Work capacity and fatigue in draught ruminant animals

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(1)ResearchOnline@JCU. This file is part of the following reference:. Martin, Donna G. (1993) Work capacity and fatigue in draught ruminant animals. Masters (Research) thesis, James Cook University of North Queensland.. Access to this file is available from:. http://researchonline.jcu.edu.au/48278/ If you believe that this work constitutes a copyright infringement, please contact [email protected] and quote http://researchonline.jcu.edu.au/48278/.

(2) WORK CAPACITY ANO FATIGUE IN DRAUGHT RUMINANT ANIMALS. Thesis submitted by Donna G MARTIN BSc (JCU). In November 1993. In partial fulfilment at the requirements for the Degree of Mastar of Science (by Research) In the Department of Biomedical and Tropical Veterinary Sciences cf James Cook University of Nor1h Queensland, Australia.

(3) DECLARATION. I declare that this thesis is. my own wor1< and has not been submitted in any rorm for. another degree°' diploma at any university« other tertiary education. Information derived from the published or unpublished worll of others has been acknowledged in the text and a. list of references is given.. Donna MARTIN November 1993. STATEMENT OF ACCESS. I, the undersigned, the author or this thesis, understand that the James COok University or North Queensland wW make It avai'lable for use within the University libraly and, by microfilm °' other photographic means, allow ac:oess to users in other approved libraries. All users consulting this thesis will have to sign the following statement. In consulting this thesis I agree not to copy or closely paraphrase ii in whole or in part without the written consent of the author; end to make proper written acknowledgment for any a&sistance which I have obtained from it.. Beyond this, I do not wish to place any restriction on access to this thesis.. Donna MARTIN November 1993.

(4) ABSTRACT The aim of this study was to define the work capacity of ruminant animals and to investigate the various factors which could limit or enhance this capacity.. A series of three experimental studies was conducted comparing work capacity and physiological and metabolic response.s to work in untrained and trained animals.. The first experiment Involved the comparison of four untrained and four trained swamp buffalo which were walked on a treadmill at 0.69 m/sec for a maximum period of three hours, while pulling four different draught loads equivalent to either 0, 5, 8 or 11% of their live weight.. In the second experiment, three Indonesian breeds of cattle (six of each), either untrained or trained, were compared. The breeds (Ongole,Bali and Madura) were worked in pairs, each pair pulling a draught load equivalent to 12% of the combined live weight of the pair, while walking around a dirt track at approximately 0.69 mlsec for a maximum period or three hours.. The third experiment involved the comparison of six untrained and six trained merino wethers which were walked on a treadmill at speed.s of either 0.67, 1.04 or 1.38 mlsec for a maximum period of three hours while pulling. a. draught load equiva.l ent to 11 % of their. respective live weight.. In all experiments, changes in body temperature, respiration rate and pulse rate were monitored as well as changes in the concentration of selected blood metabolites in the circulation. The uptake/output of the blood metabolites were also measured in wethers used In the third experiment.. The work capacity of each species was found to increase significantly after only a short period of training; the advantage of training being more ciear1y demonstrated at higher levels of work. Improvements In the cardiovascular system and the oxidative capacity of these animals were responsible for this enhanced work capacity.. Hyperthermia appeared to be the major factor causing the onset of fatigue In the working ruminants studied. The oxygen saturation of blood was reduced and the likely depletion of. ii.

(5) the glycogen reserves of the animals in hyperthermia might also have had a role in the onset of fatigue.. The accumulation of lactate in the blood occurred in animals subjected to heavy wor11 loads. This was more noticeable in the untrained animals and in the Bali and Madura breeds than in the Ongole. However, no acidosis occurred but a condition of mild respiratory alkalosis was observed.. In sheep it was found that the hind-limb muscles did not always produce lactate but in fact took up the metabolite, presumably to use as a fuel. II is likely that this would also have occurred in cattle and buffalo.. It was suggested from evidence presented in this study that under normal wor11ing conditions, it would be unlikely that the wo!X capacity of draught animals would be limited by lactic acidosis; rather, hyperthermla would mos1 probably be the major problem.. iii.

(6) Abstract . . • . . • . • . • . . . . . . . . • • . • . • • • . . . . • • • • • • • . • . . • . • . . • . • . • • . • . . . . . • Ust Of Tablas . . . . . . . . . . • . . . . . . . • . . • . • . . . . . . . . . . . • . . . . . . . . . . • . . . . . . • vl List Of FlgUIW. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • xii Ust Of p hatogra pha • • • • • • • • • • . • • . • . • • • • • • • • • • • . • • . • . • • • • . • • . • . • . . . . . XXbc Ust Of Abbnlv1atlona •..•...•. .•. .. .•• ••••.•••••••••.••.•...•••..••..• XlU(. Publications .........•.••••.•••••••••••••.......• . .............••. xxxll Acknowtldgment . • • • • • • . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • . • • • . . . • • • •. XJOtlll. 1.. General Introduction . . . . . . • • • • • . • • • • • . • • . • . . . • . • . • . . . . • • • • • • • • • • • • 1. 2.. Review Of the Lllitratura . • . . • . . • • • • • . • • • • . . . • • . . . . • . • . . • . • . . • . . . . . . 2. 2.1 Skeletal Muscle: Structure and Contraction 2.1.1 Muscle fibnl types 2.1.2 Enervetics of contraction. 2 5 6. 2.2 Fatigue. 13. 2.2.1 Central fatigue. 13. 2.2.2 Peripheral fatigue 2.2.3 Factors involved in fatigue. 14 14. 2.3 Lactic Acid. 22. 2.3.1 2.3.2 2.3.3 2.3.4 2.3.5 2.3.6. 22 24 26 26. Production of lactic acid Lactic acid transport Removal of lactic acid "Ladate shuttle" Acidosis Anaerobic threshold (An. 27. 28. 2.4 Hyperthermla. 29. 2.4.1 2.4.2 2.4.3 2.4.4 2.4.5 2.4.6. 30. Causes of hyperthermia Control of temperature regulation Coofing mechanisms (heat dissipation) Hypertllermia as a cause or latigue Lactic acid and heat stress Blood llow and heat stress. 2.5 Training 2.5.1 2.5.2 2.5.3 2.5.4 2.5.5 2.5.6. 31 31 3-4 35. 36 37. Cardiovascular system Fat utilisation. 37. Glycogen sparing. 39. Lactate and training Fibre types and size Other effects. 41 41 42. 38. 2.6 Conclusion. 42 iv.

(7) 3A.. A Comparison of Work Capaclly of Untnlined and Trained Buffalo (&.tblt*4. bu09 /ls) • • • • • • • . • . • . • • . • • • • • • • • • • • • • • • • • • • • • • • • • • • . • • . • • . • . • • • "" 3A.1 Introduction. 44. 3A.2 Materials and Methods. 45. 3A.2.1 Location 3A.2.2 Experimental animals. 45 45. 3A.2.3 Equipment. 46. 3A.2.4 Experimental design 3A.2.5 Experimental procedure. 48 49. 3A.2.6 Fat.igue assessment. 51. 3A.2.7 laboratory analyses 3A.2.8 Calculations. 53. 54 56. 3A.2.9 Statistical analysis. 56. 3A.3 Resull.s 3A.3.2 Wori\ capacity and energy expenditure. 56 56. 3A.3.3 Physiological Variables. 59. 3A.3.4 Blood Metabolites. 81. 3A.3.1 Environmental conditions. 38.. 3A.4 Discussion. 101. A Comparison of Work Capacity of UntJalned and Trained Ongole (Sos i'ldiwsj , Ball (Bos .ondekul) and Madura Cows. 106. 3B.1 Introduction. 106. 38.2 Materials and Methods. 107. 3B.2.1 location. 107. 3B.2.2 Experimental animals 3B.2.3 Implements and equipment. 108. 3B.2.4 Experimental design. 114 114. 3B.2.5 Experimental procedure 3B.2.6 Laboratory Analysis. 111. 117. 3B.2.7 Calculations 3B.2.8 Statistical Analysis. 118 118. 3B.3 Results. 118. 3B.3.1 Environmental conditions 3B.3.2 Work capacity and energy expenditure. 118 119. 3B.3.3 Physiological variables 3B.3.3 Packed cell volume. 121 130. 38.3.4 Blood metabolites. 133. 3B.4 Discussion. 139. v.

(8) 3C.. A Comparison Of Work Capacity and Musdl Metabobm Of Mer1no Wethers Walklng at 011\'erent Speeds . . . . . . . . . . . . • . . • . . . • • . • . . • . . . . . . • . • . . • 146 3C.1 Introduction. 146. 3C.2 Materials and Methods. 146. 3C.2.1 Location. 146. 3C.2.2 Experimental animals. 147 149. 3C.2.3 Feed 3C.2.4 Equipment. 150 153. 3C.2.5 Experimental Design. 154. 3C.2.6 Experimental procedure 3C.2.7 Fatigue assessment. 157 157. 3C.2.8 Laboratory analysis 3C.2.9 Calculations. 159. 3C.2.10 Statistical analysis. 159. 3C.3 Results. 160. 3C.3.1 Environmental conditions. 160. 3C.3.2 Work capacity 3C.3.3 Physiological variables. 161. 3C.3.4 Blood metabolites. 162 174. 3C.3.5 Blood parameters. 215. 3C.4 Discussion. 4.. 246. General Dlscuaslon . . . • • . . . . . . . . . • . • • . • . . . . • . . . . • • . . . . • . . • . • • . • . 256. References. 261. vi.

(9) UST OF TABLES. Page. Table 2.1. Human skeletal muscle fibre types and their properties. 5. Table 2.2. Yields of adenosine triphosphate (ATP) from anaerobic and aerobic pathways. 7. Table 2.3. Table 2.4. Table 2.5. Table 2.6. Metabolite concentration in the blood and the maximum contribution to the oxidation of skeletal muscle in the hind limb of the fed sheep and humans. 11. The maximum contribution of circulating and endogenous meta.bolites to oxidation in skeletal muscle during sustained exercise. 11. The contribution of Na•. K•-ATPase to muscle energy expenditure in vitro as detennined by 0 2 consumption and by microcalorimetry. 18. The pH values of blood associated with the onset of fatigue in different species. 27. Critical rectal temperature {°C) and behavioural changes in different animals. 33. The composition of rice straw and total dietary nitrogen content.. 46. Table 3A.2. Score card showing fatigue assessment parameters. 51. Table 3A.3. Fatigue score card for draught animals. 52. Table 3A.4. Mean duration of work, distanced travelled, energy expenditure (EE) and EE:Maintenance energy requirement (Mm) ratio of untrained (U) and trained (T) buffalo pulling four different loads. 57. Means of pooled data on duration of work, distance travelled, energy expenditure (EE) and EE:Maintenance energy requirement (Mm) ratio of buffalo for physiological states and work loads. 58. Mean fatigue scores for untrained and trained buffalo pulling four different loads.. 58. Means of pooled data on skin temperature (ST) of buffalo or physiological states, work loads and work status. 60. Means of pooled data on rectal temperature (RT) of buffalo or physiological states, work loads and work status. 62. Table 2.7. Table 3A.1. Table 3A.5. Table 3A.6. Table 3A.7. Table 3A.8. vii.

(10) Means of pooled data on respiration rate (RR) of buffalo for physiological states, work loads and work status. 64. Means of pooled data on pulse rate (PR) of buffalo for physiological states, work loads and work status. 64. Means of pooled data on packed cell volume (PCV) of buffalo for physiological states, work loads and work status. 71. Means of pooled data on haemoglobin concentration of buffalo for physiological states, work loads and work status. 72. Table 3A.13. Means of pooled data on pH in blood of buffalo for physiological states, work loads and work status. n. Table 3A.14. Means of pooled data on lactate concentration in plasma of buffalo for physiological states, work loads and work status. 82. Means of pooled data on glucose concentration in plasma of buffalo for physiological states, work loads and work status. 83. Means of pooled data on urea concentration in plasma of buffalo for physiological states, work loads and work status. 85. Means of pooled data on free fatty acids (FFA) concentration In plasma of buffalo for physiological states, work loads and work status. 86. Means of pooled data on oxygen concentrations in blood of buffalo for physiological states, work loads and work status. 91. Means of pooled data on total cwbon dioxide (TC02) concentrations in blood of buffalo for physiological S1ates, work loads and work status. 93. Means of pooled data on bicarbonate (HC0 3) concentrations In blood of buffalo for physiological states, work loads and work status. 95. Means of pooled data on partial pressure of cart>on cfioxide (pC02) 1n blood of buffalo for physiological states, work loads and work status. 97. The dry matter (OM), protein, CNde fibre (CF). ether extracilves and ash content of elephant gras.s and concentrate fed to the experimental animals. 110. The angle of pull (AP) and speed of walking (WS) while polling a load equivalent to 12% of LW for Ongole, Ball and Madura cows. 113. Means of duration of work, distance travelled, energy expenditure (EE) and EE:Malntenance energy requirement (Mm) ratio of untrained (UNT) and trained (T) Ongole, Ball end Madura cows. 119. Table 3A.9. Table 3A.10. Table 3A.11. Table 3A.12. Table 3A. 15. Table 3A.16. Table 3A.17. Table 3A.18. Table 3A.19. Table 3A.20. Table 3A-21. Table 3B.1. Table 3B.2. Table 38.3. vlil.

(11) Means or pooled data on duration or woltt and distance travelled in cattle for physiologiem swtes and breeds. 120. Mean fatigue scores tor untrained and trained Ongole, Bd and Maduni cows. 120. Means of pooled data on skin temperature (Slj of cattle for physiological slates, breeds and wont status. 122. Means or pooled data on rectal temperature (Rlj or cattle for physiological states, breeds and wont status. 124. Means of pooled data on respiration rate (RR) of cattle for physiological states, breeds and wont status. 125. Means of pooled data on pulse rate (PR) of cattle for physiological states, breeds and work status. 127. Rate of decline of pulse rate and slope 1O minutes immediately wont ceased acroH physiological states and breeds. 128. Means or pooled data on paclted cell volume (PCV) of cattle for physiological states, breeds and wont status. 130. Means or pooled data on lactate concentration in plasma of cattle for physiological states, breeds and wont status. 134. Means of pooled data on glucose concentration in plasma or cattle for physlological states, breeds and wont status. 135. Means of pooled data on urea conc:e11bation in plasma of cattle for physiological states, breeds and wont status. 137. Means or pooled data on free fatty acids (FFA) concentration in plasma or cattle for physiological states, breeds and work status. 138. Peak values for plasma lactate In O ngole, Bali and Madura cows and 11/ne of onset of blood lac:sate ac:c:umulation (OBLA). 141. The increases in skin and recta.1temperatures of untrained and trained Ongole, Bali and Madura cows during the Worlc period. 142. Peak glucose concentrations in plasma or Ongole, Bali and Madura cows during the Worlc period. 144. Table 3C.1. The composition of sorghum hay. 150. Table 3C.2. The c:omposrtion of the Rumevite mineral stodt block. 150. Table 3C.3. Mean walking speed, duration and distance travelled by untrained and trained Merino wethers. 161. Table 3B.4. Ta.ble 3B.5. Table 3B.6. Table 3B.7. Table 3B.8. Table 3B.9. Ta.ble 3B.10. Table 38.11. Table 3B.12. Table 3B.13. Table 3B.14. Table 3B.15. Teble 38.16. Table 3B.17. Table 38.18. ix.

(12) Table 3C.4. Table 3C.5. Table 3C.6. Table 3C.7. Table 3C.8. Table 3C.9. Table 3C.10. Table 3C.11. Table 3C.12. Table 3C.13. Table 3C.14. Table 3C.15. Table 3C.16. Table 3C.17. Means of pooled data on duration of work and distance travelled in sheep across physiological states and walking speeds. 161. Pooled data on energy expenditure (EE) and EE:Maintenance energy requirement (Mm) of sheep across walking speeds. 162. Means of pooled data on skin temperature (ST) of sheep for physiological states, walking speeds and work status. 163. Means of pooled data on redal temperature (RT) of sheep for physiological states, walking speeds and work status. 166. Means of pooled data on respiration rate (RR} of sheep for physiological state.s, walking speeds and work status. 168. Means of pooled data on pulse rate (PR) of sheep for physiological states, walklng speeds and work status. 170. Rate of decline in pulse rate (PR) during the first 4 minutes of the Recovery period and shape of lines across physiological states and walking speeds. 172. Comparison of arterial concentration (A} and arterio-venous concentration difference (A-V) of ladate in plasma of sheep, between physiological states, walking speeds and work status. 176. Comparison of arterial concentrations (A) and arterio-venous concentration difference (A-V) of glucose in plasma of sheep, between physiological states, walking speeds and work status. 180. Comparisons of arterial concentration of urea in plasma of sheep, between physiological states, walking speeds and work status. 184. Comparison of arterial concentration (A) and arterio-venous concentratlon difference (A-V) of free fatty acids (FFA) in plasma of sheep, between physiological states, walking speeds and work status. 186. Comparison of arterial concentrations (A)and arterio-venous concentration difference (A-V) of crealinine in plasma of sheep, between physiological states, walking speeds and work status. 193. Comparison of arterial concentrations (A) and arterio-venous concentration difference (A-V) of potassium in plasma of sheep, between physiological states, walking speeds and work status. 200. Comparisons of arterial concentrations (A) and arterio-venous concentration difference (A-V) of ammonia-nitrogen (NH3-N) in plasma of sheep, between physiological states, walking speeds and work status. 207. x.

(13) Table 3C.16. Table 3C.19. Table 3C.20. Table 3C.21. Table 3C.22. Table 3C.23. Table 3C.24. Table 3C.25. Comparison of arterial concentrations (A) and arterio-venous concentration difference (A-V) of protein/amino acid-nitrogen (Protein/AA-N) in plasma of sheep, between physiological states, walking speeds and wont status. 211. Means of pooled data on packed cell volume (PCV) of sheep, for physiological states, walking speeds and wont status. 216. Comparison or arterial concentrations (A) and arterio-venous concentration (A-v) of haemoglobin in blood of sheep, between physiological states, walking speeds and wont status. 217. Comparison of arterial concentrations (A) and arterio-venous concentratlon difference (A-V) of pH in blood of sheep, between physiological states, walking speeds and wont status. 225. Comparison of arterial concentrations (A) and arterio-venous concentration differences (A-V) of oxygen saturation (02SAT) In blood of sheep, between physiological states, walking speeds and wont status. 231. Comparison of arterial concentrations (A) and arterio-venous concentration differences (A-V) or total carbon dioxide (TC0 2) in blood of sheep, between physiological states, walking speeds and wont status. 235. Comparison of arterial concentrations (A) and arterio-venous concentration differences (A-V) of bicart>onate in blood of sheep, between physiological states, walking speeds and wont status. 237. Comparison of arterial partlal pressure of carbon dioxide (pC02) in blood of sheep, be~n physiological states, walking speeds and wor1t status. 244. -xi.

(14) UST OF FIGURES. Page. Figure 2.1. Figure 2.2. Figure 2 .3. Figure 2.4. Figure 2.5. Figure 2.6. Figure 2.7. Figure 2 .8. Figure 2.9. Figure 2.10. Figure 2.11. Figure 2.12. Figure 2.13. Figure 2.14. Diagram of muscle structure showing the arrangements of fibres in a striated muscle. The cross-striations on the myoflbrils can be seen with light microscopy.. 2. Sliding is probably brought about, by a rotation of the S 1 subunits about their points of attachment to the thin filament (actin).. 3. Schematic representation of the coupling process in skeletal muscle.. 4. Diagram showing substrates for ATP production via the TCA cycle and respiratory chain.. 7. Muscle concentrations of ATP, ADP, AMP during exercise in dogs.. 8. Muscle concentrations of creatine phosphate and creatine during exercise in dogs.. 9. Muscle concentrations of glycogen and glucose during exercise in dogs.. 9. Muscle concentrations of pyl\IVate and lactate during exercise in dogs.. 10. K" concentrations in humans (arm) with increasing exercise intensity (% V 0 2 max) and duration of exercise.. 18. The effect of exercise Intensity on the mean concentration of lactate in arterial blood of five sheep exercised for 25 minutes.. 25. The effect of worlt rate 0N) on the concentration of lactate in venous blood.. 25. Regression lines showing anaerobic threshold from the inflection point of the V-slope, co2 production (VC02) versus 0 2 uptake (V02).. 29. Diagrams showing (A) the carotid rete and (B) Circle of Willis, cavenous sinus and nasal cavity Illustrating the direction of venous and arterial bloodflow.. 32. Regression of muscle lactate (LA) content on muscle temperature in dogs.. 35. Mean increases in plasma. xii.

(15) Figure 2.15. Figure 2.16. Figure 2 .17. Figure 3A.1. Figure 3A.2. Figure 3A.3. Figure 3A.4. Figure 3A.5. Figure 3A.6. Figure 3A.7. Figure 3A.8. Figure 3A.9. Figure 3A.10. -.. Data on the distribution of blood to various tissues in sheep during exercise and mild heat stress.. 37. Blood flow and oxygen uptake in the trained (1) and nontrained (NT) leg muscle of humans.. 38. Contribution of carbohydrate (CH) and fat oxidation to the total leg oxygen uptake in the trained (1) and the nontrained (NT) leg during the post-training metabolic study.. 40. The daily time schedule for the Pre-worl<, Worl< and Recovery periods for each pair of animals during the fourday measurement period.. 48. Relationship between angle force (AF), draught force (DF), angle of pull (a) and load.. 55. Mean fatigue score (0-20) for loads 1 to 4 for untrained and trained buffalo.. 59. Mean skin temperature of untrained and trained buffalo under different work status (Pre-worl<, Worlc and Recovery) and work loads (1 -4) where 1-4 are equivalent to 0, 5, 8 and 11 % of live weight respectively.. 60. Means of pooled data on skin temperature of buffalo during the Pre-worl<, Worl< and Recovery periods.. 61. Mean rectal temperature of untrained and trained buffalo under different work status (Pre-worlc, Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11 % of live we.ight respectively.. 62. Mean respiration rate of untrained and ttalned buffalo under different work status (Pre-worl<, Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, B and 11 % of live weight respectively.. 63. Mean pulse rate of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery) and work loads (1-4) whare 1-4 are equivalent to 0, 5, 8 and 11 o/o of live weight re.spectively.. 65. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of untrained buffalo over work time across work status (Worlc and Recovery) at a work load equivalent to 0% of live weighl. 66. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 0% of live weight.. 66. xiii.

(16) 'igure 3A.11 .. =igure 3A.12. Figure 3A.13. Figure 3A.14. Figure 3A. 15. Figure 3A.16. Figure 3A.17. Figure 3A.18. Figure 3A.19. Figure 3A.20. Changes in (A) rectal temperature (RT), (B) skin te!T)perature (ST), pulse rate (PR) and respiration rate (RR) of untrained buffalo over work time across work status (Worlc and Recovery) at a work load equivalent to 5% of live weight.. 67. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of trained buffalo over work time across work status (Worlc and Recovery) at a work load equivalent to 5% of live weight.. 67. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of untrained buffalo over work time across work status (Wol1< and Recovery) at a work load equivalent to 8% of live weight.. 68. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of trained buffalo over work time across work status (Wol1< and Recovery) at a work load equivalent to 8% of live weight.. 68. Changes in (A) rectal temperature (RT), (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of untrained buffalo over work time across work status (Wol1< and Recovery) at a work load equivalent to 11% of live weight.. 69. Changes in (A) rectal temperature (RT) , (B) skin temperature (ST), pulse rate (PR) and respiration rate (RR) of trained buffalo over work time across work status { Worlc and Recovery) at a work load equivalent to 11% of live weight.. 69. Mean packed cell volume of untrained and trained buffalo under different work status (Pre-work, Work and Recover/) and work loads (1 -4) where 1--4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 70. Means of pooled data on packed cell volume of buffalo during the Pre-work, Work and Recovery periods. Vertical lines. 72. Mean concentrations of haemoglobin in blood of untrained and trained buffalo under different work status (Pre-work, Work and Recovery) and work loads (1--4) where 1--4 are equivalent to 0 , 5, 8 and 11% of live weight respectively.. 73. Means of pooled data on concentration of haemoglobin in untrained and trained buffalo. Vertical lines. 74. xiv.

(17) Figure 3A.21. Figure 3A.22. Figure 3A.23. Figure 3A.24. Figure 3A.25. Figure 3A.26. Figure 3A.27. Figure 3A.28. Figure 3A.29. Figure 3A.30. Figure 3A.31. Changes in packed cell volume and haemoglobin concentrations of (A) untrained and (B) trained buffalo over work time across work status (Worlc and Recovery) at a work load equivalent to Oo/o of live weighl. 75. Changes in packed cell volume and haemoglobin concentrations of (A) untrained and (B) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 5% of live weight.. 75. Changes in packed cell volume and haemoglobin concentrations of (A) untrained and (B) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 8% of live weight.. 76. Changes in packed cell volume and haemoglobin concentrations of (A) untrained and (8) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 11% of live weight.. 76. Mean pH of untrained and trained buffalo under different work status (Pre-worlc, Worl< and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 77. Means of pooled data on pH of buffalo during the Pre-worlc, Worlc and Recovery periods.. 78. Changes in the pattern of pH of (A) untrained and (B) trained buffalo over work time across work status (Worlc and Recovery) at a work load equivalent to 0% of live weight.. 79. Changes in the pattern of pH of (A) untrained and (8) trained buffalo over worll time across worll status (Worlc and Recovery) at a work load equivalent to 5% of live weighl. 79. Changes in the pattern of pH of (A) untrained and (B) trained buffalo over worll time across work status (Worlc and Recovery) at a work load equivalent to 8% of live weight.. 80. Changes in the pattern of pH of (A) untrained and (B) trained buffalo over worll time across wor111 status (Worlc and Recovery) at a worll load equivalent to 11 % of live weight.. 80. Mean concentrations of lactate in plasma of untrained and trained buffalo under different work status (Pre-worlc, Worlc and Recovery) and worll loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11 % of live weight respectively.. 81. xv.

(18) Figure 3A.32. Figure 3A.33. Figure 3A.34. Figure 3A.35. Figure 3A.36. Figure 3A.37. Figure 3A.38. Figure 3A.39. Figure 3A.40. Figure 3A.41. Mean concentrations of glucose in plasma of untrained and trained buffalo under different work status (Pre-worl<, Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 83. Mean concentrations of glucose in plasma of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respedively.. 84. Mean concentrations of urea in plasma of untrained and trained buffalo under different work status (Pre-worlc, Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 85. Mean concentrations of free fatty acids in plasma of untrained and trained buffalo under different work status (Pre-worlc. Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11 % of live weight respedively.. 87. Means of pooled data on FFA concentrations in plasma of buffalo during the Pre-worl<, Worlc and Recovery periods.. 88. Changes in plasma lactate (LA). glucose (G), urea (U) and free fatty acids (FFA) concentrations of (A) untrained and (B) trained buffalo over work time across work status (Preworlc, Work and Recovery) at a load equivalent to 0% of live weight.. 89. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acids (FFA) concentrations of (A) untrained and (B) trained buffalo over work time across work status (Prework, Worl< and Recovery) at a load equivalent to 5% or live weight. I. 89. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acids (FFA) concentrations of (A) untrained and (B) trained buffalo over work time across work status (Preworlc, Worl< and Recovery) at a load equivalent to 8% of live weight.. 90. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acids (FFA) concentrations of (A) untrained and (B) trained buffalo over work time across work status (Preworl<, Worlc and Recovery) at a load equivalent to 11% of live weight.. 90. Mean concentrations of oxygen in blood of untrained and trained buffalo under different work status (Pre-worl<, Worlc and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 91. xvi.

(19) Figure 3A.42. Figure 3A.43. Figure 3A.44. Figure 3A.45. Figure 3A.46. Figure 3A.47. Figure 3A.48. Figure 3A.49. Figure 3A.50. Figure 3A.51. Mean concentrations of oxygen in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery).. 92. Mean concentrations of total carbon dioxide in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 93. Mean concentrations of total carbon dioxide in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery).. 94. Mean concentrations of bicarbonate in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 95. Mean concentrations of bicarbonate in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery).. 96. Mean concentrations of partial pressure of carbon dioxide in blood of untrained and trained buffalo under different work status (Pre-worl<, Worl< and Recovery) and work loads (1-4) where 1-4 are equivalent to 0, 5, 8 and 11% of live weight respectively.. 97. Mean concentrations of partial pressure of carbon dioxide in blood of untrained and trained buffalo under different work status (Pre-worl<, Worlc and Recovery).. 98. Changes in concentrations of total carbon dioxide (TC02). oxygen (0 2) and bicarbonate (HC03) in blood of (A) untrained and (B) trained buffalo ovef work time across work status (Worl< and Recovery) at a work load equivalent to 0% of live weight.. 99. Changes in concentrations of total carbon dioxide (TC0 2), oxygen (02) and bicarbonate (HC0 3) In blood of (A) untrained and (B) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 5% of live weight.. 99. Changes in concentrations of total carbon dioxide (TC02), oxygen (0 2) and bicarbonate (HCOJl in blood of (A) untrained and (B) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 8% of live weight.. 100. xvii.

(20) Changes in concentrations of total carbon dioxide (TC02 ) , oxygen (02) and bicarbonate (HCO~ in blood of (A) untrained and (B) trained buffalo over work time across work status (Worl< and Recovery) at a work load equivalent to 11 % of live weighl. 100. The relationship between rectal temperature and respiration rate.. 103. Figure 3A.54. The relationship between respiration rate and pulse rate.. 103. Figure 38.1. The location of Grati in the province of East Java, Indonesia.. 107. The angle force (AF) measured by the load cell (LC) attached between the yoke and the load.. 112. The relationship between draught force (y) and weight of load (x).. 113. Diagram of square track showing location of sampling and environmental stations.. 115. Mean fatigue score (0-20) for untrained and trained Ongole, Bali and Madura cows.. 121. Mean skin temperature of untrained and trained Ongole, Bali and Madura cows under different work status (Prework, Work and Recovery).. 122. Mean rectal temperature of untrained and trained Ongole, Bali and Madura cows under different work status (Prework, Work and Recovery).. 123. Mean respiration rate of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery).. 125. Mean pulse rate of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery).. 126. Mean pulse rate decline during Recovery in (A) untrained and (B) trained Ongole, Bali and Madura cows.. 128. Changes in rectal temperature (RT), skin temperature (ST), pulse rate (PR) and respiration rate (RR) of (A) untrained and (8) trained Ongole cows over work time across work status (Pre-work, Work and Recovery).. 129. Changes in rectal temperature (RT), skin temperature (ST), pulse rate (PR) and respiration rate (RR) of (A) untrained and (8) trained Bali cows over work time across work status (Pre-work, Work and Recovery) .. 129. Figure 3A.52. Figure 3A.53. Figure 38.2. Figure 38.3. Figure 38.4. Figure 3B.5. Figure 3B.6. Figure 3B.7. Figure 3B.8. Figure 3B.9. Figure 3B.10. Figure 38.11. Figure 3B.12. -.. xviii.

(21) Figure 38.13 · Changes in rectal temperature (Rn, skin temperature (Sn, pulse rate (PR) and respiration rate (RR) of (A) untrained and (8) trained Madura cows over work time across work status (Pre-work, Work and Recovery).. 130. Mean packed cell volume of untrained and trained Ongole, Bali and Madura cows under different work status (Prework, Work and Recovery).. 131. Means of pooled data on packed cell volume of cattle during the Pre-work, Work and Recovery periods.. 132. Changes in packed cell volume of untrained and trained cattle, over work time across work status (Pre-work, Work and Recovery).. 132. Mean concentrations of lactate in plasma of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery) .. 133. Mean concentrations of glucose in plasma of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery) .. 135. Mean concentrations of urea in plasma of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery).. 137. Mean concentrations of free fatty acids in plasma of untrained and trained Ongole, Bali and Madura cows under different work status (Pre-work, Work and Recovery).. 138. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acid (FFA) concentrations of (A) untrained and (B) trained Ongole cows over work time across work status (Pre-work, Work and Recovery).. 140. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acid (FFA) concentrations of (A) untrained and (8) trained Bali cows over work time across work status (Prework, Work and Recovery).. 140. Changes in plasma lactate (LA), glucose (G), urea (U) and free fatty acid (FFA) concentrations of (A) untrained and (B) trained Madura cows over work time across work status (Pre-work, Work and Recovery).. 141. The relationship between respiration rate and rectal temperature.. 143. Figure 3C.1. Treadmill modified to accommodate sheep.. 151. Figure 3C.2. The daily time schedule for the Pre-work, Work and Recovery periods for each pair of animals during the three day measurement period.. 153. Figure 38.14. Figure 38.15. Figure 38.16. Figure 38.17. Figure 38.18. Figure 38.19. Figure 38.20. Figure 38.21. Figure 38.22. Figure 38.23. Figure 38.24. --. xix.

(22) Figure 3C.3. Figure 3C.4. Figure 3C.5. Figure 3C.6. Figure 3C.7. Figure 3C.8. Figure 3C.9. Figure 3C.10. ~igure. 3C.11. Figure 3C.12. Figure 3C.13. Figure 3C.14. Figure 3C.15. --. Diagram illustrating the point at which the heparin enters the blood line.. 156. Means of ambient temperature (AmT), black bulb (88) temperature and relative humidity (RH) during the measurement period.. 160. Means of pooled data of skin temperature of untrained and trained sheep during the Pre-worlc, Worlc (1 hour) and Recovery periods and at the slow and medium walking speeds.. 163. Means of pooled data on skin temperature of trained sheep during the pre-worlc, Worlc (30 minutes) and Recovery periods and at slow, medium and fast walking speeds.. 164. Means of pooled data on rectal temperature of untrained and trained sheep during the Pre-worlc, Worlc (1 hour) and Recovery periods and at slow and medium walking speeds.. 165. Means of pooled data on rectal temperature of trained sheep during the Pre-worlc, Worlc (30 minutes) and Recovery periods and at slow, medium and fast walking speeds.. 166. Means of pooled data on respiration rate of untrained and trained sheep during the Pre-worlc, Worlc (1 hour) and Recovery periods and at slow and medium walking speeds.. 167. Means of pooled data on respiration rate of trained sheep during the Pre-worlc, Worlc (30 minutes) and Recovery periods and at slow, medium and fast walking speeds.. 168. Means of pooled data on pulse rate of untrained and trained sheep during the Pre-worlc, Worlc (1 hour) and Recovery periods and at slow and medium walkil'.lg speeds.. 169. Means of pooled data on pulse rate of trained sheep during the Pre-worlc, Worlc (30 minutes) and Recovery periods and at slow, medium and fast walking speeds.. 171. Means of pooled data on pulse rate decline in (A) untrained and (B) trained sheep during the first 10 minutes of the Recovery period across walking speeds.. 172. Changes in rectal temperature (RT), skin temperature (ST), pulse rate(PR), and respiration rate (RR) in (A) untrained and (B) trained sheep over time across work status (Preworlc, Worlc, and Recovery) at slow speed.. 173. Changes in rectal temperature (RT), skin temperature (ST), pulse rate (PR), and respiration rate (RR) in (A) untrained and (B) trained sheep over time across work status (Pre- . worlc, Worlc, and Recovery) at medium speed.. 173. xx.

(23) Figure 3C.16. Figure 3C.17. Figure 3C.18. Figure 3C.19. Figure 3C.20. Figure 3C.21. Figure 3C.22. Figure 3C:23. Figure 3C.24. Figure 3C.25. Figure 3C.26. -. '. Changes in rectal temperature (RT), skin temperature (ST), pulse rate (PR), and respiration rate (RR) in trained sheep over time across work status (Pre- work, Work, and Recovery) at fast speed.. 174. Mean concentrations of lactate in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 175. Mean arterio-venous concentration difference of lactate in plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 175. Mean concentrations of lactate in arterial plasma of trained and untrained sheep during the Pre-work, Worl< (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 177. Mean arterio-venous concentration difference of lactate in plasma of trained and untrained sheep during the Pre-worl<, Worl< and Recovery periods and at slow, medium and fast walking speeds. Vertical bars above lines represent standard errors.. 177. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of lactate in untrained (A) and trained (B) sheep during the first 20 minutes of the Worl< period for slow, medium and fast walking speeds.. 179. Mean concentrations of glucose in arterial piss.ma of untrained and trained sheep during the Pre-work, ~orl< (1 hc;>ur) and Recovery periods and at the slow and medium walking speeds.. 181. Mean arterial-venous concentration difference of glucose in plasma of untrained and trained sheep during the Pre-work, Worl< (1 hour) and Recovery periods and at the slow and medium walking speeds.. 181. Mean concentration of glucose in arterial plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 183. Mean arterio-venous concentration difference of glucose in plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 183. Mean concentration of urea in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 184. xxi.

(24) Figure 3C.27. Figure 3C.28. Figure 3C.29. Figure 3C.30. Figure 3C.31. Figure 3C.32. Figure 3C.33. Figure 3C.34. Figure 3C.35. Figure 3C.36. Mean concentration of urea in arterial plasma of trained sheep during the Pr&-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 185. Mean concentration of free fatty acids in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 187. Mean arterio-venous concentration difference of free fatty acids in plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 187. Mean concentration of free fatty acids in arterial plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and of the slow, medium and fast walking speeds.. 189. Mean arterio-venous concentration difference of free fatty acids in plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 189. The concentration in arterial· plasma and the arterio-venous (A-V) concentration difference of lactate (LA), glucose (G), urea (U) and free fatty acids (FFA) in untrained (A) and trained (8) sheep over time across work status (Pre-work, Work and Recoveryr at the slow walking speed.. 190. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of lactate (LA), glucose (G), urea (U), and free fatty acids (FFA)in untrained (A) and trained (8) sheep over time across work status (Pre-work, Work and Recovery) at the medium walking speed.. 191. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of lactate (LA), glucose (G), urea (U) and free fatty acids (FFA) in trained sheep over time across work status (Pre-work, Work and Recovery) at the fast walking speed.. 192. Mean concentration of creatinine in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 194. Mean arterio-venous concentration difference of creatinine in plasma of untrained and trained sheep during the Prework, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 194. xxii.

(25) Figure 3C.37. Figure 3C.38. Figure 3C.'3 9. Figure 3C.40. Figure 3C.41. Figure 3C.42. Figure 3C.43. Figure 3C.44. Figure 3C.45. Figure 3C.46. Figure 3C.47. Mean concentration of creatinine in arterial plasma of trained sheep during the Pre-work, Work (30 minutes} and Recovery periods and at the slow, medium and fast walking speeds.. 196. Mean arterio-venous concentration difference of creatinine in plasma of trained sheep during the Pre-work, Work (30 minutes} and Recovery periods and at the slow, medium and fast walking speeds.. 196. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of creatinine in untrained (A} and trained (B} sheep over time across work status (Prework, Work and Recovery) at the slow walking speed.. 197. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of creatinine in untrained (A) and trained (B} sheep over time across work status (Prework, Work and Recovery) at the medium walking speed.. 198. The concentration in arterial plasma and arterio-venous (AV) concentration difference of creatinine in trained sheep over time across work status (Pre-work, Work and Recovery} of the fast walking speed.. 199. Mean concentration of potassium in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 201. Mean arterio-venous concentration difference of potassium in plasma of untrained and trained sheep during the Prework, Work (1 hour} and Recovery periods and at the slow and medium walking speeds.. 201. Mean concentration of potassium in arterial plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 203. Mean arterio-venous concentration difference of potassium in plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 203. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of potassium in untrained (A) and trained (B} sheep over time across work status (Prework, Work and Recovery) of the slow walking speed.. 204. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of potassium in untrained (A} and trained (B) sheep over time across work status (Prework, Work and Recovery) of the medium walking speed.. 205. xxiii.

(26) Figure 3C.48. Figure 3C.49. Figure 3C.50. Figure 3C.51. Figure 3C.52. Figure 3C.53. Figure 3C.54. Figure 3C.55. Figure 3C.56. Figure 3C.57. Figure 3C.58. The concentration in arterial plasma and arterio-venous (AV) concentration difference of potassium in trained sheep over time across woric status (Pre-work, Work and Recovery) at the fast walking speed.. 206. Mean concentration of ammonia-nitrogen in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 208. Mean arterio-venous concentration difference of ammonianitrogen in plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 208. Mean concentration of ammonia-nitrogen in arterial plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 210. Mean arterio-venous concentration difference of ammonianitrogen in plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 210. Mean concentration of protein/amino acid-nitrogen in arterial plasma of untrained and trained sheep during the Pre-work, Worl< (1 hour) and Recovery periods and at the slow and medium walking speeds.. 212. Mean arterio-venous concentration difference of protein/amino acid-nitrogen in plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 212. Mean concentration of protein/amino acid-nitrogen in arterial plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 214. Mean arterio-venous concentration difference of protein/amino-nitrogen in plasma of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 214. Mean packed cell volume in untrained and trained sheep during the Pre-work, Work and Recovery periods and at the slow and medium walking speeds.. 215. Packed cell volume in untrained and trained sheep over time across woric status (Pre-work, Work and Recovery) at slow (A) and medium (8) walking speeds.. 216. --. xxiv.

(27) Figure 3C.59.. Figure 3C.60. Figure 3.C.61. Figure 3C.62. Figure 3C.63. Figure 3C.64. Figure 3C.65. Figure 3C.66. Figure 3C.67. Figure 3C.68. Figure 3C.69. Mean concentration of haemoglobin in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 218. Mean arterio-venous concentration difference of haemoglobin in plasma of untrained and trained sheep during the Pre-work, Worl< (1 hour) and Recovery periods and at the slow and medium walking speeds.. 218. Mean concentration of haemoglobin in arterial plasma of trained and untrained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the sJow, medium and fast walking speeds.. 220. Mean arterio-venous concentration difference of haemoglobin in plasma of trained and untrained sheep during the Pre-worl<, Worl< and Recovery periods and at slow, medium and fast walking speeds. Vertical bars above lines represent standard errors.. 220. The concentration in arterial blood and the arterio-venous (A-V) concentration difference of haemoglobin in untrained (A) and trained (B) sheep over time across work status (Pre-worl<, Worl< and Recovery) at the slow walking speed.. 221. The concentration in arterial blood and the arterio-venous (A-V) concentration difference of haemoglobin in untrained (A) and trained (8) sheep over time across work status (Pre-worl<, Worl< and Recovery) at. the medium walking speed.. 222. The concentration in arterial blood and the arterio-venous (A-V) concentration difference of haemoglobin in trained sheep over time across work status (Pre-worl<, Worl< and Recovery) at the fast walking speed.. 223. Mean pH in untrained and trained sheep during the Prework, Worl< (1 hour) and Recovery periods and at the slow and medium walking speeds.. 224. Mean arterio-venous difference in pH of blood of untrained and trained sheep during the Pre-worl<, Worl< (1 hour) and Recovery periods and at the slow and medium walking speeds.. 224. Mean concentration of pH of blood in arterial plasma of trained and untrained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 226. Mean arterio-venous concentration difference in pH of blood in trained sheep during the Pre-worl<, Worl< (30 minutes) and Recovery periods and at slow, medium and fast walking speeds.. 226. xxv.

(28) Figure 3C. 70.. Figure 3C.71. Figure 3C.72. Figure 3C. 73. Figure 3C.74. Figure 3C. 75. Figure 3C.76. Figure 3C. 77. Figure 3C.78. Figure 3C.79. Figure 3C.80. The concentration of pH in arterial blood and the arteriovel")ous (A-V) concentration difference in untrained (A) and trained (B) sheep over time across work status (Pre-work, Work and Recovery) at the slow walking speed.. 227. The concentration of pH in arterial blood and the arteriovenous (A-V) concentration difference in untrained {A) and trained (B) sheep over time across work status {Pre-work, Worlc and Recovery) at the medium walking speed.. 228. The concentration of pH in arterial blood and the arteriovenous (A-V) concentration difference in trained sheep over time across work status {Pre-work, Work and Recovery) at the fast walking speed.. 229. Mean saturation of oxygen in arterial blood of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 230. Mean arterio-venous saturation difference of oxygen in blood of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 230. Mean saturation of oxygen in arterial blood of trained sheep during the Pre-work, Work (30 minutes) and Recovery period and at slow, medium and fast walking speeds.. 232. Mean arterio-venous saturation difference of oxygen in blood of trained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 232. Mean concentration of total carbon dioxide in arterial plasma of untrained and trained sheep during th~ Pre-work, Worlc (1 hour) and Recovery periods and at the slow and medium walking speeds.. 234. Mean arterio-venous concentration difference of total carbon dioxide in plasma of untrained and trained sheep during the Pre-worlc, Worlc (1 hour) and Recovery periods and at the slow and medium walking speeds.. 234. Mean concentration of total carbon dioxide in arterial plasma of trained and untrained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 236. Mean arterio-venous concentration difference of total carbon dioxide in plasma of trained and untrained sheep during the Pre-worlc, Worlc (30 minutes) and"Recovery periods and at slow, medium and fast walking speeds. Vertical bars above lines represent standard errors.. 236. xxvi.

(29) Figure 3C.81. Figure 3C.82. Figure 3C.83. Figure 3C.84. Figure 3C.85. Figure 3C.86. Figure 3C.87. Figure 3C.88. Figure 3C.89. Figure 3C.90. Mean concentration of bicarbonate in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking ·speeds.. 238. Mean arterio-venous concentration difference of bicarbonate in plasma of untrained and trained sheep during the Prework, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 238. Mean concentration of bicarbonate in arterial plasma of trained and untrained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 240. Mean arterio-venous concentration difference of bicarbonate in plasma of trained and untrained sheep during the Prework, Work and Recovery periods and at slow, medium and fast walking speeds. Vertical bars above lines represent standard errors.. 240. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of oxygen saturation (02 SAT), total carbon dioxide (TC0 3) and bicarbonate (HC0 3) in untrained (A) and trained (B) sheep over time across work status (Pre-work, Work and Recovery) at the slow walking speed.. 241. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of oxygen saturation (02SAT), total carbon dioxide (TC03) and bicarbonate (HCO~ in untrained (A) and trained (B) sheep over time across work status (Pre-work, Work and Recovery) _a t the ~edium walking speed.. 242. The concentration in arterial plasma and the arterio-venous (A-V) concentration difference of oxygen saturation (02 SAT), total carbon dioxide (TC03) and bicarbonate (HC03) in trained sheep over time across work status (Prework, Work and Recovery) at the fast walking speed.. 243. Mean concentration of partial pressure of carbon dioxide (pC02) in arterial plasma of untrained and trained sheep during the Pre-work, Work (1 hour) and Recovery periods and at the slow and medium walking speeds.. 244. Mean concentrations of partial pressure of carbon dioxide (pC02) in arterial plasma of trained and untrained sheep during the Pre-work, Work (30 minutes) and Recovery periods and at the slow, medium and fast walking speeds.. 245. Changes in partial pressure of carbon dioxide (pC02) in arterial blood in untrained (A) and trained (B) sheep over time across work status (Pre-work, Work and Recovery) periods at slow, medium and fast walking speeds.. 246. xxvii.

(30) Figure 3C.91. Figure 3C.92. Figure 3C.93. Figure 3C.94. Figure 3C.95. Changes in respiration rate (breaths/min) during work for trained sheep #193 walking at the slow speed.. 248. The relationship between plasma lactate and glucose concentration (mM).. 250. The relationship between lactate concentration in arterial plasma and speed of walking during the first 30 minutes of the Work period.. 252. The relationship between rectal temperature and respiration rate in working sheep.. 254. The relationship between rectal temperature and pulse rate in working sheep.. 254. xxviii.

(31) UST OF PHOTOGRAPHS. Page. Photograph 3A.1. Buffalo walking on the treadmill. A diagrammatic representation showing the components of the treadmill.. 47. Photograph 36.1. An Ongole oow.. 108. Photograph 36.2. A Bali cow.. 109. Photograph 38.3. A Madura oow.. 109. Photograph 38.4. A pair of wor11ing Ongole.. 111. Photograph 3C.1. Experimental sheep in the fibreglass metabolism crates.. 148. Treadmill adapted for two sheep to pull draught loads.. 151. Six sheep walking on the treadmill.. 152. Photograph 3C.2. Photograph 3C.3. xxix.

(32) UST OF ABBREVIATIONS. AA. Amino acid. ADP. Adenosine diphosphate. AMP. Adenosine monophosphate. AmT. Ambient temperature. ARC. Agricultural Research Council. AT. Anaerobic threshold. ATP. Adenosine triphosphate. AV. Arteriovenous. BB. Black bulb. CF. Crude fibre. CNS. Central nervous system. co. Cardiac output. C0 2. Carbon dioxide. er. Creatine. OM. Dry matter. EE. Energy expenditure. FFA. Free fatty acids. G-1-P. Glucose-1-phosphate. H+. Hydrogen ion. H 2P0 4. Diprotonated inorganic phoshate. IMP. lnosine monophosphate. LOH. Lactate dehydrogenase. MAFF. Ministry of Agriculture, Fisheries and Food. Mm. Maintenance energy requirement. N. Nitrogen. NAO•. Nicotinamide-adenine dinucleotide. NAOH. Nicotinamide-adenine dinucleotide, reduced. NaHC0 3. Sodium bicarbonate. NH 3. Ammonia. 02 0 2SAT. Oxygen. OBLA. Onset of blood lactate accumulation. PC02 Per. Partial pressure of carbon dioxide Phosphocreatine. PCV. Packed cell volume. PFK. Phosphofructokinase. Pi. Inorganic phosphate. P02. Partial pressure of oxygen. PR. Pulse rate. PVC. Polyvinyl chloride. RR. Resoiration rate. RT. Rectal temperature. Oxygen saturation. xxx.

(33) SR. Sarcp1asmic reticulum. ST. Skin temperature. T-tubules. Transverse-tubules. TCA TC02. Tricarboxcylic acid Total carbon dioxide. VE. Ventilation. xxxi.

(34) PUBLICATIONS. Kartiarso, Martin D and Telenl E (1987) Nutritional study using a treadmill at James Cook University.. OAP Project Bulletin 3, 11- 15 Kartiarso, Martin. D and Talenl E (1989) The pattem of utlllzation of body reserves by working cattle. and buffalo. DAP Project Bulletin 8, 7 Komarudin-Ma'sum, Teleni E, Martin DG and Aflhandy L (1991) A comparison of the work capacity of Ongole (Sos lndicus), Bati (Sos sondicus) and Madura cows. I. Untrained. DrellflJI Anfmal. Bulletin 2, 7- 15 Komarudln-Ma'sum. Teleni E, Martin DG and Aflhandy L (1991) A comparison of the work capacity of Ongole (Bos incfcus). Bai (Bos sonc:kus} and Madura. cows. II. Trained. Draught Animal. Bulletin 2, 16-22 Martin DG and Telanl E (1988) Blood flow, blood pH and C02 output across the hind-limb muscles of working ruminants. Proc Nut Soc Aust 13, 1OS Martin D and Teleni E (1989) Fatigue in buffaloes on different work loads. DAP Project BuHetin 8,. 2-6 Martin D, Kartiarso and Telanl E (1989) Muscle blood flow in draught animals. DAP Project Bulletin 9, 17-18 Teleni E and Martin DG (1991) Concentrations of some blood metaboites in trained and untrained cows. Proc Nutr Soc Aust 11, 25 Telenl E. Bakrie B, Martin 0, Boniface AN and Murray RM (1988) Physlologicl responses and changes In some blood metabolites In working ruminants. Proc VI v.t>rld Conference on Animal Productlof), Helsin~. p467. Telenl E, Komarudin-Ma'sum, Martin DG and Bakrle B (1991) Concentration of blood metaboites in working Ongole, Bali and Madura. cows. Draught Animal Bulletin 2, 27- 33. xxxii.

(35) ACKNOWLEDGMENT. 1would like to thank my supervisor, Dr E Teleni for his guidance and support in writing this thesis.. I wish to thank the Head of Department, Professor Phil Summers, for the use of the faciiitie.s at the Department of Biomedical and Tropical Veterinary Sciences.. The assistance in some of the field work by Janet Richards, Paula Tomkins, Paula Coombe and the staff at BPT Grati is gratefl.Jlly acl\nowledged with a special thanks to Janet Richards and Syriana for their help in some of the laboratory analysis.. I wish to thank my fellow students, Dwatmadji, Keu Mbwambo and Mohammad Bigdeli for their assistance in sampling during one of the experiments and Mrs Kaye Griffrths for typing this thesis. The help of Jennifer Baillie in typing some of this thesis and help with the final collation is gratefully acl\nowledged.. A special thanks to my family, especially my father and my husband Julian for their moral. support and encouragement throughout the writing of this thesis. Lastly a big hug for "Lucy• my best friend and constant companion whilst writing, who was always by my side (usually because she was asleep on my bed beside my desk).. xxxiii.

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