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UNIVERSITI PUTRA MALAYSIA COMPARATIVE STUDIES ON DIGESTIVE EFFICIENCY AND UREA KINETIC BETWEEN GOAT AND SHEEP DARLIS FPV 1995 3

COMPARATIVE STU DIES ON DIGESTIVE EFF IC IENCY AND UREA KINETIC BETWEEN GOAT AND SHEEP DARLI S MASTER OF SCIENCE UNIVERSITI PERTANIAN MALAYSIA 1995

COMPARATIVE STU DIES ON DIGESTIVE EFF ICIENCY AND U REA KINETIC BETWEEN GOAT AND SHEEP By DA RLIS Thesis Submitted in Fulfillment of the Requirements for the Degree of Master of Science in the Faculty of Veterinary Medicine and Animal Science Universiti Pertanian Malaysia September, 1995

ACKNOWLEDGEMENTS The author wishes to express his sincere appreciation and gratitude to Assoc. Prof. Dr. Norhani Abdullah, chairman of the supervisory committee for her invaluable guidance and untiring assistance throughout the period of his studies. Sincere thanks are also due to Prof. Dr. Ho Yin Wan and Assoc. Prof. Dr. Ridzwan Abdul Halim for serving as members of the supervisory. committee and for their invaluable time and advice towards the completion of this work. The author also wishes to thank Dato' Prof. Dr. Syed Jalaludin bin Syed Salim for his invaluable advice and encouragement. The author would like to thank SEAMEO-SEARCA for the financial assistance given to him to pursue a Master of Science degree, the University of Jambi for granting permission for a leave of absence to undertake this study and the Department of Animal Science, Faculty of Veterinary Medicine and Animal Science, Universiti Pertanian Malaysia for providing the research facilities. ii

The author is also grateful to Dr. 1. V. Nolan of the Animal Science Department, University of New England, Armidale, Australia for his advice, comments and suggestions in the urea kinetics study. Sincere appreciation is extended to all the staff of the rumen microbiology laboratory and the animal nutrition laboratory for their cooperation and assistance. Finally, the author would like to express his sincere gratitude and affection to his loving wife Dra. Mardiani, Apt. for her love, prayers, patience and encouragement during the period of this challenging work. And above all, to Almighty God Allah S.W.T., for all blessings on the author and his family. iii

TABLE OF CONTENTS Page ACNOWLEDGEMENTS... ii LIST OF TABLES... vii LIST OF FIGURES... ix ABSTRACT... xi ABSTRAK... xiv CHAPTER I INTRODUCTION... II LITERATURE REVIEW... Goat and Sheep Production............ 1......... 3...... 3 Agricultural By-Products as Feed Material for Ruminant Animals...... 6 Digestion in Ruminant Animals.... Rumen Microbial Population... Bacteria...................... 11.................. 13................................. 14 Protozoa............ 15 Fungi............... 17 Characteristics of Rumen Fermentation............. 18 Rumen Flow Rate.............. 19 Urea-Nitrogen Metabolism......... 20 IV

III GENERAL MATERIALS AND METHODS... 24 Location..................... 24 Animals................. 24 Diets...... 25 Chemical Analyses. Dry Matter... Ash and Organic Matter... Crude Protein... Crude Fiber... Gross Energy... Neutral Detergent Fiber... Acid Detergent Fiber...... 27... 27..... 28...... 28........ 29.......... 30........ 31... 32 IV DIGESTION, RUMEN FERMENTATION, LIQUID AND SOLID RUMENFLOW RATE AND MICROBIAL POPULATION IN GOAT AND SHEEP................ 33 Materials and Methods................. 34 Animals and Diet......... 34 In situ Degradability... 34 In vivo Digestibility.......... 37 Rumen Fermentation Activity... 38 Solid and Liquid Flow Rate... 41 Microbial Population....... 45 Statistical Analysis......... 48 Results........... 48 Discussion....... 78 V UREA KINETIC STUDY IN GOAT AND SHEEP....... 88 Material and Methods......... 89 Animals and Diet... 89 Tracer Solution....... 89 v

Procedures for Urea Kinetic Study.......... 90 Analysis of 1 4 C in Plasma............ 91 Analysis of Urea in Plasma and Urine Samples..... 91 Isolation of CO2........................... 92 BaC03 Precipitation and Purification... 92 Precipitate Solubilisation...................... 93 Radioactivity Measurement................... 93 Calculation... Statistical Analysis......... 94....................... 95 Results... 96 Discussion...................... 109 VI GENERAL DISCUSSION AND CONCLUSION.. General Discussion.................... 114 Conclusion........................ 121......... 114 BIBLIOGRAPHY. APPENDIX : Additional Tables... BIOGRAPHICAL SKETCH......................................... 123................................. 138.................... 141 vi

LIST OF TABLES TaMe Page 1. Goats and Sheep Production in Malaysia........... 5 2. Composition of Diet.................. 25 3. Chemical Composition of Feed Materials...... 26 4. Supplement Intake (g/day)...... 27 5. Dry Matter (DM), Nitrogen (N), Organic Matter (OM), and Energy (E) Intake of Goats and Sheep Fed SSB or SSG Diets... 49 6. Potential Degradability (a+b) and Degradation Rate (c) ofdm, OM and NDF of Rice Straw After 72 h Incubation in Rumen of Goats and Sheep Fed SSB or SSG Diets...... 51 7. Potential Degradability (a+b) and Degradation Rate (c) of DM and N of SBM After 48 h Incubation in Rumen of Goats and Sheep Fed SSB or SSG Diets......... 56 8. Digestibility of DM, CP, OM, NDF, ADF and E of SSB and SSG Diets in Goats and Sheep............ 60 9. Rumen Fluid ph, Ammonia and VFAs Concentration of Goats and Sheep Fed SSB or SSG Diets......... 62 10. Dilution Rate Constant (k), Water Intake (WI), Rumen Volume (RV), Mean Retention Time (MRT) and Liquid Out Flow Rate (FR) of Goats and Sheep Fed SSB or SSG Diets......... 72 vii

11. Dry Matter Intake (DMI), Passage Rate Constant (k), Pool Size (PS), Mean Retention Time (MRT) and Out Flow Rate of Particle (FR) of Goats and Sheep Fed SSB or SSG Diets....................................... 75 12. Bacterial and Protozoal Populations in the Rumen of Goats and Sheep Fed SSB or SSG Diets...... 77 13. Plasma Urea N, Urea N Pool Size, Urea N Net Flux, Fraction of Urea C From Blood to Rumen, Urea N Degradation in the Gut and Urea Excretion in the Urine of Goats and Sheep Fed SSB or SSG Diets......... 97 14. Correlation Coefficient of Urea Kinetic Parameters in Goats and Sheep...... 102 15. Correlation Coefficient of Urea Kinetic Parameters in SSB and SSG Diets...... 103 16. Roll Tube Medium (Prepared for 300 ml Solution)......... 138 17. Dilution Solution Medium (Prepared for 300 ml Solution)... 18. Methyl Green Formalin Solution (MFS)...... 139... 140 viii

LIST OF FIGURES Figure Page 1. Methods Used to Treat Crop Residues............ 9 2. Nitrogen Transaction in the Rumen................ 21 3. Percentage DM Loss of Rice Straw Incubated in the Rumen of Goats and Sheep Fed SSB or SSG Diets...... 52 4. Percentage OM Loss of Rice Straw Incubated in the Rumen of Goats and Sheep Fed SSB or SSG Diets... 53 5. Percentage NDF Loss of Rice Straw Incubated in the Rumen of Goats and Sheep Fed SSB or SSG Diets... 54 6. Percentage DM Loss of Soybean Meal Incubated in the Rumen of Goats and Sheep Fed SSB or SSG Diets... 57 7. Percentage N Loss of Soybean Meal Incubated in the Rumen of Goats and Sheep Fed SSB or SSG Diets...... 58 8. Rumen ph of Goats and Sheep Fed SSB or SSG Diets..... 63 9. Rumen Ammonia N in Goats and Sheep Fed SSB or SSG Diets.......... 64 10. Total Concentration ofvfa in the Rumen of Goats and Sheep Fed SSB or SSG Diets... 66 11. Molar Percentage of Acetate in the Rumen of Goats and Sheep Fed SSB or SSG Diets................... 67 12. Molar Percentage of Propionate in the Rumen of Goats and Sheep Fed SSB or SSG Diets......... ix......... 68

13. Molar Percentage of Butyrate in the Rumen of Goats and Sheep Fed SSB or SSG Diets... 70 14. Dilution Curve for Co in the Rumen of Goats and Sheep Fed SSB or SSG Diets.............. 71 15. Dilution Curve for Cr in the Rumen of Goats and Sheep Fed SSB or SSG Diets...................... 74 16. Time Course of Isotope Dilution in Goats Fed SSB or SSG Diets... 98 17. Time Course of Isotope Dilution in Sheep Fed SSB or SSG Diets.................. 99 1' Plasma Urea N Concentration in Goats and Sheep Fed SSB or SSG Diets at Different Sampling Times After 14 C-Urea Administration............ 100 19. Relationship Between Net Flux and Urea Degradation in the Gut of Goats and Sheep Fed SSB or SSG Diets... 104 20. Relationship Between Net Flux and Pool Size in Goats and Sheep Fed SSB or SSG Diets........ 105 21. Relationship Between Net Flux and Urea Excretion in the Urine of Goats and Sheep Fed SSB or SSG Diets... 106 22. Relationship Between Net Flux and Urea Degradation in SSB and SSG Diets......... 107 23. Relationship Between Net Flux and Urea Excretion in the Urine in SSB and SSG Diets...... 108 x

Abstract of thesis submitted to the Senate of Universiti Pertanian Malaysia in fulfillment of the requirements for the degree of Master of Science. COMPARATIVE STUDIES ON DIGESTIVE EFFICIENCY AND UREA KINETIC BETWEEN GOAT AND SHEEP BY DARLIS SEPTEMBER, 1995 Chairman Faculty Assoc. Prof. Dr. Norhani Abdullah Veterinary Medicine and Animal Science A study was conducted to compare the digestive efficiency and urea kinetics between Malaysian indigenous goats and sheep fed nce straw + soybean meal (SSB) and rice straw + soybean meal + sago (SSG). The Dry Matter (DM), Nitrogen (N), Organic Matter (OM) and Energy (E) intakes in goats and sheep were similar when fed SSB diet, but with SSG diet, DM intake of goats was higher than sheep, while no significant differences in N, OM and E intakes were observed. xi

With SSB diet, goats could digest ADF significantly (P<O.05) more than sheep, but sheep showed higher (P<O.05) digestibility values for Crude Protein (CP), OM and E than those of goats. However, with SSG diet only digestibility values for E were significantly different between the two animal species. Potential degradability for DM and NDF of straw were higher (P<O.05) in goats than sheep when fed SSB diet. However, goats had lower values (P>O.05) in N potential degradability of Soybean Meal (SBM) than sheep. Studies in rumen fluid parameters showed that ammonia concentration was higher in sheep (382.89 ± 33.76 mg Nil) than goats (363.24 ± 43.42 mg NIl), while ph and total Volatile Fatty Acid (VFA) concentrations were similar between the two species when fed SSB diet. The molar proportion of acetate in goats was higher (79.13 ± 2.95%) than that in sheep (75.84 ± 3.91 %), but the reverse was true for molar proportion of propionate where the values obtained for sheep and goats were 15.57 ± 2.40% and 17.96 ± 2.72%, respectively. No significant difference in molar proportion of butyrate was observed between the two animal species. xii

With SSG diet, a higher concentration of ammonia was exhibited by goats (3 1 0.97 ± 40.52 mg NIl) than by sheep (282.48 ± 28.42 mg NIl). Goats also had a higher proportion of acetate (77.65 ± 3.22%) than sheep (73.96 ± 2.43%), but the proportion of butyrate was lower in goats (7.24 ± 1.02%) than in sheep (9.59 ± 2.27%). There were no significant differences in the dilution rate constant, rumen volume, pool size of small particles (based on bodyweight), mean retention time on both liquid and solid between the two animal species. The bacterial population in the rumen of sheep was found to be higher than that of goats, but the protozoal population was observed to be similar between the two animal species fed SSB and SSG diets. Concentration of plasma urea N, urea N synthesis rate, urea N degradation rate, fraction of urea C from blood to the rumen and urea excretion in the urine were found to be similar between the two species of animal fed both SSB and SSG diets. The results of the present study indicate differences in digestion between goats and sheep. However, the results in the rumen fermentation, solid and liquid flow rate and urea metabolism could not explain fully why differences between the two species exist. xiii

Abstrak tesis yang dikemukakan kepada Senat Universiti Pertanian Malaysia sebagai memenuhi keperluan untuk Ijazah Master Sains KAJIAN PERBANDINGAN MEN GENAl KECEKAPAN PENGHADAMAN SERTA KINETIK UREA DI ANTARA KAMBING DAN BEBIRI OLEH DARLIS SEPTEMBER, 1995 Pengerusi Prof. Madya. Dr. Norhani Abdullah Fakulti Kedoktoran Veterinar dan Sains Peternakan Kajian ini telah dijalankan untuk membandingkan kecekapan penghadaman dan kinetik urea antara kambing dan bebiri baka Malaysia yang diberi makan jerami padi + meal kacang soya (SSB) atau jerami padi + meal kacang soya + sagu (SSG). Pengambilan bahan kering (DM), nitrogen (N), bahan organik (OM), dan tenaga (E) oleh kambing dan bebiri adalah sarna bila diberi makan SSB, Bila diberi makan SSG pengambilan DM pada kambing lebih tinggi daripada bebiri manakala tidak ada perbezaan yang ketara untuk pengambilan N, OM dan E. xiv

Dengan rnakanan SSB, karnbing dapat rnenghadarni ADF lebih tinggi daripada bebiri, rnanakala bebiri lebih tinggi berbanding kambing dalam penghadarnan protein kasar (CP), bahan organik (OM) dan tenaga (E). Manakala, dengan makanan SSG hanya penghadaman E nyata berbeza antara kedua spesies haiwan tersebut. Potensi degradasi DM dan NDF jerarni padi lebih tinggi daripada karnbing berbanding bebiri bila diberi makan SSB. Manakala, karnbing lebih rendah nilainya berbanding bebiri untuk potensi degradasi nitrogen dari meal kacang soya. Kajian terhadap ciri cecair rumen memperlihatkan bahawa kepekatan amonia lebih tinggi daripada bebiri (382.89 ± 33.76 mg Nil) berbanding karnbing (363.24 ± 43.42 mg NIl), manakala ph dan jumlah kepekatan asid lernak rneruap (VF A) adalah sarna di antara kedua spesies haiwan bila diberi rnakan SSB. lumlah molar asetat lebih tinggi pada kambing (79.13 ± 2.95%) berbanding bebiri (75.84 ± 3.91%), tetapi sebaliknya dilihat untuk molar propionat di mana nilai yang diperolehi untuk kambing dan bebiri adalah (15.57 ± 2.40%) dan (17.96 ± 2.72%) masing-masing. Hasil kajian rnenunjukkan tidak ada perbezaan dalam jumlah molar butirat antara kedua spesies haiwan. xv

Dengan makanan SSG, kepekatan ammonia didapati lebih tinggi daripada kambing (310.97 ± 40.52 mg NIl) berbanding bebiri (282.48 ± 28.42 mg Nil). Kambing juga mengandungi peratusan molar asetat (77.65 ± 3.22%) lebih tinggi berbanding bebiri (73.96 ± 2.43%), tetapi nilai butirat lebih rendah pada kambing (7.24 ± 1.02%) berbanding bebiri (9.59 ± 2.27%). Tidak ada perbezaan di antara kedua spesies haiwan dalam kelajuan pengaliran ceeair dan butir-butir kecil digesta daripada rumen, isi padu bendalir, saiz gembleng (pool size) butir-butir keeil digesta rumen (berdasarkan pada berat badan), dan min waktu penyimpanan (MRT). PopuJasi bakteria campuran pada rumen bebiri didapati lebih tinggi berbanding kambing, sedangkan populasi protozoa adalah sarna di antara kedua spesies haiwan bila diberi makan SSB atau SSG. Kepekatan N urea plasma, kadar sintesis N urea, kadar degradasi N urea, pemindahan C urea daripada darah ke rumen dan urea yang dikumuhkan pada urin didapati sarna antara kedua spesies haiwan yang diberi makan SSB atau SSG. Keputusan yang diperolehi daripada kajian ini menunjukkan adanya perbezaan dalam proses penghadaman di antara kambing dan bebiri. Tetapi, xvi

data yang diperolehi daripada fermentasi rumen, pengaliran cecair rumen dan butir-butir kecil digesta daripada rumen dan metabolisme N urea tidak dapat memberi penerangan sepenuhnya terhadap perbezaan penghadaman di antara kedua spesies haiwan tersebut. xvii

CHAPTER I INTRODUCTION Goats and sheep are the most important groups of small ruminants in tropical agriculture. They produce meat, milk, wool and fiber. In the Asian countries the indigenous goats and sheep are valued for their meat (Devendra and Coop, 1982). The indigenous goats and sheep in Malaysia are known as kambing katjang and Malin, respectively, and they are usually reared on small farms where the emphasis is on intensive crop production. Fibrous residues from the crops are important feed materials for goats and sheep. The animals provide a means of converting the residues into useful animal products. Goats and sheep have distinct characteristics and grazing behaviour. Sheep are grazing animals whereas goats prefer to browse. The relative productivity of the two species depends upon the ecotypes involved (Wilson, 1982). Each species exhibits many different ecotypes, but only a few have been studied for their actual and potential productivity in all the relevant traits such as fertility, growth rate and carcass characteristics (Fitzhugh, 1987).

2 Several comparative studies between goats and sheep indicated that there are differences in the ability of these two species in digesting various feeds. Earlier studies using low-quality roughage have shown that goats are better than sheep in the digestibility of most nutrients including fiber (Devendra, 1977; Gihad et ai., 1980). The reason for the difference in the digestibility between these two species is not fully understood. There is thus a need to conduct comparative evaluation on the rumen activities of these two species in order to compare their digestive abilities. The objective of this study is to determine whether goats and sheep di f fer in their digestive efficiencies. To achieve this objective, a series of experiments were carried out to measure nutrient intake, in vivo and in situ digestion rates, liquid and solid out flow rates, rumen liquid characteristics (ammonia, ph, total VF A, acetate, propionate and butyrate), microbial population (bacteria and protozoa) and urea kinetics of the two animal species.

CHAPTER II LITERATURE REVIEW Goat and Sheep Production Goats and sheep belong to the tribe caprini of the family Bovidae in the suborder Ruminantia and the order Artiodactyla. cloven-hoofed ruminants of relatively small size. They are typical The tribe caprini comprises of five genera. The goats belong to two of these genera, the Capra and Hemitragus, the sheep belong to the genus Ovis, while the goatlike sheep or sheep-like goat belong to the other two genera, Ammotragus and Pseudo is, respectively (Wilson, 1991). Goats and sheep have spread all over the continents and inhabit almost all climatic zones, from the arctic circle to the equator (Gall, 1981). Devendra and Coop (1982) reported that, nearly 80% of all goats are found within the latitude 0-40 N, in the arid tropical and sub tropical regions. The majority lives in the belt from Bangladesh through India, Iran and Turkey to the Mediterranean countries and in Africa around Ethiopia and the Sahel. Within the tropics, there are more sheep than goats in the highland ecozones of the semi arid areas in east and north 3

4 Africa, but they are of approximately equal number to the goats in the Sahel. Goats are more important in the humid tropics such as Southern India, South East Asia, West Africa and the Caribbean. This indicates that goats are predominantly found in the drier areas, because these areas contain a wide variety of plants, shrubs and trees, which provide a diverse supply of feed for animals at different times of the year. Goats are adaptable; they graze when grasses are lush and abundant and browse the leaves and trees when the supply of grasses is sparse. This contrasts with sheep which graze almost exclusively on grasses (Wilkinson and Stark, 1987). The goats and sheep are valued for their meat, milk, and fiber. Goats contribute 16.6%, 13.1 % and 16.9% of the total world production of meat, milk and fresh skin respectively, while sheep produce 8.2%, 12.2% and 4.4% of the total world output of meat, milk and wool (Devendra, 1986). In the western countries, goats are mainly reared as dairy animals, but in most Asian and African countries, goats are considered almost exclusively as meat animals. In the developing countries the proportion of total income from goats coming from meat is very high, about 70 to 80%, with the remaining 20 to 30% from milk (Gall, 1981).

In Malaysia, goats and sheep production is mostly carried out by smallholders where the animals are kept in small flocks. They are reared primarily for mutton (Rajion et at., 1993). On the average, sheep production (for mutton) has increased about four fo ld during the last 10 years. The production from goats and sheep are 283 and 125 tons per year, respectively (Table 1). Table 1 Goats and Sheep Production in Malaysia Year Goat Sheep 1981 309.22 64.78 1983 274.49 59.73 1985 273.59 78.31 1987 269.11 128.38 1989 2'83.24 181.00 1991 288.52 234.90 Source: Department of Veterinary Service (1992). The goats reared in Malaysia are the indigenous kambing katjang. This breed is also found in Burma, Thailand, Indonesia, Philippines and

6 Taiwan (Mason, 1981). Devendra and McLeroy (1982) described the kambing katjang from Malaysia as follows: they have short horn, while the ears are short and erect. The colour is usually black, but sometimes with a few white patches. The average height of this small-built goat at withers is 56-65 cm. The average birthweight is 1.5 kg and the adult weights are 25 kg for males and 20 kg for females. This prolific breed is very hardy, well adapted to the humid environment and reared mainly for meat. According to Rajion et al. (1993) the sheep population in Malaysia is made up of the indigenous breed, Malin. The Malin, which is small-built and has a mixture of hair and long wool in assorted colours of white, brown or black is considered indigenous to Malaysia, although it may have originated from the sheep in Tibet via the Yunan province in China. The average height at wither is 46.7 cm and the adult weight is 25-30 kg for males and 20-25 kg for females. Agricultural By-Products as Feed Material for Ruminant Animals Fibrous agricultural residues represent a valuable potential source of ruminant feed in many Asian countries. Kossila (1985) estimated that Asia produces approximately 1,628,882 tons of fibrous crop residues, which