) and for the smallholdings, it is QRSHt - YRSHt * ARSHt Where QRE = QRSH =
(16)
production from estate production from smallholdings.
This form of production identity is employed by Shonkwiler and Emerson (1982). Identities
where Y Y
= =
actual yield potential (desired) yield.
We shall close the model by defining three identities, namely the world price of natural rubber in ringgit, the price received by the exporters, and total rubber production in Malaysia as follows:
Rearranging (10.1), we obtain
(17)
0 0 .2)
By process of lagging and substituting, (10.2) could be written as
PRR t = P R t - X T R t and QR t = DCt + XR t + AlRt
(18) (19)
where (10.3)
QR DC
As i ~^°°, equation (10.3) reduces to
£
AIR (10.4)
Lagging (10.4) by one period,
i=0
(10.5)
It is very clear from (10.5) that the lagged actual yield is positively related to the past potential yields. And of course these past potential yields were the result of research and development efforts in the past, which in turn, are positively related R & D expenditures. Thus, the past R & D expenditures are captured by the lagged actual yield, Y t - 1 444
= total rubber production in Malaysia in metric tons = domestic consumption of natural rubber in metric tons • change in rubber inventory in metric tons.
Equation (17) defines the domestic price of rubber as equal to the world price in foreign currency divided by the effective exchange rate in foreign currency per ringgit. Equation (18) on the other hand says that the price received by the exporters, PRR, is equal to rubber price in domestic currency less the export tax. This simulatneous. equations model has six behavioral equations and five identities. The behavioral equations are the export demand, world price, matured acreage disaggreaged into estates and smallholdings, and the yield equations for both estates and smallholdings. The identities define quantities produced in estates and small-
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MALAYSIAN NATURAL RUBBER MARKET MODEL
holdings, world price, the price received by the exporters, and the total production. The endogenous variables are: export demand, world price, price received, world price in domestic currency, harvested estates acreage, matured smallholding acreage, yield of estates, yield of smallholdings, quantities produced by estates, and quantities produced by smallholdings. The exogeneous variables (including predetermined variables) are the price of synthetic rubber, time trend, consumer price index of Malaysia, world industrial production index, effective exchange rate, export tax, lagged ratio of world stock in producing
countries compared to consuming countries, and the lagged endogeneous variables of production, acreages, and yields of both estates and smallholdings. Since a number of equations involved ratios and nonlinear in parameters, the model was estimated by nonlinear 2 SLS. The data for this study (from 1960 to 1981) were extracted from the Quarterly Economic Bulletin of Bank Negara Malaysia, Rubber Statistics Handbook of the Department of Statistics of Malaysia, and International Financial Statistics of the International Monetary Fund.
TABLE 1 Nonlinear 2SLS estimates Export Demand log (XR)t • 729.10 - 0.0912 log (PR*e/PSR)t - 96.1616 log (t) + 1.6765 log IPIWt (1.27) (3.94)** (7.07)** R2 = 0.9652, DW = 1.62 World Price log (PRWt) = 1.9232 + 0.8064 log PSRt + 0.1238 log IPIWt + 0.2714 (3.26)** (0.30) (0.1234) R2 = 0.6811, DW = 1.97 Acreage logAERt= 127.50 + 0.0067 log (PR/CPI) - 16.5320 log t + 0.7098 log ( A E R ) t l (0.77) (3.71)* (7.81)** 2 R = 0.9987, h=0.81 log ASHt • -34.3153 + 0.0065 log (PR/CPI) + 4.8236 log (t) + 0.7088 log (ASH) (0.46) (2.09) (12.30)** R2 = 0.9919, h= 1.021 Yields log (YRE)t = -44. 1963 + 5.8124 log(t) + 0.8291 log ( Y R E ) t l (0.52) (5.13)** R2 = 0.9629, hc = 2.021 log(YRSH)t = -215.00 + 28.2832 log(t) + 0.7008 log (YRSH) t l (1.65) (4.14)** R2 = 0.9373, h c = 1.81 t hc
- values are in the parentheses; - the equation was corrected for autocorrelation. PERTANIKA VOL. 11 NO. 3, 1988
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MOHAMMED BIN YUSOF
gestation period. Thus, as long as the market price RESULTS is above the average variable cost, rubber will be Generally speaking, the results of the regression harvested. Therefore, it is found that the elasticity analysis are satisfactory in terms of goodness of fit (R 2 ), significance of the explanatory variables, of acreage will with respect to price is very low, and correct signs (Table 1). almost perfectly inelastic at 0.0067. The coefficient of the time trend is negative and signifiThe foreign demand for Malaysian rubber cant at one percent level suggesting that the would depend on the performance of the world acreage is declining over time. economy represented by the world industrial production index. The elasticity of the demand This is expected since many estates had with respect to industrial production index was diverted substantial acreage under rubber to grow estimated to be 1.6765, which is significant at oilpalm. The lagged acreage is also significant and one percent level and elastic. Thus, a one percent the adjustment to the desired acreage is quite slow change in industrial production index would as indicated by the low coefficient of adjustment result in about 1.7 percent increase in the demand at 0.29, implying that about 30 percent of actual for Malaysian rubber. The coefficient of the time acreage is adjusted to the desired level per year. trend is significant at one percent level and negaThe results of the smallholding acreage equation tive suggesting that the foreign demand for rubber follow closely to that of the estate, except that from Malaysia is declining overtime, perhaps in its acreage is on an upward trend since the smallpreference of its closest substitute, synthetic holding sector has been expanding in recent rubber or that more importers are opting to years. purchase rubber from other producing countries The yield equations for both the estate and such as Indonesia, Thailand, and Sri Lanka. This smallholding produce similar results. The only trend is indeed very alarming. Malaysia therefore important determinant of yield is the lagged yield should take steps now to be more aggressive in its itself which should capture the effect of techefforts to promote rubber in terms of quality, nological progress. Both indicate that adjustment price, and services. As was found in other studies, to the desired yields are slow at 17 percent and for example Yusoff (1978), the price of natural 29 percent for estate and smallholding respectiverubber cannot explain the change in quantity lydemanded and its elasticity with respect to relative SIMULATION RESULTS price is very inelastic, at 0.0912. A dynamic simulation was carried out through the The results also suggest that the price of sample period to see how far the model could natural rubber could be significantly explained by track the path of the actual data. We shall measure the price of synthetic rubber; a ten percent this tracking ability by using the mean percent increase in the price of synthetic rubber would error and root mean-square percent error (Table increase the price of natural rubber by 8 percent. 2). The ratio of stock of natural rubber in producing countries and consuming countries is also an The mean percent error of the endogenous important determinant of the natural rubber price, variables are relatively small (less than two perwhere it is significant at 5 percent level. It is cent) except for the production of the estates estimated that a ten percent rise in the stock in the where the mean percent error is 9.40 percent. But producing nation in relation to stock in consuming the root mean-square percent error criterion countries would result in a 2.7 percent increase in indicates that the errors are more than five percent the price of natural rubber. Thus, assuming that for prices, yield of smallholders, and production the consuming countries would not release their of both smallholders and estates, although they are stocks, one of the ways to increase the price of still less than ten percent. The root mean percent natural rubber is through accumulating its stock error for the yield of estates, export, and acreages in the producing nations, if funds are available. are all less than five percent. The price of natural rubber is found not to In order to see more clearly the paths of be an important determinant of harvested acreage selected endogenous variables, their simulated and in the estate sector. This is understandable since actual values are graphed as shown in Appendices investment in rubber production involves a long 1, 2, 3, 4, 5 and 6. By and large, the model could 446 PERTANIKA VOL. 11 NO. 3, 1988
MALAYSIAN NATURAL RUBBER MARKET MODEL TABLE 2 Historical simulation results Endogenous variables
Root mean square percent error
Mean percent error
PRW
1.50
7.42
YRES
0.07
3.78
YSHR
0.29
7.42
PR
1.50
7.42
XR
0.11
4.66
AER
0.02
0.59
ASHR
0.02
0.85
QER
9.40
9.78
QSHR
0.28
7.06
trace the directions of the actual values quite well. The simulated values trace the actual world price of natural rubber satisfactorily; a number of critical turning points (1976, 1977, and 1980) were correctly predicted by the model. The acreage for estates has been declining steadily during the sample period and the simulated values trace the path quite closely. The smallholdings acreage, on the other hand is on an upward trend and its simulated values also follow the actual path satisfactorily. The model was not able to predict export as well as we would expect; it almost consistently underpredicts the export. This suggests that although an equation is satifactory in terms of correct signs, significance of the regressors, and high R 2 , it does not necessarily imply that the equation could predict very well when all the equations are simulated. The simulated values of the production of estates also follow closely to that of the actual values; it correctly predicts two critical turning points in 1970 and 1973, although it misses a turning point in 1974. The quantity of production in the smallholdings is on an upward trend and the simulated values follow the movement of the actual values quite well.
100
CONCLUSION This study is an attempt to formulate a model of the Malaysian natural rubber market. Simul-
A simulation exercise was done to ascertain the adequacy of the model in tracking the actual values. It was found that, in general, the model
World price 350 300
Actual Simulated
200
1963
1965
1970
1975
1980 Year
Appendix 1: World Price of Natural Ruboer
taneous equations model of production, acreage, yield, export, and price was developed and then estimated by using nonlinear two-stage least squares. The regression results suggest that the model is quite satisfactory in terms of correct signs, high R 2 , and significance of variables concerned.
PERTANIKA VOL. 11 NO. 3, 1988
447
MOHAMMED BIN YUSOF
Export
Actual Simulated
1600 1500
p
1250 1000
1963
1965
1970
1975
1980 Year
f
Actual Simulated
1000 Appendix 2: Acreage (estates) 8000 1963 1965
Acres
1970
1975
1980 Year
3000 Appendix 4: Export
Y Production 700
Actual Simulated
2500
650
\ 600
2000
1963
550 1965
1970
1975
1980 Year
Appendix 3: Acreage (Smallholdings)
has the ability to trace, at least, the directions of the movements of certain selected endogenous variables. The model developed here could be used to forecast the effect of a change in an exogenous variable, such as export duty, exchange rates, or
448
Actual Simulated 500 80 1963
1965
1970
1975
1980 Year
Appendix 5: Production (Estates)
PERTANIKA VOL. 11 NO. 3, 1988
MALAYSIAN NATURAL RUBBER MARKET MODEL Production
FAIR, R.C. (1980): Estimating the Uncertainty of Policy Effects in Nonlinear Models, Econometrica. 48(6): 1381-1391.
1000
FISHER, F.M and P. TEMIN. (1970): Regional Specialization and the Supply of Wheat in the US., 1967-74. American Journal of Agricultural Economics, VoL Lll,No. 2,121-134.
800
TAN, A.H. (1967): The Incidence of Export Tax on Small Producers. Malayan Economic Review, 12(2):
600
SHONKWILER, J.S. and R.D. EMERSON. (1982): Import and the Supply of Winter Tomatoes: An Application of Rational Expectations. American Journal of Agricultural Economics, 64(4): 634-641.
Actual Simulated
400
INTERNATIONAL MONETARY FUND. International Financial Statistics. Various Issues.
300
1963 1965
1970
1980 Year
1975
Appendix 6: Production (Smallholding)
recession on endogenous variables which is not done in this study. REFERENCES
YUSOFF, MOHAMMED B. (1978): An Econometric Model of the World Natural Rubber Industry. Kajian Ekonomi Malaysia, 25(2): YUSOFF, MOHAMMED. B. (1987): Exchange Rate Changes and the Terms of Trade. PKDP Report, No. 17/87, Centre for Agricultural Policy Studies, Fakulti Ekonomi & Pengurusan, Universiti Pertanian Malaysia.
BANK NEGARA MALAYSIA. Quarterly Economic Bulletin. Various Issues. DEPARTMENT OF STATISTICS, MALAYSIA. Rubber Statistics Handbook. Various Issues.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 4 August, 1987)
449
Pertanika 11(3), 451-460(1988)
Real Money Balances in the Production Function of a Developing Economy: A Preliminary Study of the Malaysian Agricultural Sector MUZAFAR SHAH HABIBULLAH Department of Economics Faculty of Economics and Management Universiti Pertanian Malaysia 43400 Serdang, Selangor, Darul Ehsan, Malaysia. ABSTRAK Bukti empirikal berpendapat bahawa baki wang benar boleh dianggap sebagai input yang produktif dalam pengeluaran. Baki wang benar dimasukkan sebagai faktor pengeluaran adalah kerana baki wang benar sebagai alat perantaraan pertukaran memudahkan penyesuaian di antara modal dan buruh untuk tujuan pengkhususan dan dengan itu menambahkan produktiviti. Kajian ini cuba menguji secara empirikal bahawa baki wang benar adalah input dalam fungsi pengeluaran di dalam sektor pertanian di Malaysia. Keputusan empirikal menyatakan bahawa baki wang benar memainkan peranan yang penting sebagai input yang produktif dalam fungsi pengeluaran sektor pertanian di Malaysia. ABSTRACT Recent empirical evidence suggest that real money balances can be treated as a productive input in production. The reason for incorporating real money balances as a factor of production is because real money balances as a medium of exchange facilitate adjustments between capital and labour for specialisation purposes and thus increase productivity. This study is an attempt to empirically test the evidence that real money balances is an input in the production function of the Malaysian agricultural sector. The results suggest that real money balances play a significant role as a productive input in the production function of the Malaysian agricultural sector. I. INTRODUCTION Traditionally in production, output has been specified as a function of capital and labour. This technical relationship between output and input has been recognised by economists for over half a century. More recently Sinai and Stokes (1972) provided empirical evidence which suggests that real money balances are a third factor input in the production function. Earlier Friedman (1959, 1969), Bailey (1971), Johnson (1969), Levhari and Patinkin (1968), Moroney (1972) and Nadiri (1969) suggested that real money balances were a factor of production. But, it is Sinai and Stokes (1972) who provided the pioneering empirical work on this issue. Despite their empirical evidence the idea of money balances as a productive input has been criticised by Fischer (1974), Nicolli (1975), Prais (1975a, 1975b), Khan and Kouri (1975), Ben-
Zion and Ruttan (1975) and Boyes and Kavanaugh (1979). They argued that incorporating real money balances in the production function is subject to specification bias. Nevertheless, recent empirical evidence by Simos (1981), Apostolakis (1983), You (1981), Short (1979), Subrahmanyam (1980) and Khan and Ahmad (1985) provide strong support that real money balances act as a productive input in production. The reason for incorporating real money balances as a factor of production is because real money balances as a medium of exchange, facilitates the exchange between capital and labour for specialisation purposes and thus increases productivity. Also, it reduces the transaction cost and therefore, increases the economic efficiency of the money market system (Sinai and Stokes, 1972; Short, 1979; Khan and Ahmad, 1985; Finnerty, 1980).
MUZAFAR SHAH HABIBULLAH
This study is an attempt to empirically test the evidence that real money balances act as a productive input in the agricultural production of Malaysia. The paper is divided into four sections. Section II provides the literature review, model used and data. In section III. empirical results are presented and discussed, and the final section contains the conclusion. II. METHODOLOGY
Since the paper by Sinai and Stokes (1972), numerous studies have been done to substantiate the role of real money balances in production. Short (1979), using both the Cobb-Douglas and translog models, came to the conclusion that, the decision to hold real money balances like any other productive input is based on rational, profit maximising considerations. He found that real money balances are positive and statistically significant for both the Cobb-Douglas and translog models. Results from You (1981) were similar to those by Sinai and Stokes (1972). Subrahmanyam (1980) and Simos (1981), both agreed with You (1981), and further concluded that real money balances are substitutes for capital, but complement with labour. The study by Apostolakis (1983), on the other hand, found that, real money balances substitutes for labour and complements capital services. All the above studies were centered on the developed countries. However, Khan and Ahmad (1985) using a multi-equation framework examined the role of real money balances in the production function of the manufacturing sector in Pakistan. Their conclusion was in accordance with the results of Sinai and Stokes (1972). The Model Following Sinai and Stokes (1972), Khan and Ahmad (1985), Short (1979) and Finnerty (1980), the general form of the three-input production function can be written as; Qt = f ( K t , L t , M t ) where Q. is quantity of output produced, K t , Lt and Mt are the amount of capital, labour and money stock employed in production. Empirical emphasis was on seven Malaysian agricultural subsectors - rubber, oil palm, tea, coconut, forestry, paddy and fishery. The production function in log-linear form was specified as: 452
Log Q i t = Oo + ax log K it + o^ log L i t + a 3 log(M j /P) t + U i t where i
j
P U
(3)
= respective sub-sectors; rubber, oil palm, tea, coconut, forestry, paddy and fishery, = alternative measurement of money stocks used in the model; Ml, M2 orM3, = price level measured by consumer price index, and = the disturbance term.
For each of the subsectors, three models one for each measure of real money balances (Ml, M2 or M3) were estimated. As suggested by economic theory, we would expect a positive relationship between output, Q and the inputs capital, labour and real money balances. The measure of real money balances would represent real purchasing power over factor inputs, and an appropriate measure for capturing the role of money in the production process (Short, 1979). Method of Estimation and Data This study is based on Malaysian time series data over the period 1960-1985. The money variables used in this study are money stock Ml (defined as currency plus demand deposits held by nonbank private sector), M2 (Ml plus saving deposits and fixed deposits held at commercial banks), and M3 (M2 plus saving deposits and fixed deposits at other financial institutions; namely, finance companies, merchant banks, National Saving Bank and Employee Provident Fund). The money stocks variables were deflated by the consumer price index (1967=100) to arrive at real money balances. All outputs are in metric tonnes except for coconut which is the number of nuts collected per year. Labour refers to the number of workers employed in the production in each subsectors. However, some problems arise in measuring capital, particularly in the developing countries. Although the above production function specified in equation (1) has been used for both the developed countries and the developing countries, the functional form will not be appropriate for some of the developing countries at least for two reasons. Firstly, the role of capital is not as evident as the role it plays in the advanced developed
PERTANIKA VOL. 11 NO. 3, 1988
REAL MONEY BALANCES IN THE PRODUCTION FUNCTION OF A DEVELOPING ECONOMY
countries. In an economy like Malaysia, producers in the agricultural sector operate their farm on a relatively smaller scale as compared to their counterparts in the more advanced countries. Thus, instances of fixed capital input ownership by the agricultural producers are rare. Secondly, published data on capital services in the developing countries are not available. Therefore, as an alternative, in the actual estimation, area planted was substituted for the measurement of capital inputs since area planted includes all the trees planted, livestock reared, labour building structures and farm implements stationed on the land which is a form of investment (Booth and Sundrum, 1984). Strong and Suhaila (1987) also employed area planted to proxy for capital services in their study on oil palm sector in Malaysia. Data on financial variables are collected from various issues of Quarterly Economic Bulletin published by Bank Negara Malaysia. Data on agriculture were compiled from publications by the Department of Statistics, Ministry of Agriculture and (their respective departments) which include Rubber Statistics Handbook, Oil Palm, Cocoa,. Coconut and Tea Statistics, Paddy Statistics, Annual Fisheries Statistics, and Forest Statistics Peninsular Malaysia. III. EMPIRICAL RESULTS
In this study all estimated regression equations were corrected for autocorelation that might arise due to the nature of time series data. The results of the estimated equations are presented in Tables 1 through 7; for rubber, oil palm, tea, coconut, forestry, paddy and fishery subsectors, respectively. For each subsector, four models were estimated. Model I was estimated without real money balances as one of the regressors. Models II, III and IV were estimated with Ml, M2 and M3 respectively. The purpose of estimating Model I is to act as a standard model for comparison between Models II, III and IV. Thus, comparisons can be made with respect to the measure of real money balances. For the rubber subsector, the results of the estimated regression equations are presented in Table 1. The results show that all estimated coefficients were not significantly different from zero, except for labour in Model I, which has a negative sign, and Ml, M2 and M3 in Models II, III, and IV respectively.
Table 2 shows the result for the oil palm subsector. The results clearly show that all variables are important except for labour in Model I, II and III, and capital in Model IV. Real money balances are significant at the one percent level for Ml, M2 and M3. With the inclusion of real money balances, the estimated coefficient of capital has reduced from 1.11 in Model I to 0.88 and 0.47 in Models II and HI. The estimated regression equations for the tea subsector are shown in Table 3. The results show that capital and labour are significant at the five percent level in Models III and IV. On the other hand, real money balances are significant at the one percent level in Models II, III and IV. In all cases, the inclusion of real money balances in the production increases the estimated coefficient of both capital and labour in the tea subsector. The results for the coconut subsector are presented in Table 4. In Models I through IV, the estimated coefficient for capital and labour employed are significant at the one percent level, however, real money balances are not significant. Furthermore, real money balances exhibit negative signs. Therefore, the results suggest that real money balances do not play an important role in coconut subsector. For the forestry subsector, the results are shown in Table 5. It can be seen that capital and labour are important except for labour in Model IV. Real money balances are only significantly different from zero in Model II. In this model we can see that the estimated coefficient for capital was reduced from 0.097 in Model I to 0.086 in Model II. The results for the paddy subsector are presented in Table 6. However, in the paddy subsector, estimations were made without including labour due to the unavailability of data on workers employed in this subsector. The results show that all variables are significantly different from zero. However, with caution the results suggest that real money balances play an important role in the paddy subsector. Lastly, Table 7 show the results for the fishery subsector. In this subsector, due to unavailability of data on capital, only labour and real money balances are included in the final estimation. Only Model IV gave the 'best' results in terms of the significance of the variables. However, as in the paddy subsector, with caution we can
PERTANIKA VOL. 11 NO. 3, 1988
453
TABLE 1 Regression coefficients and related statistics for the Rubber sector Variables3
Model I
Model II
Model III
Model IV
Constant
5.8948 (0.91680)
5.2017 (0.93214)
6.2713 (1.1132)
7.1007 (1.2252)
K
0.49035 (0.56584)
0.12910 (0.16926)
-0.13581 (-0.17182)
-0.22353 (-0.27012)
L
-0.47091 (-2.5498)**
-0.04786 (-0.22035)
0.09963 (0.38556)
0.06757 (0.25143)
3
(Ml/CPI)
0.34766 (2.9202)***
(M2/CPI)
0.30585 (2.8250)*** 0.27196 (2.5497)***
(M3/CPI)
o R-square D.W. Rho SER d.f.
0.9783 2.1125 0.9220 0.0607 23
0.9853 1.7904 0.9137 0.0528 22
0.9839 1.5793 0.9234 0.0532 22
0.9833 1.6459 0.9214 0.0546 22
Notes: independent variable = rubber produced in metric tonnes, K • area planted in hectares, L • number of workers employed. R-square • coefficient of multiple determination, D.W. = Durbin-Watson statistics, SER = standard error of regression, d.f. = degrees of freedom. All other variables are as previously defined in the text. ***Statistically significant at the one percent level ** Statistically significant at the five percent level * Statistically significant at the ten percent leveL Figures within brackets are 't-statistics\
UZAFAII SHAH HAB1BULLAH
m
TABLE 2 Regression coefficients and related statistics for the oil palm sector Model I
Model II
Model III
Model IV
Constant
-0.63747 (-2.0214)*
-0.88306 (-6.4452)***
-0.57959 (-5.3551)***
-0.76110 (-6.2661)***
K
1.1082 (6.1299)***
0.87807 (4.1033)***
0.47125 (2.1512)**
0.38522 (1.6292)
L
0.20155 (0.69516)
0.04225 (0.14554)
0.44760 (1.6461)
0.54764 (1.923?)**
73
0.64731 (5.0362)***
O
(M2/CPI)
o
0.64757 (6.2602)***
(M3/CPI)
0.64621 (5.7361)*** 0.9532 2.2608 0.7279 0.0934 23
0.9965 1.8296 0.0002 0.0818 22
0.9968 1.9615 0.0904 0.0694 22
7*
0.9958 2.0087 0.1822 0.0707 22
Notes: independent variable = palm oil produced in metric tonnes, K = area planted in hectares, L = number of workers employed. All other variables are as previously defined in Table 1. ***Statitically significant at the one percent level ** Statistically significant at the five percent level * Statistically significant at the ten percent level Figures within brackets are 4t-statistics\
i O
OF A DEVELOP
R-square D.W. Rho SER d.f.
U 2
ODUCT][ON FU!
(Ml/CPI)
MONEY 13ALAN<
m
Variables4
TABLE 3 Regression coefficients and related statistics for the tea sector
o m
Model I
Model II
Model III
Model IV
Constant
1L589 (7.2553)***
4.7573 (2.0183)*
3.3318 (1.3913)
3.3404 (1.3901)
K
-0.25495 (-1.2188)
0.45350 (1.6811)
0.60785 (2.2382)**
0.59621 (2.2078)**
L
-0.00074 (-0.00748)
0.17730 (1.5822)
0.24898 (2.1930)**
0.27182 (2.3324)**
(Ml/CPI)
0.30120 (3.0286)***
55
o
2 JZAF
I
Vaiiablesa
> 73
(M2/CPI)
Notes
0.25442 (3.6254)*** 0.9920 1.9217 0.5827 0.0659 23
0.9940 1.6150 0.8479 0.0568 22
0.9945 1.6227 0.8545 0.0538 22
0.9949 1.6863 0.8290 0.0535 22
independent variable = green leaves harvested in metric tonnes, K • area planted in hectares, L • number of workers employed. All other variables are as previously in Table 1. ***Statistically significant at the one percent level **Statistically significant at the five percent level * Statistically significant at the ten percent level Figures within brackets are 4t-statistics\
HAB IBU;
(M3/CPI) R-square D.W. Rho SER d.f.
X X
0.26929 (3.5580)***
> X
TABLE 4 Regression coefficients and related statistics for coconut sector
o Model 1
Model II
Model III
Model IV
Constant
7.4170 (16.718)***
7.7655 (14.930)***
7.7449 (15.123)***
7.7665 (15.170)***
K
0.39741 (8.2316)***
0.37920 (7.6328)***
0.38266 (7.7840)***
0.38346 (7.8639)***
L
0.26322 (6.1967)***
0.21791 (3.8920)***
0.20574 (3.2857)***
0.19702 (3.0082)***
m
I
(Ml/CPI)
-0.02936 (1.2248)
(M2/CPI)
-0.02261 (-1.2376)
o
-0.02385 (-1.3131)
(M3/CPI)
o R-square D.W. Rho SER d.f.
0.9912 1.8403 -0.1891 0.0452 23
0.9926 1.8657 -0.1983 0.0447 22
0.9919 1.8723 -0.1900 0.0447 22
0.9917 1.8731 -0.1876 0.0445 22
Notes: independent variable = number of nuts collected, K = area planted in hectares, L • number of workers employed. AU other variables are as previously defined in Table 1. ***Statistically significant at the one percent level **Statistically significant at the five percent level * Statistically significant at the ten percent level Figures within brackets are 't-statistics\
BALANCESIN THE PR ODUCTION FUNCTIOl^J OF A DEVELO
Variables'a
I O
TABLE 5 Regression coefficients and related statistics for the forestry sector
Model I
Model II
Model III
Model IV
Constant
-1.5132 (-3.0107)***
4.6087 (3.1831)***
4.7242 (3.1839)***
4.5109 (3.2423)***
K
0.09738 (2.8589)***
0.08656 (1.8339)*
0.09296 (2.0100)*
0.09241 (2.0041)*
m
L
0.87813 (14.668)***
0.34496 (1.6896)
0.35826 (1.7803)*
0.38023 (1.9728)*
z
(Ml/CPI)
0.20015 (1.6050) 0.9744 1.9539 0.3019 0.0738 20
0.9864 1.8642 0.6411 0.0875 19
0.9830 1.8078 0.5258 0.0882 19
0.9821 1.7935 0.5092 0.0886 19
Notes: independent variable • sawlogs produced in cubic metres, K = area logged in hectares, L = number of workers employed. All other variables are as previously defined in Table 1. •••Statistically significant at the one percent level ••Statistically significant at the five percent level •Statistically significant at the ten percent level Figures within brackets are 't-statistics*.
H HAB(BULLA
0.22599 (1.6515)
(M3/CPI) R-square D.W. Rho SER d.f.
6 5
0.36523 (1.7751)*
(M2/CPI)
tfUZAF
Variables*
X
REAL MONEY BALANCES IN THE PRODUCTION FUNCTION OF A DEVELOPING ECONOMY TABLE 6 Regression coefficients and related statistics for the paddy sector Variables11
Model I
Model II
Model III
Model IV
Constant
-1.5855 (-1.3284)
-0.67679 (-1.6124)
-0.57526 (-1.2945)
-0.64135 (-1.3483)
K
1.4495 (7.5944)***
1.1644 (16.085)***
1.1643 (15.322)***
1.1691 (14.358)***
(Ml/CPI)
0.19926 (9.6571)*** 0.13893 (9.3078)***
(M2/CPI)
0.13243 (8.6894)***
(M3/CPI) R-square D.W. Rho SER d.f.
0.9368 2.4801 0.9349 0.0689 23
0.9640 1.7643 -0.0924 0.0532 22
0.9650 1.8515 -0.0250 0.0525 22
0.9633 1.8841 0.0252 0.0537 22
Notes: independent variable = paddy harvested in metric tonnes, K • area planted in hectares. All other variables are as previously defined in Table 1. •••Statistically significant at the one percent level **Statistically significant at the five percent level •Statistically significant at the ten percent level Figures within brackets are ^-statistics*.
TABLE 7 Regression coefficients and related statistics for the fishery sector Variables41
Model I
Model II
Model III
Model IV
Constant
6.0595 (2.6020)**
1.9762 (0.92056)
1.2369 (0.65579)
0.73393 (0.40111)
L
0.04942 (0.09262)
0.51584 (0.93887)
0.72740 (1.4968)
0.83075 (1.7524)*
0.51874 (3.2826)***
(Ml/CPI)
0.37600 (4.3628)***
(M2/CPI)
0.35062 (4.4198)***
(M3/CPI) R-square D.W. Rho SER d.f.
0.9284 1.7104 0.8939 0.1254 23
0.9270 1.7674 0.6002 0.1296 22
0.9326 1.6271 0.4990 0.1245 22
0.9316 1.5915 0.4775 0.1254 22
Notes: independent variable = fish landing in metric tonnes, L = number of fishermen. All other variables are as previously defined in Table 1. * * * Statistically significant at the one percent level * * Statistically significant at the five percent level •Statistically significant at the ten percent level Figures within brackets are *t-statUtics\ PERTANIKA VOL. 11 NO. 3, 1988
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REAL MONEY BALANCES IN THE PRODUCTION FUNCTION OF A DEVELOPING ECONOMY conclude that real money balances are productive inputs. IV. CONCLUSION The objective of this study has been two-fold. Firstly, to formulate the aggregate production function model with respect to the agriculture sector in Malaysia. Secondly, to determine whether the data support the hypothesis that real money balances influence production. The factors of production considered in this study were capital, labour and real money balances. The production of each agricultural subsector; rubber, oil palm, tea, coconut, forestry, paddy and fishery was considered in this study. With caution, the results of the individual sectors suggest that except for the coconut subsector, real money balances play an important role as a productive input. REFERENCES APOSTOLAKIS, B. (1983) : Money Balances as Factor Inputs: An Empirical Evidence Based on Translog Specification, Europ. Econ. R. 23: 149-160. BAILEY, M.J. (1971): National Income and the Price Level, New York: McGraw-Hill Book Company. BEN-ZION, U. and V.W. RUTTAN. (1975): Money in the Production Function: An Interpretation of Empirical Results, The R. Econ. Statist. 57: 246247. BOOTH, A. and R.M. SUNDRUM. (1984): Labour Absorption in Agriculture: Theoretical Analysis and Empirical Investigations, Delhi: Oxford University Press. BOYES, W.J. and D.C. KAVANAUGH. (1979): Money and Production Function: A Test of Specification Bias, TheR. Econ. Statist. 61: 442-446. FINNERTY, J.D. (1980): Real Money Balances and the Firm's Production Function, / Money, Cred Bank. 12(4): 666-671. FISCHER, S. (1974): Money and the Production Function, Econ. Inquiry. 12(4): 517-533. FRIEDMAN, M. (1959): The Demand for Money: Some Theoretical and Empirical Results, /. Polit. Econ. 67: 327-351. (1969): The Optimum Quantity of Money and Other Essays, Chicago: Aldine Publishing Company. JOHNSON, H.G. (1969): Inside Money, Outside Money, Income, Wealth and Welfare in Monetary Theory, J. Money Cred. Bank. 1: 30-45. KHAN, A.H. and A. AHMAD. (1985): Real Money Balances in the Production Function of a Developing Country, TheR. Econ. Statist. 67: 336-340. 460
KHAN, M.S. and PJ.K. KOURI. (1975): Real Money Balances As a Factor of Production: A Comment, TheR. Econ. Statist. 57: 244-246. LEVHARI, D. and D. PATINKIN. (1968): The Role of Money in a Simple Growth Model, Amer. Econ, R. 58: 713-754. Malaysia, Bank Negara Malaysia: Quarterly Economic Bulletin, various issues. Department of Statistics: Oil Palm, Cocoa, Coconut and Tea Statistics, various issues. Department of Statistics: Rubber Statistics Handbook, various issues. Ministry of Agriculture: Annual Fishery Statistics, various issues. Ministry of Agriculture: Forest Peninsula Malaysia, various issues.
Statistics
Ministry of Agriculture: Paddy Statistics, various issues. MORONEY, J.R. (1972): The Current State of Money and Production Theory, Amer. Econ. R. 62: 335343. NADIRI, M.L (1969): The Determinants of Real Cash Balances in the U.S. Total Manufacturing Sector, Quart. J. Econ. 83: 173-196. NICOLLI, A. (1975): Real Money Balances: An Omitted Variable From The Production Function? A Comment, TheR. Econ. Statist. 57: 241-243. PRAIS, Z. (1975a): Real Money Balances as Variable in the Production Function, The R. Econ. Statist. 57: 243-244. (1975b): Real Money Balances as Variable in the Production Function, /. Money, Cred. Bank. 7(4): 535-540. SHORT, E.D. (1979): A New Look at Real Money Balances as a Variable in the Production Function, / Money Cred. Bank. 11(3): 326-339. SIMOS, E.O. (1981): Real Money Balances as a Productive Input,/ Monet. Econ. 7: 207-225. SINAI, A. and H.H. STOKES (1972): Real Money balances: An Omitted Variable from the Production Function? TheR. Econ. Statist. 54: 290-296. STRONG, S. and H.A.J. SUHAILA. (1987): Adjustment Cost Model for Labour Wage in the Oil Palm Estate Sector, Kalian EkonomiMalaysia 24(1): 36-45. SUBRAHMANYAM, G. (1980): Real Money Balances as a Factor of Production: Some New Evidence, The R. Econ. Statist. 62: 280-285. YOU, J.S. (1981): Money, Technology and the Production Function: An Empirical Study, Canad. J. Econ. 14(3): 515-524.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 26 September, 1987)
Pcrtanika 11(3), 461-468 (1988)
Some Stream Water Quality Characteristics of Two Small Logged Over Watersheds in Selangor LAI F.S. and NORAJIKI AJ.* Faculty of Forestry Universiti Pertanian Malaysia 43400 Serdang, Selangor, Malaysia. Keywords: Watersheds; water temperature; dissolved oxygen; pH; conductivity; suspended solids; dissolved solids. ABSTRAK Satu kajian berkenaan dengan beberapa parameter kualiti air sungai telah dijalankan di dua kawasan legeh hutan yang telah dibalak. Kajian ini mengambil masa selama enam bulan iaitu dari Ogos 1983 hingga Januari 1984 dimana pensampelan air telah dibuat diantara jam 1400 dan 1500. Keputusan yang diperolehi menunjukkan bahawa kesemua parameter yang dikaji itu lebih berbedza untuk kawasan legeh WA yang lebih dibalak berbanding dengan WB. Nilai purata parameter kualiti air sungai untuk WA dan WB adalah berikut: suhu air - 25.3 dan 25.2°C; oksigen terlarut -6.7 dan 6.7 ppm ; pH - 5.55 dan 5.25; kekonduksian -16.0 dan 11.3 [itnhos/cm; takat pepejal ampaian - 7 dan 22 mg/l; dan takat pepejal terlarut - 31 dan 36 mg/l. Keputusan yang diperolehi mencadangkan bahawa kesan operasi hutan atas kualiti air adalah berbedza. Walau bagaimanapun, kualiti air masih dalam keadaan baik dengan adanya pertimbangan keatas alam sekitar semasa operasi hutan dijalankan.
ABSTRACT A study of selected stream water quality parameters was carried out in two forested watersheds with varying degrees of disturbance. The study period of six months from August 1983 to January 1984 involved regular sampling at between 1400 and 1500 hours. Results indicate all parameters observed from the relatively more disturbed watershed WA were higher than the less disturbed basin WB. Mean values of stream water quality parameters of WA and WB respectively areas follows : water temperature - 25.3 and 25.2°C; dissolved oxygen - 6.7 and 6.4 ppm; pH - 5.55 and 5.25; conductivity 16.0 and 1L3 ymhos/cm; suspended solids - 22 and 7 mg/l and; total dissolved solids - 36 and 31 mg/l The findings suggest logging operations had varying influences on the water quality parameters. Although affected, the water quality remains good, aided by environmental considerations during logging operations.
INTRODUCTION It is often reported that logging of forests adversely affects stream water quality. By physical reasoning it is obvious that this effect is real. Unfortunately, little attention has actually been received to document the physical and chemical stream water quality from affected catchments. More
attention has been given, in the country, to quality of receiving waters from heavy polluting industries such as palm oil and rubber processing plants. The need for some comparison between basic data must be stressed in order to better understand not only variations within a particular watershed but also between watersheds. For this
Present address: Hano Forests Consultants, P.O. Box 169 Limbang, Sarawak.
LAI F.S. AND NORAJIKI A.J.
reason, selected stream water quality parameters pf temperature, dissolved oxygen (DO), pH, conductivity, total suspended solids (TSS) and total dissolved solids (TDS) of two logged over watersheds were measured and reported here. MATERIALS AND METHODS
General Description of Study Area A description of the study area is briefly outlined in this section. The geographic setting has been detailed by Lai and Samsuddin (1985). Two small watersheds demarcated, WA and WB are tributaries of Sungai Rasau, the main river draining the Air Hitam Forest Reserve (Figure 1). WA is larger in area covering 7.3 km 2 compared to 4.7 km 2 of WB.
Figure L
Location of study area.
Soil type for both watersheds are similar comprising the soil series of Serdang-Kedah and Durian association (Zainuddin, 1976). The Serdang series occurs along places downslope while that of Kedah and Durian are found on the ridges and upper slope regions. The local Alluvium Colluvium association occurs in the valley and foothill regions. 462
Prior to logging, the reserve was a lowland dipterocarp forest; with logging it was gradually replaced by secondary disturbed forest (Ali Riza, 1977). From past records, logging has been the main activity in the study area. Log extraction from the forest reserve began as early as 1930 and thereafter continued on commercial and subsistence basis up to 1983. In the harvesting, prior to this study, a crawler tractor-San Tai Wong system in which a Komatsu 16 was used. On record, WA is relatively more disturbed compared to WB; logging operations (including salvage logging) ceased in July 1983 in WA, a year after operations stopped in WB-about 35% and 13% of basin areas of WA and WB were logged respectively. Data Collection Over the study period, the stream water quality parameters were regularly sampled at between 1400 and 1500 hours at both stations. In addition, hourly sampling was also carried out for both stations on a 5-consecutive day basis to obtain any indication of diurnal fluctuations. A brief outline of methods used to record the selected water quality parameters in the study is given below. Dissolved oxygen readings were made on site using a YSI 54 oxygen meter and YSI 5739 DO probe with an accuracy of ± 0.1 ppm. Water temperature and conductivity readings were recorded on site using a YSI 33 S - C - T meter. The YSI 3000 Series probe for the instrument was submerged halfway from the water surface at the mid-stream section. Water temperature was checked and calibrated against a submerged laboratory thermometer. Sufficient time was allowed for stable readings before recordings were made. For comparison purpose, air temperature of the surroundings was measured using a laboratory thermometer well-placed in shaded areas - readings were noted at about the same time water temperature was recorded. Measurement of pH was also carried out on site using a Mini-Mite pH meter (PA-11). The electrode was placed in the same section of the stream, as the temperature/conductivity and DO probe described earlier. A one-litre plastic bottle sampler was used to collect stream water samples for suspended and dissolved solids analyses. The container was fixed with two copper tubes, one as intake and
PERTANIKA VOL. 11 NO. 3, 1988
SOME STREAM WATER QUALITY CHARACTERISTICS OF TWO SMALL LOGGED OVJER WATERSHEDS
the other as air exhaust constructed following the sampling mechanism of a point integrating sampler. The water sample collected was later analysed in the laboratory using 4 replicates of individual samples. Concentrations of total suspended and dissolved solids were determined following the procedures outlined by Wang and Ong (1978). RESULTS
\ i
Water Temperature Analysis of variance for instantaneous water temperature data showed no significant difference (P <0.05) between stations WA (x = 25.3°C) and WB (x = 25,2°C). Similarly, the range noted throughout the study period does not indicate obvious variation (Table 1) although the lowest values recorded were in the order of 25.0°C and 24.2°C for WA and WB respectively. In addition, a fair number of recordings taken during time of sampling were higher in WA than in WB. This can be attributed to the relatively exposed condition in certain stream reaches in WA mainly resulting from logging; direct sunlight provides the major energy source for heating the exposed streams. In small shaded streams, temperature changes induced by logging is directly proportional to the amount of exposure given the stream surface (Brown, 1974). It was not possible to ascertain, however, if the temperature has an indirect relationship with discharge and by how much from the two basins since most of the recordings were done during low flow conditions. Although monitored separately, the diurnal variation of the water temperature for both WA and WB showed similar patterns (Figures 2a and 2b). The maximum water temperature occurred in the late afternoon (approx. 1600 and 1800 hours) while minimum readings were recorded during early parts of the day (approx. 700 to 1000 hours) after the highest and lowest air temperatures had been attained respectively. The pattern is characteristic of tropical streams (Edinger etal, 1968) because of shortwave solar radiation influence which causes the maximum equilibrium temperature to occur near noon in the diurnal cycle while bottom heating gives a higher value at the end of the day (Geijskes, 1942).
I'
'
3411
Figure 2a:
5 consecutive day air and water temperature -WA
I t »• TIM ( i n )
Figure 2b.
5 consecutive day air and water temperature -WB.
Throughout the diurnal water temperature observation, minor variations between hourly readings for both watersheds were noted. That small temperature variations do occur is because the amplitude of the actual water is only a small portion of the diurnal amplitude of the water equilibrium temperature (Edinger etal, 1968), Dissolved Oxygen Dissolved oxygen range for the two watersheds are summarized in Table 1. The results for WA are significantly higher (P <0.01) than WB although the mean values are 6.7 and 6.4 ppm respectively.
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LAI F.S. AND NORAJIKI A.J. TABLE 1 Selected water quality characteristics for WA and WB (August 1983 to January 1984) Watershed A
Watershed B Average
Min
Max
Standard Deviation
0.4
25.2
24.2
26.2
0.58
7.1
0.4
6.4
5.9
6.9
0.31
5.70
0.09
5.25
5.06
5.48
0.11
Parameter
Average
Min
Max
Standard Deviation
Temperature
25.3
25.0
26.2
Dissolved Oxygen (ppm)
6.7
6.0
PH
5.55
5,25
Conductivity (//mhos/cm)
16
9
30
5.5
Total Suspended Solids (mg/1)
22
4
56
Total Dissolved Solids (mg/1)
36
18
77
11.3
5
28
5.22
10.7
7
0
23
5.26
13.4
31
18
42
6.30
Note : Number of observations for both watersheds - 44
It is suggested that the relatively higher flow velocities measured with a current meter at WA has some influence on the aeration of the stream. Logging can affect the amount of oxygen in streams in many ways: increased water temperature (thereby decreasing DO) as a result of clear cutting along streams - this is not clear in the study since the water temperature range data appear similar and are not statistically significant; and increase in BOD because of higher microorganism activity due to accumulation of debris. Recent logging in WA has not resulted in a substantial decrease in DO which can also be attributed to low logging intensity and the fact that no clearcutting along river banks was permitted. Diurnal DO decline towards the evening hours was observed especially in WA (Figure 3a). It is not clear, however, why this trend is such. The 5-day diurnal DO range is slightly higher for WB (0.9 ppm) than WA (0.5 ppm) which can be attributed to a number of factors. With weather fluctuations - since photosynthetic oxygen production by aquatic plants is propor464
tionl to light intensity (Krenkel and Ruane 1979), DO can be reduced compared with normal 'bright' days. Additionally, storm flows can also affect DO concentrations due largely to dilution and transportation of large amounts of suspended matter - a drop in DO level from 6.8 to 6.0 ppm occurred just before the storm hydrograph peaked and increased steadily after that to assume the "normal" diurnal fluctuation (Fig. 3b). PH pH recorded for WB is significantly lower than WA (P<0.01) (Table 1). The difference can be explained by the higher organic matter concentration from WB reported by Shamsuddin (1984) in a separate study. Organic matter has an important influence on the pH concentration; higher organic matter content usually leads to a higher occurrence of decomposition, thus affecting pH. The 5-day diurnal pH for WA and WB shows higher levels of occurrence during the day and lower levels during the night (Figures 3a and
PERTANIKA VOL. 11 NO. 3, 1988
SOME STREAM WATER QUALITY CHARACTERISTICS OF TWO SMALL LOGGED OVER WATERSHEDS
am
I
xrt*.
I
mm
Till IB) I am
Figure 2a. 5 consecutive day water quality survey - WA.
tivity level. The findings of this study are in agreement with those reported by Hatch and Shea (1977), Thanasilungkoon and Ruangpanit (1981) where higher conductivity levels were obtained from disturbed forested watersheds compared with undisturbed ones. Conductivity levels remained consistent over the 5-day diurnal recordings for WA. For WB however, an increase from 12 to 18 /imhos/cm at 25°C was recorded during the occurrence of a storm of which higher readings were gradually registered initially with a rise in water level. For 18 hours, even after peak discharge had occurred and the stormflow had begun to recede, conductivity readings remained stable. Total Suspended and Dissolved Solids
I
Ii-l
Figure 3b.
!«)
5 consecutive day water quality survey- WB.
3b) that pH remains almost consistent in terms of daily variation for both watersheds until the occurrence of stormflow This suggests that the dependence on the hydrologic response is more dominant. Conductivity Relatively higher conductivity levels were observed in WA and significantly different (P<0.01) compared with WB (Table 1). Because of recent exposure and disturbance resulting from logging roads and skid trails, the role of surface wash becomes dominant. This can lead to an increase in dissolved solids load in the stream. Consequent to which, the increase in dissolved solids improves the ionic strength of the water, thereby raising the conduc-
Table 1 shows the data range obtained for both watersheds. Statistically, TSS concentrations are significantly different (P <0.01) between WA and WB. This is expected because of the more recent disturbance in WA due to logging operations in terms of soil exposure and erosion. Although the difference is significant however, a maximum of 77 mg/1 and 58 mg/1 recorded for WA and WB respectively, is not indicative of the whole TSS rating range. Most samples were, unfortunately, taken during lowflows or when stormflows had receded. As commonly reported, (e.g. Heidel, 1956; McPherson, 1971; Walling, 1977; Finlayson and Wong, 1982; Carling, 1983; Lai and Samsuddin, 1985) most of the organic and inorganic material supply would have already been "flushed" out of the system during the first and earlier storms. As a result, TSS concentration is reduced after this or even after a rapid succession of storm events. Once the available material supply is reduced, concentrations decline even though discharge may continue to rise due chiefly to dilution. It also follows that the concentration of suspended solids will vary within a particular storm depending on the water stage, that is, rising, peak or falling. This is evident as Samsuddin (1984) reported maximum concentration of 362.2 mg/1 for the same basin during stormflows. Nevertheless, the results provide some indication of forest disturbance on TSS concentrations during receding and low flows. The variation of TDS concentration is not definite and was not significantly different
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LAI F.S. AND NORAJIKI A.J.
between watersheds. Because sampling was not intensively carried out during stormflows, it is not possible to suggest any probable TDS concentration range for the two watersheds. Much also depends on the influence of geomorphology and geology of the study area (Kunkle and Meiman 1967; Finlayson and Wong, 1982). DISCUSSION AND CONCLUSION
Although the six month study on the water quality was largely confined to low flow samples, it is clear from the general variation patterns that logging in the watersheds has resulted in varying changes insofar as the selected water quality parameters are concerned (Table 2),
The study showed no significant difference in water temperature between watersheds although a fair number of readings were higher in WB. It has been indicated by Crowther (1982) that fotest clearance would lead directly and indirectly to an increase in water temperature and temperature fluctuation. This could be mainly due to greater exposure of the streams in WB to incoming solar radiation at which point this hypothesis would have been better established with a good network of continuous temperature data loggers installed at strategic locations along streams draining the watershed. Although water temperature has a direct relationship with DO, the small effect logging
TABLE 2 Summary of selected parameter concentration range of present and previous studies in Selangor v. Description Area (Km2)
Temp (°C)
D.O. (ppm)
pH
Conductivity (/Anhos/cm)
Source
Study Area Sg. Gombak* Station I
123.3
22.5 - 25.0
24.4 - 29.0
6 . 8 6 - 7.52
24.8 - 30.5
4.78 - 6.67
6.65 - 7.35 6.70- 7.39 6.70 - 7.64 6.30-7.10 6.23 - 6.95
_
4.80 - 7.45
5.50-6.50
9.20 - 27.0
Ho (1973)
6.0-7.1 5.9 - 6.9
5.25-5.70 5.06 - 5.48
9.0 - 30.0
Present study
7.04-7.88
Station II
(total
22.2 - 24.5
7.34 - 8.00
Station III
catch-
23.8-28.5
7.25 - 7.92
Station IV
ment)
Station V
31.92 - 48.00
Bishop (1971)
26.40-33.60 2 6 . 4 0 - 34.71 25.20-36.54 35.40-50.40
Sg. Renggam** Station V
20.0 (total catchment)
Sg. Rasau WA
7.3
2 5 . 0 - 26.2
WB
4.7
24.2 - 26.2
5.0 - 28.0
Notes : *Landuse of total catchment area : forest - 56.7%; rubber - 23.7%; rice and irrigated land - 2.5%; urban and paved roads - 11.9%; tin mining - 4.9%; quarrying - 0.3% **Land use of total catchment ar&: undisturbed forest - 1%; logged over forest - 21% cleared land, urban development - 68%; industrial - 8%; oil palm - 2% 466
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SOME STREAM WATER QUALITY CHARACTERISTICS OF TWO SMALL LOGGED OVER WATERSHEDS
operations had on the former suggests that there are other factors that could in combination, result in the higher DO levels recorded for one of the two watersheds. One such factor is the higher occurrence of algae and aquatic plants observed in the stream draining WA compared to WB. As pointed out by Krenkel and Novotny (1979) the most important factor which constitutes the effect of aquatic plants and DO within a particular time of day is plant density. As suggested earlier, :he relatively higher flow velocity at the sampling site WA could also have contributed to the high DO concentrations. pH levels recorded during the study period suggest an increase after disturbance in the watersheds. WA recorded significantly higher pH values than WB although both drainage basins have similar geology and soil types. The reasons are not clear although it is probable that photosynthesis from the fairly abundant aquatic plants in WA using CO2 can raise the pH level (Thanasilungkoon and Ruangpanit, 1981). It is also likely that pH is largely influenced by hydrological conditions. Ramberg (1981) observed a significant increase in pH after improved drainage in coniferous forest due to lesser acidification of groundwater although this observation was made in temperate conditions, it is interesting to note whether the presence of a small well-drained oil palm stand in the upper catchment of WA could also have affected pH in the same way. The dominant role of surface wash after logging can affect the water chemistry in some ways. In this study, for instance, conductivity, TSS, and TDS were found to have been affected in varying degrees. Significantly higher conductivity levels sampled at disturbed basins compared with undisturbed ones have been reported by Hatch and Shea (1977), Thanasilungkoon and Ruangpanit (1981). Although statistical tests did not show any significant difference in dissolved solids between watersheds, there were indications that TDS can be higher in concentrations for WA. This is also evident from readings obtained for a stormflow on 12/12/1983 during the 5-day survey (Fig. 3b) - conductivity increased from 12 to 18 jLtmhos/cm. It was unfortunate, however, that only a small discharge range was sampled during the entire study preventing an otherwise detail data analysis and interpretation. Nevertheless, data variation for the parameters reflect
much dependence on hydrological conditions. The stream water quality of WA and WB has remained good after logging. As a comparison, studies on water quality conducted by Bishop (1971) and Ho (1973) in Selangor also showed variations for selected parameters (Table 2). Land use in their studies were more diverse, thereby explaining the wide range recorded. For example, temperature varied from 22.5 to 25.0°C in the upper catchment which is 99% forest and 24.8 to 30.5°C in the lower catchment of Sg. Gombak the results suggest marked temperature changes compared to the present study. Additionally, DO range also appears wide from 4.78 to 6.6 ppm for Sg. Gombak, 4.80 to 7.45 ppm for Sg. Renggam compared with 6.0 to 7.1 and 5.9 to 6.9 ppm for WA and WB respectively. The comparison largely indicates the important influence of land use on the selected water quality parameters. Land development including urbanisation and industrialisation are frequently considered to have drastic environmental consequences. Essentially, the concentration variation reflects a change to the previous ecosystem which almost always results in inadvertent deterioration of water quality provided adequate measures are undetaken to minimise environmental damage. Logging can affect water quality as suggested by the results in the study. In contrast, for undisturbed watersheds, the constancy and systematic variations in the water chemistry is influenced by various geological and biological processes. It must be noted, however, that environmental considerations were observed during logging such as the use of previous logging roads, avoidance of stream crossings and cutting along stream banks. Coupled with low timber production, i.e. 14-17 m 3 /ha, the impact on the selected water quality parameters can be considered minimal. However, it is not known if the impact can be more serious during logging operations. Finally, despite the short period of study, the results provide some indication of stream recovery since logging in both watersheds were carried out at different times. ACKNOWLEDGEMENTS
The authors wish to thank the Dean of the Faculty of Forestry, Universiti Pertanian Malaysia for facilities, Saad Nyan for technical assistance, Tuan Hj. Wan Mamat Saufi Wan Yusuf, Encik Abdul Kadir Jaafar and Encik Bashri Razali for their kind
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LAI F.S. AND NORAJIKI A.J.
cooperation. The authors are also grateful for the financial assistance granted by Universiti Pertanian Malaysia. REFERENCES ALI RIZA (1977): A long range development plan for main access road of Air Hitam Forest. Unpublished B. Sc. (Forestry) Project. Univ. Pertanian Malaysia, Serdang. BISHOP, J.E. (1971): Limnology of a small Malayan river: Sg. Gombak. Unpublished Ph.D thesis. Univ. Malaya. Kuala Lumpur. BROWN, G.W. (1974): The impact of timber harvest on soil and water resources. School of Forestry, Oregon St. Univ. Oregon. CARLING, P.A. (1983). Paniculate dynamics, dissolved and total load, in two small basins, northern Pennines,UK. Journ. Hydrol Sc., 28(3): 355-375. CROWTHER, J. (1982): The thermal characteristics of some West Malaysian rivers. Mai. Nat. Journ. Vol 35: 99-108. EDINGER, J.E., D.W. DUTTWEILER and J.C GEYER. (1968): The response of water temperatures to meteorologic conditions. Wat. Resour. Res. 4(5): 1137-1144, EDINGTON, J.M. (1965): Some observations on stream temperature. Oikos 15: 263-273. FINLAYSON, B.L. and N.R. WONG. (1982): Storm runoff and water quality in an undisturbed forested catchment in Victoria. Aust. For. Res. 12: 303-315. GEIJSKES, D.C. (1942): Observations on temperature in a tropical xivti.Ecology. 23(1): 106-110. HATCH, A.B. and S.R. SHEA. (1977): Water quality in allan road Catchment, Western Australia. Res. Pap. No. 30. For Dept West. Aust. HEIDEL, S.G. (1956): The progressive lag of sediment concentration with flood waves. Trans. Am. Geophy, Union. 37(1): 56-65. HO, S.C. (1973): The ecology of a lowland stream: Sg. Renggam with special references to water pollution. Unpublished M. Sc. thesis. Univ. Malaya. Kuala Lumpur.
468
KRENKEL, P.A. and V. NOVOTNY. (1979): River water quality model construction: 17/1-17/22 in Modelling of rivers (Shen, H.W., Ed.) John Wiley & Sons. N. York. KRENKEL, P.A. and RUANE R.J. (1979): Basic approach to water quality modelling: 18/1-18/40 in Modelling of Rivers (Shen, H.W., Ed.) John Wiley & Sons. N. York. KUNKLE S.H. and J.R. MEIMEN. (1967): Water quality of mountain in watersheds. Hydro. Pap. No. 21. Colorado St. Univ. U.S.A. LAI, F.S. and M. SAMSUDDIN. (1985): Suspended and dissolved sediment concentrations of two disturbed lowland forested watersheds in Air Hitam Forest Reserve, Selangor. Pertanika 8(1): 115-122. McPHERSON, H.J. (1971): Dissolved, suspended and bed load movement patterns Jour Hydrol 12: 221-233.
RAMBERG, 1. (1981): Increase in stream pH after a forest drainage, Ambio. 10(1): 34-35. SAMSUDDIN, M. (1984): Suspended and dissolved sediment concentration of two disturbed forested watersheds in Air Hitam Forest Reserve, Selangor. Unpublished B. Sc. (Forestry) Project. Univ. Pertanian Malaysia, Serdang. THANASILUNGKOON, C. and N. RUANGPANIT. (1981): Some physical and chemical aspects of water quality under various land use of Sakaerat Environmental Research Station. For. Res. Bjill. No. 81 Kasetsart Univ. Thailand. WALLING, D.E. (1977): Assessing the accuracy of suspended sediment rating curves for a small basin. Wat. Resour. Res. 13(3): 531-538. WANG, C.W. and K.C ONG. (1978): Standard manual of chemical and biological examinations for the wastewater project network. Wastewater Reclamation Project (Mai.). ZAINUDDIN, M.A. (1976): Long range development plan for road network and a detailed plan for the main access road of Air Hitam Forest. Unpublished B. Sc. (Forestry) Project. Univ. Pertanian Malaysia, Serdang.
PERTANIKA VOL. 11 NO. 3, 1988
(Received 3 December 1987)
Pertanika 11(3), 469-474(1988)
Kesan Saliniti dan Keamatan Cahaya ke atas Pertumbuhan Chlorella virginica Effect of Salinity and Light Intensity on the Growth of Chlorella virginica ABDULLAH ZAINI ALIAS Pusat Perikanan dan Sains Samudra Universiti Pertanian Malaysia Mengabang Telipot, 21030 Kuala Terengganu, Malaysia Kata punca: Saliniti; keamatan cahaya; Chlorella virginica ABSTRAK Satu kajian mengenai kesan saliniti dan keamatan cahaya ke atas tumbesaran Chlorella virginica telah dijalankan dalam bilik kawalan sekitaran pada suhu 22 ± 2?C. Saliniti yang ditetapkan ialah 15ppt, 20pptf 25ppt dan 30ppt dan keamatan cahaya pula pada 1140 lux, 2260 lux dan 3040 lux. Kesan saliniti dan keamatan cahaya didapati mempunyai keertian berbeza (P <0.01 - 0.05). Saliniti optimum untuk tumbesaran Chlorella virginica ialah 15 ppt dan keamatan cahaya optimum pada 3040 lux. Interaksi di antara saliniti dengan keamatan cahaya juga mempunyai keertian berbeza (P <0.01 - 0.05) di mana tumbesaran maksimum (39375 sel/jjl) diperolehi pada saliniti 15 ppt dan keamatan cahaya 3040 lux. ABSTRACT A study on the effects of salinity and light intensity on the growth of Chlorella virginica was conducted in an environmental control room at a temperature of 22 ± 2°C Salinities were set at 15 ppt, 20 ppt, 25 ppt and 30 ppt while light intensities were at 1140 lux, 2260 lux and 3040 lux. The effect of salinity and light intensity was found to be significantly different (P <0.01 - 0.05). Optimum salinity for the growth of Chlorella virginica was at 15 ppt whilst optimum light intensity was at 3040 lux. Interaction between salinity and light intensity was also significantly different (P K0.01 - 0.05) whereby maximum growth (39375 cells/yi) was obtained at a salinity of 15 ppt and light intensity of 3040 lux. PENGENALAN Alga merupakail makanan yang penting untuk menternak invertebrat dan ikan (Helm et aL, 1973; Walne, 1974; Trotta, 1981; Laing dan Helm, 1981). Bekalan alga diperlukan segera secara berterusan untuk mendapatkan pertumbuhan atau pengeluaran yang baik (Spectorova et al.> 1981; Rodhouse et at., 1983; Juario dan Storch, 1984). Chlorella merupakan alga yang mempunyai dinding sel yang tebal dan tegar (Howell, 1979). Walau bagaimanapun ia penting sebagai makanan kepada rotifer yang mana seterusnya menjadi makanan larva ikan (Jurio dan Storch, 1984; Liao, 1975; Gatesoupe dan Luquet; 1981). Secara tidak langsung Chlorella sp. memainkan peranan
penting sebagai sumber makanan di samping alga lain seperti Tetraselmis sp. dan Isochrysis galbana yang boleh dimakan secara langsung, Di dalam kultur alga, faktor-faktor sekitaran seperti suhu, pH, saliniti, kepekatan karbon dioksida keamatan cahaya, mineral dan juga nutrien organik mestilah dikawal dengan baik untuk mendapatkan hasil yang tinggi (Trotta, 1981; Laing dan Helm, 1981; Fabregas et ah, 1984). Keamatan cahaya merupakan faktor yang penting di mana dalam kultur alga tanpa cahaya tiruan, nilai ketumpatan maksimum amatlah sukar untuk dicapai (Goldman, 1980). Menurut Presscott (1981) cahaya amat penting untuk pengoksidanan dan fotosintesis. Laing (1985) mendapati kultur Skeletonema sp. dipengaruhi oleh
ABDULLAH ZAIN1 ALIAS
cahaya di mana hasil meningkat sebanyak 60% jika cahaya ditambah keamatannya daripada 28,000 lux kepada 40,000 lux. Air laut mempunyai tahap kemasinan yang tinggi. Oleh itu ianya mempunyai keupayaan osmotik yang tinggi. Proses osmoregulasi alga adalah berbeza mengikut spesis (Greenway dan Setter, 1979). Julat saliniti yang optimum untuk Isochrysis galbana dan Tetraselmis suecica adalah 15 ppt hingga 25 ppt dan 25 ppt hingga 30 ppt masing-masing (Laing dan Utting, 1980). Laing (1985) pula mendapati Chaetoceros calcitrans berkembang dengan baik pada julat saliniti 15 ppt hingga 20 ppt. Memandangkan kepentingan yang terdapat pada Chlorella virginica, kajian kesan cahaya dan saliniti ke atas pertumbuhan Chlorella virginica telah dijalankan di dalam bilik kawalan sekitaran pada suhu 22 ± 2°C. TATACARA DAN PERALATAN
Stok Chlorella virginica yang mempunyai kepadatan lebih kurang 18,000 sel//il dimasukkan ke dalam kelalang kultur 500 ml (mengandungi 250 ml air pada saliniti 15 ppt, 20 ppt dan 30 ppt; dibajakan dengan medium (Conway). Kelalang ditutup dengan kapas bagi menghalang kemasukan bendasing dari luar. Pengudaraan diberikan pada tiaptiap kelalang dengan menggunakan pengudara. Air terlebih dahulu ditapis dengan menggunakan penapis kartus pita berukuran 1 mikron dan disteril dengan menggunakan ultra lembayung dan ozon selama lebih kurang 15 minit. Kultur ini kemudiannya didedahkan pada keamatan cahaya 1140 lux (sebuah lampu kalimatang), 2260 lux (dua buah lampu kalimatang) dan 3040 lux (tiga lampu kalimatang). Keamatan cahaya ini ditentukan dengan menggunakan salinometer optikal. Kajian dijalankan dalam bilik kawalan sekitaran pada suhu 22 ± 2°C, menggunakan dua replikat bagi setiap rawatan. Pertumbuhan sel dikira setiap 24 jam dengan menggunakan himositometer. Jika sekiranya keadaan sel terlalu pekat, pencairan dilakukan untuk raemudahkan pengiraan. Pengiraan dijalankan sehingga pertumbuhan mencapai ketumpatan maksimum. Data-data yang didapati dianalisis dengan menggunakan analisis varian dua hala (Zar, 1974) dengan perbezaan min diuji dengan menggunakan Duncan's New Multiple Range Test (Duncan, 1955). 470
KEPUTUSAN DAN PERANCANGAN
Saliniti Daripada Jadual 1, saliniti pada keseluruhannya menunjukkan keertian berbeza (P < 0.01-0.05) kecuali pada hari pertama (P > 0.05) di mana bilangan sel bagi saliniti 15 ppt, 20 ppt, 25 ppt dan 30 ppt adalah 2358 sel//il, 2175 sel/jul, 2550 sel//Ld dan 2424 sel/jul masing-masing. Ini bermakna di peringkat awal kesan saliniti tidak menunjukkan perbezaan yang ketara. Keputusan ini sama dengan yang diperolehi oleh Laing (1985) yang mendapati saliniti mempunyai kesan terhadap pertumbuhan Tetraselmis suecica. Selain dari itu, Febregas et al. (1984) juga mendapati bahawa saliniti memberikan kesan yang tinggi terhadap pertumbuhan Tetraselmis suecica di dalam perbandingan hasil jisim yang dicapai pada fasa yang tetap. Pada saliniti 30 ppt tidak ada keertian berbeza (P > 0.05) kecuali pada hari keempat (P < 0.05) dengan saliniti 25 ppt (Jadual 1). Pada saliniti 25 ppt terdapat keertian berbeza (P < 0.05) jika dibandingkan dengan saliniti 20 ppt pada hari kelima, keenam dan ketujuh. Saliniti 20 ppt pula menunjukkan keertian berbeza (P < 0.05) dengan saliniti 15 ppt (Jadual 1) melainkan pada hari ketiga, keempat dan keenam. Bagi saliniti 15 ppt ianya mencapai kepadatan sel yang tertinggi antara saliniti yang lain (Rajah 1) di mana nilai maksimumnya adalah lebih kurang 25,400 sel//il pada hari keempat. Nilai maksimum bagi saliniti 20 ppt terdapat pada hari kelima (lebih kurang 17,400 sel//il), saliniti 25 ppt pada hari keempat (lebih kurang 13,400 sel//il) dan saliniti 30 ppt pula pada hari kelima (lebih kurang 8,800 sel/jtd). Pada keseluruhannya kepadatan yang paling tinggi ialah pada saliniti 15 ppt di hari keempat. Menurut Greenway dan Setter (1979), air laut mempunyai keupayaan osmotik yang rendah sedangkan dalam keadaan osmotik yang tinggi, pertumbuhan Chlorella sp. adalah baik. Ini jelas menunjukkan pertumbuhan Chlorella sp. adalah baik pada saliniti yang rendah berbanding dengan saliniti yang tinggi kerana semakin tinggi kadar kemasinan, semakin rendah pula keupayaan osmotik. Keamatan Cahaya Ketiga-tiga keamatan cahaya yang diberikan (1140 lux, 2260 lux dan 3040 lux) menunjukkan
PERTANIKA VOL. 11 NO. 3, 1988
KESAN SALINITI DAN KEAMATAN CAHAYA KE ATAS PERTUMBUHAN CHLORELLA VIRGINICA JADUAL 1 Min bilangan sel/ul setiap hari bagi Chlorella virginica yang dhikur pada keamatan £ahaya 1140 Lux, 2260 Lux dan 3040 Lux pada saliniti 15 ppt, 20 ppt, 25 ppt dan 30 ppt. \ ^
Perkaia
No.
Bilangan sel/)LQ untuk hari 1
2
3
4
5
6
7
1763ab 1387a 1400a 1363a 2513cd 2413cd 2738cd 2188bc 2800d 2725cd 3513e 3723e
2725a 3088a 1875a 2325a 7888cd 7713cd 7013c 4713b 13388e 8250cd 8663d 7988cd
4988bc 2800ab 863a. 538a 14550e 13213e 10850d 5950c 18225g 1725 Ofg 15575ef 15100ef
12750b 2355a 250a 775a 24129c 17650b 15675b 1500a 39375d 28875cd 24375cd 18000b
10375b 925a 192a 150a 285OOe 20250d 13375c 225a 27 5 OOe 3125Of 12875bc 26125e
1225a 635a 343a 266a 29125d 23513bc 3625a 188a 31250d 28375cd 19375b 22750b
8000b 513a 838a 405a 28875e 27750de 1213a 1101a 3l875e 22750cd 19000c 22250cd
1478x 246 3y 3190z
2503x 6831y 9572z
2297x 11141y 16538z
4033x 14739y 27656z
291 Ox 15588y 24438z
617x 14113y 25438z
2439x 14735y 23969z
2358 2175 2550 2424
8000m 63502 5850£ 5008k
12588m 25418m 22125m 11088xm 16293xm 174752 9096kE 134332 8814k 8833k 7196k 6758k
204332 175082 7781k 7735k
22917m 170042 7017k 7919k
#* 304.9
** 385.3
** 186.3
** 236.8
** 134.4
** 28.6 ** 12.6
** 30.9 * 3.8
** 59.8
** 50.4 ** 19.8
** 50.1 ** 16.8
Rawatan A. B. C. D. E. F. G. H. I. J. K. L.
1140 Lux, 15 ppt 1140 Lux, 20 ppt 1140 Lux, 25 ppt 1140 Lux, 30 ppt 2260 Lux, 15 ppt 2260 Lux, 20 ppt 2260 Lux, 25 ppt 2260 Lux, 30 ppt 3040 Lux, 15 ppt 3040 Lux, 20 ppt 3040 Lux, 25 ppt 3040 Lux, 30 ppt Min untuk cahaya
A. B. C.
1140 Lux 2260 Lux 3040 Lux Min untuk saliniti
A. B. C. D.
15 ppt 20 ppt 25 ppt 30 ppt Nilai untuk F
*•
A.
Cahaya
104.9
B.
Saliniti
2.6
C.
Interaksi
4.8
* 4.6
** 660.6 ** 185.2 ** 71.1
Nilai min yang diikuti oleh huiuf abjad menunjukkan tiada keertian berbeza (p > 0.05). *
Terdapat keertian berbeza pada p < 0.05.
** Terdapat keertian berbeza pada p < 0 . 0 1 .
PERTANIKA VOL. 11 NO. 3, 1988
471
ABDULLAH ZAINI ALIAS
kesan keertian yang berbeza (P < 0.01) di sepanjang kajian dijalankan (Jadual 1). Dari segi perbandingan antara ketiga-tiga keamatan cahaya itu terdapat keertian berbeza (P < 0.05) di mana keamatan cahaya 3040 lux adalah yang terbaik. Ini mungkin disebabkan oleh terdapatnya perbezaan dalam tahap keseimbangan antara fotosintesis dan pengoksidaan yang berbeza-beza bagi setiap keamatan cahaya.
Rajah 1:
Pertambahan bilangan sel bagi Chlorella virginica yang dikultur pada saliniti 15 ppt, 20 ppt, 25 ppt dan 30 ppt.
Bagi keamatan cahaya 2260 lux dan 304C lux, pertambahan bilangan sel selepas hari pertama adalah tinggi. Nilai tertinggi yang diperolehi bagi kedua-dua keamatan cahaya ini adalah lebih kurang 27,600 sel//il bagi 3040 lux (hari keempat) dan lebih kurang 15,500 sel//il bagi keamatan cahaya 2260 lux pada hari kelima (Rajah 2). Walaupun kepadatan sel bagi keamatan cahaya 2260 lux tidak memperolehi tahap tertinggi pada hari keempat, bilangan selnya nyata lebih tinggi daripada keamatan cahaya 1140 lux iaitu lebih kurang 14,700 sel/pd (2260 lux) berbanding dengan 4,000 sel/jil (1140 lux). Daripada keputusan yang didapati ini, semakin tinggi keamatan 472
cahaya yang diberikan lebih tinggi pula hasil yang diperolehi. Ini adalah bersamaan dengan kenyataan Jereos (1977) yang menyatakan antara julat 200 lux hingga 25,000 lux yang diberikan kepada Chaetoceros cakitrans keamatan cahaya 12,000 lux adalah terbaik. Oleh itu, penambahan cahaya daripada 200 lux telah memberikan hasil yang lebih baik. Menurut Laing (1985) dengan menambahkan keamatan cahaya dalam kultur Skeletonema sp. dari 4 buah lampu kalimatang (80 watt setiap satu lampu kalimatang) kepada 6 buah lampu kalimatang akan dapat menambahkan lagi hasil.
Rajah 2:
Pertambahan bilangan sel bagi Chlorella virginica yang dikultur pada keamatan cahaya 1140 lux, 2260 lux dan 3040 lux,
Interaksi antara Saliniti dan Keamatan Cahaya Interaksi antara saliniti dan keamatan cahaya mempunyai keertian berbeza (P < 0.01-0.05) sepanjang kajian dijalankan (Jadual 1). Daniel et al (1980) mendapati pertumbuhan Nannochloris oculata dipengaruhi oleh kesan interaksi antara saliniti dan keamatan cahaya. Chlorella virginica yang dikultur pada keamatan cahaya 1140 lux dan saliniti 20 ppt,
PERTANIKA VOL. 11 NO. 3, 1988
KESAN SALINITI DAN KEAMATAN CAHAYA KE ATAS PERTUMBUHAN CHLORELLA VIRGINICA
25 ppt dan 30 ppt tidak mempunyai keertian berbeza (P > 0.05) sepanjang kajian dijalankan (Jadual 1). Bagi saliniti 15 ppt dan 20 ppt, terdapat keertian berbeza (P < 0.05) pada hari keempat, kelima dan ketujuh. Kultur yang dijalankan pada keamatan cahaya 2260 lux dan saliniti 15 ppt, 20 ppt, 25 ppt dan 30 ppt, pada hari pertama tidak mempunyai keertian berbeza (P > 0.05) tetapi pada hari berikutnya terdapat beberapa perubahan (Jadual 1). Pada hari kelima kesan interaksi antara saliniti dan keamatan cahaya menunjukkan keertian berbeza (P < 0.05). Bagi keamatan cahaya 3040 lux dan saliniti 15 ppt, 20 ppt, 25 ppt dan 30 ppt juga terdapat beberapa perbezaan dan menunjukkan keertian berbeza (P < 0.05) pada hari kelima (Jadual 1). Pertumbuhan Chlorella virginica adalah yang terbaik pada keamatan cahaya yang tinggi dan saliniti yang rendah. Pertumbuhan maksimum (39375 sel/jLd) diperolehi pada saliniti 15 ppt dan keamatan cahaya 3040 lux (Jadual 1). Walau bagaimanapun, untuk memperolehi hasil yang baik, kajian mengenai faktor-faktor lain seperti suhu, pH, kepekatan karbon dioksida> mineral dan nutrien perlu dijalankan. Ini adalah berpandukan kenyataan yang dibuat oleh Trotta (1981), Laing dan Helm (1981) dan Fabregas et al (1984) yang mengatakan faktor-faktor tersebut perlu dikawal dengan baik untuk memperolehi hasil yang tinggi.
FABREGAS, J., J. ABALDE, C. •HERRERO, B. CABEZAS and M. VEIGA (1984): Growth of the marine mircoalga Tetraselmis suecica in batch cultures with different salinities and nutrient concentrations. Aquaculture, 42; 207-215.
PENGHARGAAN
LAING, I. (1985): Factors affecting the large-scale production of four species of commercially important marine algae. Aquaculture, 44: 161-166.
Pengarang ingin mengucapkan terima kasih kepada semua di Fakulti Perikanan dan Sains Samudera terutama pekerja-pekerja di Pusat Penetasan Ikan Universiti Pertanian Malaysia kerana pertolongan mereka sama ada secara langsung atau tidak dalam menjayakan kajian ini. Terima kasih istimewa diucapkan kepada Puan Siti Shapor Hj. Siraj kerana membaca manuskrip dan Puan Jamilah Abdol kerana menaip manuskrip ini. RUJUKAN
GATESOUPE, F.J. and P. LUQUET (1981): Practical diet for mass culture of the rotifier Brachionus plicitillis: application to the larva rearing of sea bass, Dicentrarchus labrax: Aquaculure, 22: 149-163. GOLDMAN, J.C. (1980): Physiological aspects in algal mass cultures, p. 343-359. In: Algae Biomass. Shelef G. and C.J. Soeder (Eds). Elsevier, North-Holland Biomedical Press, Amsterdam. 825 pp. GREENWAY, H. and T.L. SETTER (1979): Accumulation of proline and sucrose during the first hours after transfer of Chorella emersonii to high NaCl.
Aust. J. PhntPhysiol 6: 69-79. HELM, M,M., D.L. HOLLAND and R.R. STEPHENSON (1973): The effect of supplementary algal feeding on a hatchery breeding stock of Ostrea edulis L. on larval vigour. /. Mar. Biol, Assoc. U.K. 53: 673-684. HOWELL, B.R. (1979): Experiments on the rearing of larval turbot, Scopthalmus maximus L. Aquaculture, 18:218 225. JEREOS, E.C. (1977): Effect of different light intensities on the growth ot the diatom Chaetoceros calcitrans. Q. Rep. Aquacult. Dept. Southeast Asia Fish. Div. Cent, 1(3): 17-18. JUARIO, J.V. and V, STORCH (1984): Biological evaluation of phytoplankton {Chlorella sp., Tetraselmis sp. and Isochrysis galband) as food for milkfish (Chanos chanos) fry. Aquaculture, 44: 161-166.
LAING, I. and S.D. HELM (1980): Factors affecting the Semi-continuous productions of Tetraselmis suecica (Kglin) Burch. In 200 i, vessels. Aquaculture, 22; 137-148. LAING, I. and S.D. UTTING (1980): The influence of salinity on the production of %o commercially important unicellular marine algae. Aquaculture, 21:79-86. LIAO, I.C. (1975): Experiments on induced breeding of the grey mullet in Taiwan from 1963 to 1973. Aquaculture, 6: 31-58,
DANIEL, E., J. TERLIZZI, and P.E. KARLANDER, (1980): Growth of coccoid nanoplankter (Eustigmatophyceae) from the Chesapeake Bay as influenced by light, temperature, salinity and nitrogen source in factorial combination. /. Phycol. 60; 364-368.
PRESSCOT, I. (1981): The Algae: A review (J.R. Stein, ed.) Cambridge University Press. 385 pp.
DUNCAN, D.B. (1955): Mulple range and multiple F tests. Biometrics 11: 1-42.
SPECTOROVA, L.V. O.I. GORONKOVA, L.P. NOSOVA and O.N. ALBITSKAYA (1981): High-density
RODHOUSE, P.G,, C. RODEN and M.E. SOMMERVILLE-JACKLIN (1983): Nutritional value of microalgal mass cultures to the oyster Ostrea edulis L. Aquaculture; 32: 11-18.
PERTANIKA VOL. 11 NO. 3, 1988
473
ABDULLAH ZAINI ALIAS
culture of marine microalgae-promising items for maricurture I. Mineral feedings regime and installations for culturing Dunaliella tertiolecta Butch. Aquaculture, 26: 289-302. TROTTA, P. (1981): A simple and inexpensive system for continous monoxenic mass culture of marine microalgaeM
474
WALNE, P.R. (1974): Culture of bivalve molluscas, 50 years experience at Conway. Fishing News (Books) Ltd. West Byfleet 177 pp. j H ( 1 9 7 4 ) : s t a t i s t i c a l Analysis. Prentice-Hall, Inc., Englewood Cliffs, New Jersey,
ZAR
PERTANIKA VOL. 11 NO. 3, 1988
(Terima 8 Ogos, 1987)
Pertanika 11(3), 475-481 (1988)
Pengubahsuaian Bentuk Selang Kecerunan Dalam Kaedah Newton bagi Suatu Kelas Fungsi Satu Pembolehubah The Improved Interval Forms of the Gradient in Newton's Method for a Class of Functions with One Variable ISMAIL BIN MOHD Jabatan Matematik, Fakulti Sains dan Pengajian Alam Sekitar, Universiti Pertanian Malaysia, 43400 Serdang, Selangor Darul Ehsan, Malaysia. Kata punca: Kaedah Newton; aritmetik selang; bentuk tersarang. ABSTRAK Bentuk terubahsuai kaedah Newton selang yang dihuraikan oleh Hansen (Hansen, 1978b) untuk mengira dan membatasi pensifar f:Rl ~+Rl adalah diberikan. Untuk meningkatkan sifat-sifat penumpuan kaedah-kaedah itu, bentuk tersarang digunakan untuk mengira dan membatasi terbitan ff;Rl ~->R};. ABSTRACT Improved forms of some interval Newton methods which are described by Hansen (Hansen, 1978b) for computing and bounding the zeros of f:Rl -• Rl are given. In order to improve the convergence properties of the methods, the nested form has been used for computing and bounding the derivative 1. PENGENALAN Sudah menjadi pengetahuan umum di kalangan cendikiawan matematik bahawa kaedah Newton dapat digunakan untuk mencari pensifar nyata persamaan f(x) = O (1.1) dengan diberinya fungsi f : D C R 1 -+R 1 . Moore (Moore, 1966) menggunakan aritmetik selang kepada kaedah Newton dalam menyelesaikan persamaan (1.1) dan penumpuannya telah dihuraikan oleh Nickel (Nickel, 1971). Hansen (Hansen, 1987b) menunjukkan bahawa dengan penggantian beberapa selang tertentu dalam kaedah Moore tadi akan meningkatkan kadar penumpuan kaedah Newton selang. Dalam makalah ini pengubahsuaian kaedah Hansen tadi akan dihuraikan dan dalam perbincangan ini kita hanya mempertimbangkan fungsifungsi polinomial sahaja. 2. TATATANDA Selang x = [ x p x s ] • ( x E R 1 l x j < x < x s ; I1] jadix= [x!,x s ] dengan masing-masing Xj dan xs disebut influmum dan supremum. Tandakan set
l^K ) - JX - |Xj , XgJ
| Xj A
Pemetaan-pemetaan lebar w( . ) : dan titik tengah m(. ) :.I(R ! ) "^R 1 masing-masingnya ditakrif oleh w(x) = x s - X j ,
(2.1)
m (x) = (x, + x s )/2
(2.2)
dan
Operasi aritmetik peraduaan + , - , . , / ditakrif kepada set I(R*) sebagai x*y={x*y I xEx,yEyj
(2.3)
(x, y E KR1)) dan *E{+. - , . , / ] kecuali x/y tak tertakrif jika 0 ^ . Penjelasan mengenai (2.3) diberikan dalam (Mohd, 1987b). Persilangan antara x dan y kepunyaan I(R*) ditakrif oleh
f 0 ((xs
atau
L[maks minim i x g , y s ] ] selainnya
(2.4)
ISMAIL BIN MOHD
Selang x E I (R 1 ) disebut meftyusut jika dan hanya x{ = x s , Jadi selang menyusut berbentuk [x,x] (Vx G R 1 ) . Jika OGc dan n integer genap positif maka
xn = [O.maksWJ1, x§ } ].
(2.5)
3. PERLUASAN SELANG Katakan diberi fungsi f: DCR1 ^ R 1 dan fQC.^D). Pemetaan f:I(D) -> I(R*) bagi f ditakrif oleh fix) = { f ( x ) | x e c ; x € I ( D ) } (3.1) disebut perluasan seseunit bagi f. Katakan f:I(D) -M(R ! ) perluasan selang bagi f:D -> R 1 , maka f dikatakan berekanada rangkuman jika dan hanya jika (xCyEI(D)) mengimplikasikan (f(x) Cf (y)). Jelas'f yang ditakrif oleh (3.1) satu perluasan selang berekanada rangkuman. Fungsi selang nisbah fA : I(D) -* I(R l ) yang diperoleh daripada ungkapan nyata nisbah fungsi f menerusi penggantian semua xQ*. dalam ungkapan f(x) dengan x E I (R) dan aritmetik nyata dalam f(x) dengan aritmetik selang disebut perluasan selang asli bagi f.
5. KAEDAH NEWTON SELANG
Katakan diberi f : DC R1 -•R 1 dengan : Jika x E I(D) maka (V x, yE x) wujud £ Ex sehingga f(y)=f(x)+f'(£)(y-x)
Misalkan f' : I(D) - ^ ( R 1 ) perluasan selang berekanada rangkuman bagi f* : D -* R 1 . Oleh kerana x padat jadi x cembung dan £ E x. Justeru itu f(y)Ef(x) + f ' ( x ) ( y - x ) (5.2) Jika yEx pensifar nyata bagi f dalam x maka f(y) = 0, jadi y memenuhi untuk £ E x. Jadi sebarang pensifar y E x terkandung dalam set D'= {y|f(x) + f ' ( f t ( y - x ) = O] (5.4) 1 Kita akan mencariyEI (R ) sehingga D' Cy. Jika kita takrifkan y EI (R l ) sebagai y = x - f(x)/f (x)
k)
Katakan fungsi f: D C R 1 -» R1 ditakrif oleh +
)
(4.1)
+ a j X + aQ
dan bentuk tersarang bagi f(x) seperti berikut
(5.5)
maka 0 (£ f' (x) mengimplikasikan D' C y seterusnya menerbitkan kaedah Newton selang yang terdiri daripada penjanaan jujukan (x' ^) daripada
4. BENTUK TERSARANG f(x) = a n x n + a ^ x * " 1
(5.1)
(5.6a) (5.6b)
biasanya sama dengan
dengan
dan
f(x) = (. . . (((anx + a n _j)x + a n _ 2 )x + a n _ a )x + . . . + a i ) x + a0-
(4.2)
Perluasan selang l s : I(D)->I(R 1 ) bagi f yang ditakrif oleh (4.2) diberi oleh f s (x) • ( ; . - ( ( (a n x + an_ i)x + a n _ 2 )x + ^_ 3 )x + . . . + aj)x + a0 ; (4.3) sedangkan perluasan selang JA : ^ D ) - * ! ^ 1 ) bagi f diperoleh menerusi (4.1), iaitu fA (x) = ^x 1 1 + a n . j x 1 1 " 1 + . . . +
aix
+ a0.
Jika 0 E f ' (x), D' masih boleh dibatasi seperti ditunjuk dalam (Mohd, 1987a). 6. PROSES PENGUBAHSUAIAN HANSEN
Katakan kuasa kepada pembolehubah x dalam ungkapan f(x) lebih besar dari satu. Gantikan satu x atau lebih dengan xx dan bakinya dengan x 2 . Jika ungkapan baru ditandai g(x x , x 2 ) maka untuk x = X! = x2 diperoleh x 2 ) = g(x,x) = f(x)
(4.4) Telah dibuktikan (Moore, 1966) bahawa
l s (x)) + I A (x)
(4.5)
w(f s (x))
Menurut kembangan Taylor (Vxj ,yx Ex dan ( V x 2 , y 2 E x 2 ) wujud ^ i ^ X ! dan sehingga
sehingga diperoleh
(6.1)
(4.6) PERTANIKA VOL. 11 NO. 3, 1988
PENGUBAHSUAIAN BENTUK SELANG KECERUNAN DALAM KAEDAH NEWTON
(6.2) dan x{ j = x untuk i, j yang berkaitan. Dengan Jika X! = x2 = x dan yt =
mengambil N(x) = £ dalam (5.5) diperoleh = y maka
N(x) = x-f(x)/f'(x)
(6.9)
h
(6.3)
Kaedah lelaran menggunakan (6.7) adalah P e r t i berikut: Mulai dengan x ( 0 ) = x janakan jujukan (x ( k ) ) dari se
sehingga menerusi (6.1) diperoleh (6.10) (6.4)
Jika y pensifar nyata bagi f maka f(y) = 0 dan menerusi (6.4) diperoleh
Nickel (Nickel, 1971) telah menunjukkan bahawa kaedah Newton selang satu matra yang diberi oleh (5.6) menumpu asalkan 0 G f (x) sedangkan untuk 0 Ef' (x) ditunjuk dalam (Mohd, 1987a) dan (Hansen, 1978a). Sekarang dari (6.8) diperoleh
f(x)
Jika x E x dan y G x maka £i, 7? Ex dan pensifar nyata y terkandung dalam selang f(x)
N(x) = x -
(6.6)
..,x)+ A ox2
t
dengan x n , X2i> 2^22
sama
+ ...+ A g(x,...,x) dengan x. Namun
begitu kita masih menggunakan tatatanda xx x,
a*m
x2 x, x 2 2 untuk menyatakan yang mereka merdeka
ax,
serta masing-masingnya membatasi | 1 ; yt dan TJ2.
- f'(x)
Sekarang jika kuasa bagi x dalam ungkapan f(x) lebih besar atau sama dengan m maka kita dapat menggantikan f(x) dengan g(xj, . . . , x m ).
Justeru itu lebar selang (6.10) lebih kecil dari (5.6).
Dengan cara yang sama seperti di atas diperoleh Nm(x) = x-f(x)/S'(x)
(6.7)
dengan g'W •
~
7. PROSES PENGUBAHSUAIAN TERSARANG Katakan fungsi f : D C R1 -* R1 berbentuk polinomial
g(xU)x,...,x) + (7.1) K=0
(6.8)
yang mempunyai pensifar nyata dalam x. Kita mempunyai dua kes untuk dibincangkan di sini, iaitu kes-kes 0 £x dan 0 E x .
PERTANIKA VOL. 11 NO. 3, 1988
477
ISMAIL BIN MOHD
Katakan 0 € x maka untuk m < n, g' (x) — dibina menerusi (6.8). Oleh kerana f berbentuk polinomial dan m >2 maka ~- g(Xjl >..., Xii» x , . . . , x) (7,2) 1
seperti yang dinyatakan dalam (6.8), berbentuk polinomial juga sehingga g' (x) dapat diubah menjadi bentuk tersarang g^(x) seperti diterangkan dalam Bahagian 4. Dengan demikian
k
a,
x
~ m + 1 Xo . . x
Oleh kerana OG x dan Xj j = x untuk i, j yang berkaitan maka seperti yang diterangkan dalam Bahagian 6 pilihlah x = 0 sebab x boleh dipilih sebarang sehingga diperoleh 9 ( 9x. - i l ' '"""' '
dan (7.4) n Jika 0 Ex tulislah polinomial (7.1) menjadi
/
a. \ k ~ m
+ l
x.. , x . . . x
(7.5) dan Terbitkan separa (7.5) sehingga diperoleh i
J-g(x1)...,xm) = a1+a2x2
a
x
k—m 1
d
-+am_1x2...xm_1
k=m
3 dx
lX3
+ ...+
2
Oleh sebab itu menerusi (6.8) diperoleh . ^' (x) = aj + a2 x + a3 x 2 + . . . + a. x 1 " l + . . . d
m-l^
k=m
Z
k-m+1X a x k l 3 • • -xm'
... X
(7.6)
•E
k=m
-, +
Jika %t • . . . « x _ * x dalam (7.5) maka diperoleh g(xi ,• - , x m ) = g(x sehingga k=m
k^x k - 1
dan k=l
478
PERTANIKA VOL. 11 NO. 3, 1988
x)=f(x)
PENGUBAHSUAIAN BENTUK SELANG KECERUNAN DALAM KAEDAH NEWTON
yang jelas memenuhi
Diketahui (Moore, 1966) bahawa
dan
Oleh sebab itu w(gf(x))
dengan g' (x) diberi oleh (7.6).
Jadi jika 0€x maka kita gunakan rumus
Sekarang daripada (7.6) kita bina bentuk tersarang berikut 2
4
n
2
gg(x) = a! + (a2 + a4 x + a6 x + . . . an x ~ )x + (a 3 + a 5 x 2 + a 7 x 4 + . . . + a
g' TB (x) = g' T (x) ^ g ' S A (x)
bagi menggantikan f' (x) dalam (5.5). Jelas bahawa sTB ^ ^ —£.T ^
x 11 " 4 ) x 2
n l
(7.7) untuk n genap dan g§ (x) = &i + (a 2 + a4 x 2 + a 6 x 4 +
(7.12)
8. CONTOH-CONTOH Kaedah-kaedah di atas diilustrasikan oleh contohcontoh berikut dalam satu lelaran sahaja. Contoh 8.1: Misalkan f : R1 -^R 1 ditakrifkan oleh f(x) = x 4 - 1
n
3
...+ anx ~ )x
2
untuk n ganjil yang kedua-duanya memenuhi (7.3) dan (7.4).
yang mempunyai pensifar 1 di dalam selang [0.5,3.5] danpilihlahx = 2. Menerusi (6.9) diperoleh N(x) = 2 - f(2)/4x3
Daripada (7.7) diperoleh g'T (*) = a! + x (a 2 + x 2 (a 4 + x 2 (a 6 + . . . +
yang memberikan N ([10.5, 3.5])= [-28, 1.913], Jika dibentuk g(x!,x2,x3,x4) =
2^2 (a 5 + x 2 (a 7
x2x3x4-
maka menerusi (6.7) diperoleh (7.9)
N
(v\
zz
9 _
X 3 +X 2 X 2 1 + XX
untuk n genap dan g'T (x) = ax + x (a 2 + x 2 (a 4 + x 2 (a 5
+2U 1X42X43
dengan g' (X) = X3 + X 2 X 2 , + XX3 ! X 3 2 + X4 12U2X43
dan setiap XJJ = [0.5,3.5] (1-2,3,4; j = l , 2 , 3)
x 2 (a 3 + x 2 (a 5 + x 2 (a 7 + . . . (7.10) untuk n ganjil.
sehingga memberikan N 4 ( [ 0 . 5 , 3 . 5 ) * [0.5882,1.8330] Dengan menggunakan (4.3) kita mengira
Persamaan (7.6) dapat ditulis dalam bentuk tersarang Alefeld dan Herzberger (Alefeld, Herzberger 1983) seperti berikut. g' SA (x) = 3LX + x (a 2 + x (a 3 + x(a 4 + . . . + x(a
- +a x ) . . . ) ) )
,
(7.11)
N (x) - 2 —
~s
"
f (2)
x 3 +x(x 2 +x(x + x
yang memberikan N s ([0.5, 3.51])= [0.5882, 1.8322] dengan gs(x) = x 3 + x(x 2 + x(x + x) ).
PERTANIKA VOL. 11 NO. 3, 1988
479
ISMAIL BIN MOHD
Jelas didapati bahawa
dan menerusi (7.9) (sebab n genap) diperoleh g^(x) - -60 + x(47 + x 2 (l) ) + x 2 (-12)
N s (x)CN 4 (x)CN(x) untuk x = [0.5,3.5].
sehingga memberikan
Contoh 8.2: Misalkan f : R1 ^ R 1 ditakrif oleh
g ^ h l , 21])= [-159,42] . Akan tetapi menerusi (7.11) diperoleh
f(x) = x 4 - 12x 3 +47x 2 - 6 0 x yang mempunyai pensifar 3 dalam selang [2.5, 3.7] danpilihlahx=3.1. Dengan menggunakan fA (x), £ (x) dan gg(x) yang masing-masing berbentuk f A GO • 4x3 - 36x2 + 94x - 60, I GO • (x3 - 12x2 + 47x - 60) +
gsAGO = -60 + x(47 + x (-12 + x) ). memberikan Oleh sebab itu dengan menggunakan (7.12) diperoleh lVBGO= [-120,42]. Jadi jelas bahawa
(xx2 - 12xx + 47x) + ( x 2 x - 12x2) + x 3 ,
dan
dan gs(x) = x 3 - 12x 2 +47x-60 + x(x 2 - 12x + 47 + x ( x - 12 + x)) memberikan _f;(x)= [-225.34, 265.412], jj'fe)= [-125.224, 134.72] dan
Oleh kerana f'A (x) dalam Contoh 8.2 dan f'AGO , g' (x) , g'T(x) , g' S A 00 » I T B ^ d a l a m contoh 8.3 menganduhgi 0 maka kaedah yang disarankan dalam (Mohd, 1987a) hendaklah digunakan untuk mengira N([-1,2]),N4([-1,2]),NT([-1,2]), NSA([-l,2])danNTB(hl,2]).
gg(x)= [-15.628,23.888] sehingga kita pasti memperoleh N s (x)CN 4 (x)CN(x) dengan masing-masing Ns(x) , N4(x) dan N(x) diperoleh dengan menggunakan g^(x), g'(x) dan f'A(x) dalam rumus Newton selang untuk selang x= [2.5, 3.7] Contoh 8.3; Misalkan f : R1 ->RJ ditakrif oleh f(x) = x4 - 12x3 + 47x2 - 6Gx + 24 yang mempunyai pensifar 1 dalam selang [-1, 2]. Jadi pilihlah x = 0 sebab 0€ [-1, 2]. Jelas didapati bahawa 4(hl,2])=[-314,160], menerusi (7.6) diperoleh £ ( [ - 1 , 2 ] ) = [-159,42] 480
untuk x= [-1,2] .
9. KESIMPULAN
Berdasarkan perbahasan dalam Bahagian 4 dan Bahagian 7, dan contoh-contoh 8.1, 8.2 dapatlah kita katakan bahawa kaedah pengubahsuaian tersarang lebih baik daripada kaedah pengubahsuaian Hansen dalam mengira dan membatasi pensifar syata fungsi polinomial (1.1). Begitu juga kaedah tersarang (7.12) lebih baik daripada kaedah tersarang (7.11). Walau bagaimanapun pengiraan g'Tfi (x) menerusi (7.12) memerlukan masa yang lebih kerana ia melibatkan pengiraan Dengan cara yang sama di atas kaedah yang disarankan di sini dapat dikembangkan untuk f : Rn -> Rn yang akan kita terangkan dalam makalah yang lain.
PERTANIKA VOL. II NO. 3, 1988
PENGUBAHSUAIAN BENTUK SELANG KECERUNAN DALAM KAEDAH NEWTON RUJUKAN
Bounding Real Zeros of a Class of Functions with one Variable, Pertanika, 10(1) : 89-95.
ALEFELD, G., and J. HERZBERGER. (1983): Introduction to Interval Computations, Academic Press, New York. HANSEN, E.R. (1987a) : A Globally Convergent Interval Method for Computing and Bounding Real Roots, BIT, 13: 4 1 5 4 2 4 . HANSEN, E.R. (1978b) : Interval Forms of Newtons Method, Computing. 20: 153-163.
MOHD, I.B. (1987b) : Aritmetik Selang, Serita Matematik, 32 : 15-22. MOORE, R.E. (1966) : Interval Analysis, Prentice-Hall. NICKEL, K. (1971) : On the Newton Method in Interval Analysis, Mathematics Research Center Report 1136, University of Wisconsin.
MOHD, I.B. (1987a) : The Comparison Between Hansen's Method and AlefekTs Method for Computing and
PERTANIKA VOL. 11 NO. 3, 1988
(Terima 13 November, 1987)
481
Pertanika 11(3), 483-485(1988)
COMMUNICATION I Survival and Isolation of Avian Mycoplasmas from Drinking Water of Infected Chickens. ABSTRAK
Mikoplasma telah disuntik ke dalam air minum ayam dan diambil contoh pada waktu yang berlainan, Pada kepekatan yang tinggi, organisma-organisma boleh didapati kembali daripada air 24 jam pos inokulasi. Mikoplasma tidak boleh didapati kembali daripada air minum ayam yang berinfekst
ABSTRACT
Mycoplasma was inoculated into the drinking water of chicken and sampled at different time intervals. At a higher concentration, the organisms could be recovered from the water 24 hours post inoculation. Mycoplasma could not be recovered from the drinking water of infected chickens. INTRODUCTION
The survivial of avian mycoplasmas within and outside the body of the host is important in epidemiology (Jordan, 1985). In the study of the gpidemiology of avian mycoplasmas, little has been reported on the survival of these organisms in the environment. Perlstein (1969) cited by Polak-Vogelzang (1977) reported that Mycoplasma gallisepticum could be recovered from drinking water of infected chickens and several strains of M. gallisepticum remained viable for at least 48 hours after inoculating broth cultures into water. Polak-Vogelzang (1977) stated that survival of M. gallisepticum in mains water at room temperature depended on the initial number of organisms and the incorporation of mycoplasma broth in the suspending medium. In a survey on the prevalence of mycoplasmas in the indigenous fowls, Shah-Majid and Nihayah (1987) isolated mycoplasmas from 64 out of 205 fowls sampled. The mycoplasmas were isolated from the choanal cleft of these fowls. Perhaps, at the time of drinking, the mycoplasmas that are present in the choanal cleft may be introduced into the water and other fowls that share the same drinker may be infected. This study was conducted to determine if mycoplasma could survive in the drinking water of chicken and to isolate mycoplasmas from the drinking
water consumed by infected chickens. MATERIALS AND METHODS
Mycoplasma spp. isolated from the choanal cleft of chicken during the preliminary survey was cultured in mycoplasma broth (Mycoplasma Supplement-G, Oxoid Limited, England). After 2 - 3 days of incubation, the viable count was determined using the plate count method and was diluted to contain approximately 102 and 10 8 colony-forming units (CFU) per ml. The mycoplasma was dispensed in 1 ml aliquot, frozen at - 2 0 C. On the day of inoculation, the vial was thawed before use. After inoculation, the exact numbers of mycoplasmas used was determined by the plate count method. In the first study, a rectangular and round drinkers were filled with tap water (ph 7.6) and kept at room temperature. The rectangular (62 x 5 x 7.5 cm 3 ) drinker has a capacity of 2,16 litres of water when filled up to the brim while the round drinker has a capacity of 3.28 litres. One millilitre of 4x 102 cfu/ml of mycoplasmas were inoculated into the rectangular drinker and another 1 ml of the same concentration was inoculated into the round drinker. The mycoplasmas were inoculated into the drinking water with a 1 ml pipette and attempt were made to cover the entire length of the water surface with mycoplasma. A sterile dry
MOHD SHAH-MAJID
cotton swab was used to sample the water surface for mycoplasmas. At the time of sampling, about 1 inch of the cotton swab was dipped into the water and was dragged slowly to the other end and back to its starting point. A different swab was used for different drinkers and for sampling at different time intervals. The swabs were then inoculated onto Mycoplasma Supplement -G agar medium. The water was sampled after 10, 20, 30 mins and 1, 4, 8, 12 and 48 hours after initial inoculation. The same procedure was repeated with 5.5 x 108 cfu/ml inoculum. All plates were incubated at 37 C in the presence of 5 per cent carbon dioxide and were examined for growth at days 2, 4, 7, 10 and 14 of incubation. In the second study, two-month-old indigenous fowls belonging to the university farm were initially swabbed for the presence of mycoplasmas
in the choanal cleft region. Ten mycoplasma positive birds were then selected and kept in individual cages. These birds were allowed to drink and the remaining water was immediately sampled for mycoplasma using a dry cotton swab. Samples were then inoculated onto Mycoplasma Supplement—G agar medium. Five water samples (after drinking) were taken from each bird. RESULTS AND DISCUSSION Table 1 shows the recovery of mycoplasmas from the drinking water at different time intervals following inoculation. Mycoplasmas remain viable in the drinking water when the inoculum used is high. Mycoplasmas could be recovered from the tap water 48 hours post inoculation and this is similar to the observation made by Perlstein (1969) and Polak-Vogelzang (1977). Polak-
TABLE 1 Recovery of mycoplasmas from the drinking water at different time interval following inoculation
Inoculum
Time interval after initial inoculation
4x10' organisms/ml
10 mins 20 mins 30 mins lhr 4hrs 8hrs 12hrs 24 his
5.5 xlO 8 organisms/ml
10 mins 20 mins 30 mins lhr 4hrs 8hrs 12hrs 24hrs 48hrs
+-H-
484
Oto 10 colonies 10 to 100 colonies > 100 colonies No Isolation PERTANIKA VOL. 11 NO. 3, 1988
Isolation of mycoplasmas from: rectangular water drinker
round water drinker
SURVIVAL AND ISOLATION OF AVIAN MYCOPLASMAS FROM DRINKING WATER OF INFECTED CHICKENS
Vogelzang (1977) stated that the survival of mycoplasma strains in mains water at room temperature depend on the number of initial organisms used and the suspending medium of the mycoplasma broth. Polak-Vogelzang (1977) was able to isolate mycoplasmas at 6 to 8 log 10 cfu/ml for 4 to 5 days when suspended in 1% broth in sterile water at ph 7.7. However, at concentrations less than 6 log 10 cfu/ml, the mycoplasmas survive for less than one day. In the present study, mycoplasma could not be reisolated from both drinkers as early as 10 mins post inoculation when the inoculum used was 102 cfu/ml. Mycoplasmas could not be recovered from all the water samples taken from all the infected birds. The infected birds used in this stifdy were clinically normal with no nasal discharges, sneezing or coughing but only the presence of mycoplasmas in the choanal cleft region. Polak—Vogelzang (1977) suggests that under certain environmental conditions or when well protected by exudates or tissue debris, the survival of mycoplasma in the environment may be longer. However, this study shows that mycoplasma that were present in the choanal cleft region cannot be isolated from the water sample at the time of drinking using the swabbing technique. A small number of mycoplasmas could have been transmitted into the water but the technique of swabbing the water surface may not be efficient in detecting small numbers of mycoplasmas as shown in the drinker studies (Table 1). Perlstein (1969) cited by PolakVogelzang (1977) observed than when infected nasal discharges were placed in water, the Myco-
plasma gallisepticum could be recovered from the water for two days and drinking water of infected chicks yielded Mycoplasma gallisepticum. It is probable that the transmission of Mycoplasma gallisepticum into the water at the time of drinking is from the nasal discharges. In the present study the infected birds were clinically normal with no nasal discharges. Under farm conditions and at room temperature, it is probable that the important factors which will influence the survival of mycoplasma in drinking water are the presence of agents which would protect against osmotic lysis (Polak-Vogelzang, 1977) and high concentration of mycoplasmas as shown in this study. REFERENCES JORDAN, F.T.W. (1985): Gordon Memorial Lecture: People, poultry and pathogenic mycoplasmas. Br. Poult. Set 26; 1-15. POLAK-VOGELZANG, A.A. (1977): Survival of Mycoplasma gallisepticum in mains water. Avian Pathol. 6: 93-95. SHAH-MAJID. M and M. NIHAYAH. (1987): Choanal cleft as an alternative site for the isolation of avian mycoplasmas. Vet. Rec. 121: 153. (Recieved 14, May, 1988)
MOHD SHAH-MAJID Dept. of Veterinary Microbiology and Pathology Faculty of Veterinary Medicine and Animal Sciences, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia.
PERTANIKA VOL. 11 NO. 3, 1988
485
Perlanika 11(3), 487-490 (1988)
COMMUNICATION II Preliminary Estimates of Growth and Mortality in Nemipterus bathybus Snyder (Pisces: Nemipteridae) from the Coast off Sarawak, South China Sea ABSTRAK Sampel kekerapan panjang Nemipterus bathybus Snyder (Pisces: Nemipteridae) yang diperolehi dari kajiselidik di luar pantai Sarawak, Malaysia Timur (Laut China Selatan) telah diasingkan ke dalam komponen dengan kaedah Bhattacharya. Panjang min komponen berdekatan yang dianggap mewakili kelas tahun telah digunakan untuk menganggarkan parameter L^ dan K dalam persamaan Bertalanffy, manakala bilangan ikan dalam komponen ini digunakan untuk menganggarkan jumlah kematian. Ke~ putusan adalah: TL^ = 312, K • ft36 -7 tahun dan Z = L4 tahun ~J. Keputusan telah dibincang berdasarkan anggaran lain bagi N. bathybus dan spesisikan kerisilain.
ABSTRACT Length-frequency samples of Nemipterus bathybus Snyder (Pisces: Nemipteridae) obtained in a trawl survey off the coast of Sarawak, East Malaysia (South China Sea) were separated into components using the Bhattacharya method. The mean length of adjacent components, assumed to represent year classes, were used to estimate the parameters Loo and K of the von Bertalanffy equation, while the numbers of fisah in these components were used to estimate total mortality. Results were: TLOO =31.2, K = 0.36 year~ and Z = IA year . The results are discussed in relation to other estimates o/N. bathybus and other species of threadfin breams. INTRODUCTION A brief survey was conducted from 17 to 21 June 1987 on board R/V Kagoshima Maru in the waters off Sarawak, South China Sea. A total of 12 trawl hauls were made out of which 7 were successful and positive for Nemipterus bathybus Snyder visees: Nemipteridae). Length measurements were taken for the purpose of growth and mortality estimation, as needed for future assessment of the stock. MATERIAL AND METHODS The length-frequency data used here are summarized in Table 1. They were used in their original form (sample i (a) ) and after smoothing over 3 classes as recommended in Pauly and Morgan (1987) for irregular data (samples i (b) ). The method of Bhattacharya (1967) as implemented in the Compleat ELEFAN software package (Pauly and Morgan 1987) was used to
separate samples i (a) and i (b) into their normally distributed component distributions (Fig, 1). Assuming, along with Weber and Jothy (1977) and Pauly and Martosubroto (1980) that the major components identified represent year classes (i.e. At = 1, the mean length between adjacent groups (AL) was computed, and related to the mean length of all components involved in the computation of A L(L). A rough estimate of asymptotic length (Loo) was obtained from Lmax/0.95 = L^ where Lmax is the maximum size in Table 1 (30.5 cm, TL). A preliminary estimate of the parameter K of the von Bertalanffy equation was then obtained using a "forced Gulland and Holt Plot" (Pauly 1984, see also Gulland and Holt 1959) i.e. from K = (AL/A t) ( L ^ - L) . . . .1) while a preliminary estimate of tQ was obtained from the empirical equation 10g 10 (-t Q ) - -0.3922-0.2752 x log 10
MOHD. AZMI AMBAK, ABU KHAIR MOHAMMAD MOHSIN AND MOHD. ZAKI MOHD. SAID
where Nt, Nt + 1 represent the sum of number of fish in adjacent groups (Table 1).
Sample 3 a
A
RESULTS AND DISCUSSION
t
Table 2 summarizes the results of the Bhattacharya analyses and identifies the components used to estimate K and Z. With Loo estimated as 32.2 cm, K is computed, following (1), as K = 5.1/(32.2-17.9) = 0.36 . . . .4) while t = -0.45 based on equation (2). Thus the von Bertalanffy equation, based on our data for the stock of Sarawak is:
Sample 3 b
L t = 32.2[l-e- 6 - 36 ( t+0 - 45 )]
Sample 5 a
1 Fig. 1:
i
Sample 5 b
ACKNOWLEDGEMENTS Length cm
Sample outputs of Bhat tar chary a's method for separation of composite length-frequency samples into their component distributions, as inplemented on the Compleat ELEFAN program (a; data of Tables l;b: smoothed data; see Table 2).
1 ^ - 1 . 0 0 3 xlogi 0 K ....2) (from Pauly and Martosubroto 1980, with a correction for the sign of the intercept from Pauly, pers. comm.). Total mortality (Z) was estimated, assuming representativeness of the samples in Table 1, and constancy of recruitment from Z--log, 488
....5)
Our estimate of total mortality, based on the N values in Table 2, is Z = 1.37 year" 1 , corresponding to a survival rate of 2.5% year . This latter result is compatible to those of Weber and Jothy (1977) who estimated values of Z = M ranging from 0.4 to 2.2 off Sarawak and Sabah. Similarly, the (mean = 1.31) for nemipterids growth parameter estimates are comparable to these reported for N. bathybus by Pauly (1978), based on Eggleston 1972 i.e. L^ = 30.2 cm, K = 0.324 and to • -0.80. This correspondence is encouraging, although due to the nature of our data, our results must be considered as preliminary.
•3)
We would like to thank JICA for enabling us to spend, in April 1988 a study period at ICLARM, and Dr. Pauly, for his assistance with the completion of this paper.
MOHD. AZMI AMBAK ABU KHAIR MOHAMMAD MOHSIN MOHD. ZAKI MOHD. SAID
Department of Fisheries Biology Faculty of Fisheries and Marine Science Universiti Pertanian Malaysia 43400 UPM, Serdang, Selangor, Malaysia.
PERTANIKA VOL. It NO. 3, 1988
PRELIMINARY ESTIMATES OF GROWTH AND MORTALITY IN NEMIPTERUS BATHYBUS
£ 3
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PERTANIKA VOL. 11 NO. 3, 1988
489
MOHD. AZMI AMBAK, ABU KHAIR MOHAMMAD MOHSIN AND MOHD. ZAKI MOHD. SAID TABLE 2 Summary of results of Bhattacharva analysis of length-frequency data of Nimepterus bathybus. Mean length (cm) Sample*^ la lb 2a 2b 3a 3b 4a 4b 5a 5b 6a 6b 7a 7b
T
t
t
17.3 17.5 17.3 16.9 16.3 16.6 17.8 18.1 15.8 15.8 14.5 14.3 11.7 11.4
S.D. of Groups
_ 21.9 21.7 21/9 22.2 20.5 21.0 17.7 17.3
1.61 1.85 10.7
t+1
N in Group Nt
1.39 1.14 1.39 1.74 2.12
0.91 0.91 0.70 (0.70) 0.99 0.99 -
1.17 1.35
0.82 0.82
105 100 31 31 189 214 208 236 258 265 81 83 52 50
Sumb>
1472
1.41 0.95 1.36 1.04
-
a)
a refers to estimates based in the original length frequency data, b to data smooted over 3 length classes.
b)
refers to N-values for which both N. of Nt + ^ w-ere available, as used in equation (3).
REFERENCES BHATTACHARYA, C.G. (1967): A simple method of resolution of a distribution into Gaussion components. Biometrics 23: 115-135. EGGLESTON, D. (1972): Pattern of biology in the Nemipteridae. /. Mar. Biol, Assoc. India, 14(1): 357-364. GULLAND, J.A. and S.J. HOLT. (1959): Estimation of growth parameters for data at unequal time intervals. /. Cons. Int. Explor. Mer 25: 47-49. PAULY, D, (1978): A preliminary complilation of fish length growth parameters. Ber. Inst. Meeresk. Univ. Kiel No. 55, 200 p.
_ 67 43 61 33 74 77 9 11 375
culators. ICLARM studies & Reviews 8, 325 p. ICLARM, Manila, Philippines, PAULY, D and P. MARTOSUBROTO. (1980): The population dynamics of Nemipterus marginatus (Cuvier & Val.) off Western Kalimantan, South China Sea, J. Fish. Biol. 17: 163-273. PAULY, D. and G,R. MORGAN. (1987): (eds.). LengthBased Methods in Research. ICLARM Conference Proceedings 13. Manila, 468. p. WEBER, W. and A.A. JOTHY (1977): Observations on the fish Nemipterus spp. in the coastal waters of East Malaysia. Arch Fisch wiss., 28(182): 109-122.
PAULY, D. (1984): Fish population dynamics in tropical waters: A manual for use with programmable cul-
490
Nt + 1
PERTANIKA VOL. 11 NO. 3, 1988
(Received 17 May,1988)
Subject Index for Volume II Achras sapota harvester 161-164 Alluvial soils acidity 15-23 aluminium 15—23 charge characteristics 15-23 Ammonia volatilization, effect or air-flow rates 323-325 Arsenobetaine hydrobromide preparation 437—439 Ascorbic Acid see Vitamin C
Embryo culture media formulation 357-363 recalcitrant species 357-363 Floating refuse Gombak River, Kuala Lumpur 151-155 Fluorescent esterase 7-13 Forest, hill soil compaction 31-37 Fusarium oil palms 317-321
Automorphisms braid groups 115-123 Fuchsian groups 115-123 mapping class groups 115-123, 299-303 Seifert fibre groups 115-123, 299-303 surface groups 299-303 Biological activity see medicinal properties Buffaloes disease conditions 51-55 Serdang, Selangor 51-55 Capsicum annuum Colletotrichwn capsici, infection 341-344 Chlorella virginica Light intensity 469-474 salinity 469-474
Gallus gallus lung, qualitative studies 299-238 Glomus fasciculatum mass production 143-145 Glycine max breeding nurseries 33-340 response to row spacing 333-340 Hexokinase 7-13 Hevea latex drying, microwave 289-297 total solid content 289-297 Howea forsteriana dormancy, breaking 137—141 gibberellic acid 137-141 Hydrilla verticillata herbicide Aquathol K, effect of 191-124 Hyptis suaveolens essential oils, composition 239-242 Imperata cylindrica allelochemicals 175-189 Ion-exchange resin poly (hydroxamic acid), preparation, properties 255-259 Ionization coefficient, secondary excited level, 3 P 0 and 3 P 2 311-315
Conopomorpha cramerella polymorphisms 7—13 Corcyra cephalonica insecticides, effect of 195-201 Corn oil frying 393-398 Cucumis melo storage, physico-chemical characteristics 203-209
Lead
Dermochelys coriacea hatchings, feeding 147-149 Dehydroascorbic acid see Vitamin C Domestic fowl see Gallus gallus
Malate dehydrogenase 7-13 Malic enzyme 7-13 Mangrove Rhizophora species, stocking density 365-374 soils, chemical characteristics 385-391
Kuala Lumpur 157-160 urban dust 157-160
PERTANIKA VOL. 11 NO. 3, 1988
491
Mentha arvensis medicinal properties 249-253 Metaphycus helvolus host-searching 1—5 Muskmelon see Cucumis melo Mycoplasmas, avian drinking water, chickens 483-485 N-methoxy-N-methylaminocarbonyl-1, 4-benzoquinone synthesis 87-90 Nemipterus bathybus growth and mortality 487-490 Nephelium lappaceum chemical dipping, shelflife 407-417 Nephelium lappaceum seeds, proximate analysis of 211-215 shelflife, packaging, modified atmosphere 217-228 Newton methods interval analysis 475-481 nested form 475-481 Newton polyhedral method cardinality 125-131 congruence equation 125-131 Normal populations, heteroscedastic chi-square test 133-135 coefficient of variation 133-135 maximum likelihood 133-135 Ocimum sanctum medicinal properties 249-253 Oil palm wastes furfural production 243-248 Organic chemistry problem solving strategies 261-271 Ortoshipon staminea medical properties 249-253 Oryza sativa ammonium nitrogen 25-30 fertilizer nitrogen 25-30 nitrogen efficiency 25-30 nitrogen balance 25-30 Oryza sativa submergence, chemical characteristics 385- -391 Padina alginates 79-85 Palm Oil Nutritional evaluation 399-406
492
Palm olein frying 393-398 nutritional evaluation 399-406 Palm stearin nutritional evaluation 399—406 Pisum sativum post-harvest storage 45—50 root, genetic control 165-173 sugar content 45-50 titratable acidity 45-50 total soluble solids 45-50 Potato crisps frying, shelf-life 393-398 Psidium guajava Ascomycetes 349-355 Deuteromycetes 349-355 Protein film ckickpeadhal 327-331 Rainfall data analysis 101-106 Rambutan see Nephelium lappaceum Real money balances Malaysian Agricultural Sector 451-460 Rice drying, milling cost functions 283-288 malathion residues 13—1% Rubber, natural market model 441—449 Saissetia coffeae natural enemies 1-5 Sargassum alginates 79-85 Scutellista cyanea host-searching 1-5 Sheep Brachiaria decumbens 57-61 hepatotoxicity 67-61 stall-fed 57-61 Soil properties metamorphic rocks, parent materials 375-384 Soil series Carey Island 15-23 Soya Oil frying 393-398 Stock market returns market behaviour 107-114 weekend effect 107-114
PERTANIKA VOL. 11 NO. 3, 1988
Transportation, off-road soil compaction 31-37 Tragulus javanicus
anatomy, abdomen 419-426 radiographic anatomy 427-435 Trawling drag force 91-100 ground drag 91-100 ground friction 91-100 ground spread 91-100 spread force 91 — 100 Van der Pol equation small parameters 305-309
Vesicular-arbuscular mycorrhizal see Glomus fasciculatum Vitamin C dye-titration 39-44 micro-fluorometry 39-44
Waterquality Selangor, Sg. Langat 273-281 Waterquality Watersheds, Selangor 461-468 Wood moulding marketing mix 63-72
PERTANIKA VOL. 11 NO. 3, 1988
493
Author Index for Vol. 11 1988 AbasMazni Othman 57 Abdul Ghani Ibrahim 1 Abdul Jalil Hassan 243 Abd. Rahman Kasmani see Kasmani, A.R. Abdulah Zaini Alias 469 Abu Khair Mohammad Mohsin see Mohsin, AK.M. Abu Osman M.T. see Abu Osman bin Md. Tap Abu Osman bin Md. Tap 115, 299 Ahmad, F.B.H. 87 Aini Mohd Salehuddin 73 Alang,Z.C. 141,357 Andrews, A.C. 175 Anhar Suki see Suki, A. Annuar bin Md. Nasir 107 Ashari Muktar 365 Asman Hassan 283 Atan M.Sharif 243 Augustin, M.A. 45, 203, 211, 393 Au,WaiFong 195 Awan, M.A. 345 Aziz Saharee, A. 51 Azmi, T.L 419,427 Azizah Osman 45, 203
Ismail bin Mohd
Bagyaraj,D.J. 239 Brophy, Joseph 239
Mahmood bin Mat Yunus, W. 311 Marcus Jopony, 385 Mashhor Mansor 191 Mat Rofa bin Ismail 305 Mohammad bin Yusof 441 Mohamad Ali Abdul Hamid 107 Mohamed Ali Abudl Hamid 107 Mohamad Nooh Hassan see Hassan, M.N. Mohamad Nordin bin Abdul Karim 163 Mohd. Aspollah H. Sukari 243, 249 Mohamad Azmi Ambak 488 Mohd Kamil Yusof see Kamil, M. Mohd. Nasir Azudin 203 Mohd Nawi Abd. Rahman 133 Mohd. Zaki Mohd Said 488 Mohsin, A.K.M. 488 Muzafar Shah Habibullah 451 Mohd Shah Majid 485 Mohd Wahid Samsudin 239 Md. Zin bin Wan Yunus, Wan 255 Mohd Zin Yusop 63 Mok, T.P. 273 Mok, Thean Poe see Mok, T.P.
Chan,E.H. Chew, T.A. Chin,H.F. Chong, Y.H. Chua, B.C.
148 283 141,357 399 211
Eshah Othman
217,407
Faridawati, M. 385 Fashihuddin Ahmad 79 Faujan bin H. Ahmad see Ahmad, F.B.H. Fauziah Abdullah, 407 Fuzina,N.H. 419,427 Gan, Y.Y. 7 Ghizan bin Saleh 165,333 Gritton.E.T. 165,333 Hassan, M.N 101 Hazar Ismail 437 Heng, L.K. 393 Ho, S.K. 39 Ho,Y.W. 320 494
475
Joyce, Joseph 191 Juzu Hayati Arshad 73 Kaida bin Khalid see Khalid, K.B. Kamaruzaman Jusoff 31 Kamel Ariffin Mohd Attan 125 Kamil, M. 273 Kasmani, A.R. 289 Kassim, H. 229 Khalid, K.B. 289 Khanif, Y.M. 25,325 King, A.S. 299 * Khor, H.T. 399 Krishnapillay, B. 141,257 Lai, F.S. 461 Laily bin Din 239 Lee,CK. 160 Lim, C.G. 160 Lim, T.K. 349 Lovett,J.V. 175 Low,K.S. 160
Ng. T.K.W. 399 Nor Aini Idris 393 Noorma Osman 195 PERTANIKA VOL. 11 NO. 3, 1988
Norajiki AJ. 461 Norasmah Ahmad 79 Nordin Hj. Lajis 243 Norlia Abdullah 331 Pancras, H. 325 Paramananthan, S.
15
Rajan, A. 175 RehanHj.MohdNor 261 Rita Muhamad 7 Rita, S. 7 Richardson, K.C. 419,427 Roslan A. Ghaffar 283 Sajjad, M. Siddique 345 Sakri bin Ibrahim 91-100 Sariah Meon 341 Saw, Leng Guan 365 Shamsher Mohamad 107 Shamsuddin, J, 15 Sharif Haron 57 Shukri Mohamed 63 Siraj Omar 79 Siti Maizura, S. 39 Sreenivasa,M.N. 145 Srivastava, P.B.L. 365
Stern, R.D. Suhaila Mohamed Suki, A. 273
203, 217, 331, 407
Takahashi Shozo 249 TanKaKheng 155 Tan, S.G. 7 Tee, E.S. 39 Thyer, Dan 191 Toia,T.F. 239,437 Varghese, G. 320 Vidyadaran, M.K. 229, 419, 427 Wahab, Z.A. 289 W. Zainun A. Nik see Zainun W. Nik, W. YongEuFan 155 Young, S.I. 39 YusufHadi 63 Zaharuddin bin Ahmad 255 Zaidan B.A. Wahab see Wahab, Z.A. Zainun W. Nik, W. 341 Zalma,M.L. 385 Zamri Saad,M. 51 Zauyah, S. 375 Zuriati Zakaria 239
PERTANIKA VOL. 11 NO. 3, 1988
495
Acknowledgement The Editorial Board acknowledges the help of the following reviewers in the preparation of Volume 11. Dr. Abdul Aziz Ariffin Mr. Abd. RahimHj.Nik Dr. Abdullah Hassan Dr. Abu Bakar Husin Dr. Abu Bakar Jaffar Assoc. Prof. (Dr.) Abu Osman Mr. Ahmad Ismail Prof. (Dr.) Ahmad Mahdzan Ayob Dr. Alias Hussin Mr. Ambrin Buang Dr. Ann Anton Dr. E.C. Appleby Dr. Ashfaque H. Khan Prof. Augustine S.H. Ong Dr. Baki bin Bakar Prof. (Dr.) J.M.Bruce Prof. K.A. Buckle Prof. (Dr.) J.R. Cannon Dr. H.T. Chan Mr. Charanpal Singh Dr. T.A. Chew Dr. Y. Chin Wee Dr. T.H. Chua Dr. Ir. Fr. De Connick Dr. E. Craswell Prof. R. Edwards Prof.(Dr.)I.R.P. Fillery Dr. L. Fredericks Dr. F.E. Gardiol Dr. Habibah Suleiman Dr. Hafiz Abdul Samad Prof. (Dr.)A,H.Haines Prof. (Dr.)J.G.Harvery Dr. Y.W. Ho Dr. Y.L. Hor Prof. G.M, Hughes Dr. Jinap Selamat Prof. (Dr.) A.H.Johnstone Dr. R.A. Jones Prof. (Dr.) P.G.Joseph Dr.J.C.Katyal Assoc. Prof. G. Kawamura Assoc. Prof. Khalip Abdul Raffar Dr. K.C. Khoo Dr. G.S. Khush Prof. D.T.Kowalski Assoc.Prof. (Dr.) A.J. Kuthubutheen
Assoc. Prof. (Dr) K.S. Low Prof. J.H. Loxton Prof.(Dr.)J.C.Madgwick Dr. Mansor Monsi Dr. Mat Isa Awang Dr. W.A.Mattews Prof. (Dr.) B.L. McNeal Dr. Mohd Fadzil Yahya Dr. Mohd. Hilmi Abdullah Assoc. Prof. (Dr.) Captain Mohd. Ibrahim bin Mohd Assoc. Prof. (Dr.) Mohd Khalid Md. Nor Dr. Mohd Kidin Shahran Prof. (Dr.) Mohd Nawi Abd. Rahman Assoc. Prof. (Dr.) Mohammad Noor bin Hj. Salleh Assoc. Prof. (Dr.) Mohd. Rashidi Razali Assoc. Prof. (Dr.) Mohamed Suleiman Dr. Mokhtar Abdullah Dr. Mokhtaruddin Abd. Manan Assoc. Prof. (Dr.) Mokhtar Bidin Dr. S.K. Neoh Dr. Nik Fuad Kamil Prof.(Dr.)T.N.Nisra Prof. Nordin Hassan Dr. Nordin Mohd Som Mr. K.S. Ong Assoc. Prof. (Dr.) S. Paramananthan Prof. (Dr.) Paula Michalik Dr. Rasedee Abdullah Dr. Razali Abdul Kadir Mrs. Rita Muhammad Prof. G. Rosenberger Dr. Roslan A, Ghaffar Dr. Rosli bin Mohamad Dr. Salleh Kadzimin Prof. (Dr.) Sham Sani Assoc. Prof. (Dr.) Shamsuddin Jusop Prof. Sanit Aksornkoae Prof. Emeritus Saul bin Saila Prof. (Dr.) Shigeo Hayashe Dr. M. Siddique Sajjad Mr.K.T.Siow Dr. Soleha Ishak Prof. (Dr.) E.L. Springer Prof. (Dr.) S.S. Stuchly Dr. Sulaiman Abd. Ghani Dr. Surjit Singh
PERTANIKA VOL. 11 NO. 3, 1988
Mr. P.F. Lam Prof. Max R. Langham Dr. B. Larsen Dr. B.S. Lee Ms. S.S. Lee Prof. (Dr.) Lee Sing Dr. S.C. Lim Dr. T.K. Lim Dr. W.H. Lim Dr. W.H. Loke Assoc.Prof.(Dr.)C.K.Wan Dr. Wan Muhamad Wan Muda Prof. R.B.H. Wills Dr. P.C.Wong Prof. R.S.Bob Wyburn Prof. R.S. Yanagawa Dr. Yahya bin Mat
Prof. T.C. Yap Dr. C.B. Yee Assoc. Prof. (Dr.) Yusof Hussein Assoc. Prof, (Dr.) Zaharah Abdul Rahman Dr. Zainol Eusof Prof. Zakri A. Hamid Assoc. Prof. (Dr.) Mohd. Zohadie Bardaie Prof. (Dr.) D.M.Sylvia Mr. E.S. Tee Prof. W.Taylor Dr. Y.T. Teng Prof. B.C. Tan Dr. B X Tan Prof. (Dr.) G. Varghese Prof. (Dr.) F. Vernon Dr. M.K. Vidyadaran Dr. S.S. Virmani
PERTANIKA VOL. 11 NO. 3, 1988
497
Agronomy Journal The American Society of Agronomy's bimonthly journal is highly respected for its technical reports on all aspects of plant and soil science. Original research findings in crop science; soil science; biometry; turf grasses; agroclimatology and agronomic modeling; crop, soil, pasture, and range management; and crop, forage, and pasture production and utilization are presented. Letters, invited reviews and notes on apparatus, experimental techniques, and observations/preliminary results and computer programs are also presented. $65.00 per year or $13.00 per issue in the U.S. $70.00 per year or $14.00 per issue elsewhere. Back volumes are available at $78.00 per volume ($13.00 per issue) in the U.S. Add $5.00 more per volume of $1.00 per issue elsewhere
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Footnotes - Footnotes are numbered consesecutively and typed at the bottom of the page below a horizontal rule. Key to author's name - Where an author's name does not conform with the Western practice of a surname appearing after the first and second names, the author should provide a key to his (her) name to facilitate the compilation of subsequent indices and bibliographies. Illustrations (a) All figures should be numbered consecutively from 1 onwards, in arabic numerals, and each figure should be referred to in the text. The title of the paper and the figure number should be written on the back of the sheet in pencil or (preferably) on a gummed label. Figures should be about twice the size of the printed reproduction. (b) Photographs: All photographs (glossy black-and-white prints) should be supplied with appropriate scales. The scales should be drawn on a separate sheet (not on the photograph). (c) Lettering and numbering on the face of the figure itself should be kept to a minimum. Unit of Measure - Metric units must be used for all measurements. Reprints - Authors receive 20 free copies of their articles. Manual - A manual, "Notes on the Preparation of Papers", is available on request from the Chief Editor.
