- Email: [email protected]
Effect of feeding Leucaena leucocephala supplemented rations on thyroid hormones and fasting heat production in Jamunapari goats
Small Ruminant Research ELSEVIER
Small Ruminant Research 19 (1996) 29-33
Effect of feeding Leucaena leucocephala supplemented rations on thyroid hormones and fasting heat production in Jamunapari goats N. Haque”, V.P. Varshney’, M.Y. Khan, M. La1 Division of Animul Nutrition. Indian Veterinary Research Institute, Izahzagar-243 12.2, Etareilly (U.P.), India Accepted 25 January 1995
Abstract Fifteen intact male Jamunapari goats, average BW 22.0 kg, were divided into three groups to investigate the effects of longterm consumption of leucaena on fasting heat production and thyroid hormones. Animals were offered rations containing leucaena leaves and twigs to provide about O%, 40% and 50% CP in L, (control), L2 and L3 groups, respectively, in Phase I ( 165 days). In the second phase (210 days) L, and L3 were similar to the first phase whereas in L2 the proportion of leucaena was reduced to provide 25% CP. Basal roughage was Zea mays (maize), aerial part, in Phase 1 and Aoena saliva (oat) hay in Phase II. Blood samples were collected at loo-day intervals. The mean value of triiodothyronine was 0.82 ng ml-‘, 1.00 ng ml-‘, 1.65 ng ml-‘; 0.84 ng ml-‘, 0.91 ng ml-‘, 0.96 ng ml-’ and 0.76 ng ml-‘, 0.66 ng ml-’ and 0.67 ng ml-’ in L,, L2 and L, groups, respectively, on Day 135 (Stage I), Day 240 (Stage II) and Day 340 (Stage III) of feeding. Triiodothyronine was higher (P< 0.01) for the L3 group in Stage I with no differences at later samplings. Thyroxine level was not affected in this study. Fasting heat production (FHP) was not affected by diet (269 + 9 kJ per kg W”.75 day- ‘). It is concluded that feeding leucaena up to a level to provide 50% CP in the rations to Jamunapari goats had no effect on FHP and serum thyroxine level. Ke.vword.r Leucaena leucucephala; Jamunapari goat; Fasting heat production; Triiodothyronine; Thyroxine
1. Introduction Feeding of Leucaena leucocephala to farm livestock has generally been associated with thyroid gland enlargement. Cattle grazing on mixed grass-leucaena pasture had enlarged thyroid glands with low serum thyroxine level (Jones et al., 1976). Non-significant elevation of thyroxine was observed along with hyperactivity and clinical symptoms of leucaena toxicity in cattle consuming daily 1.3 kg leucaena DM (Falvey, 1976). No significant change in thyroid hormones has * Corresponding author. ’ Present address: Division of Physiology Veterinary India.
Research
and Climatology, Indian Institute, Izatnagigar-243 122, Bareilly (U.P.),
0921.4488/96/$15.00 0 1996 Elsevier Science B.V. All rights reserved .SSDfO921-4488(95)00736-9
also been reported in sheep (Pessoa et al., 1988)) goat (Rai et al., 1991; Senani, 1992), cattle (Wahyuni et al., 1982; Kailas, 1991) consuming leucaena. The objective of this experiment was to study the effect of long-term feeding of leucaena to goat on fasting heat production (FHP) and serum thyroid hormones level.
2. Materials
and methods
Fifteen intact male goats of the Jamunapari breed, 12-15 months old and weighing 22 kg were divided into three groups of five animals in a completely randomised design.
30 Table 1 Ingredients
N. Haque et al. /Small Ruminant Research 19 (1996) 29-33
(g kg-‘)
in the diets (DM basis)
Ingredient
L,
LZ
L,
Maize grain, crushed Groundnut cake, solvent extracted Wheat bran Molasses Maize green, aerial part Leucaena leaves and twigs
207 124 83 172 414
153 45 30 187 390 195
188
The animals on the control treatment (Li ) received a conventional ration consisting of concentrate mixture (crushed maize grain 50%, solvent extracted groundnut cake 30% and wheat bran 20%) and Zea mays (maize) green aerial part, to provide nutrients as per recommendations of the NRC ( 198 1) . The concentrate mixture was supplemented with 2% mineral mixture (contained moisture max. 5%, calcium min. 28%, phosphorus min. 12%, iodine (as KI) 0.02&0.130%, copper 0.077-O. 130%, fluorine max. 0.04%) and 1% common salt. Molasses was added to meet energy requirements, Animals in L2 and L, groups received similar rations except that leucaena leaves and twigs provided about 40% and 50% CP of total ration, respectively, by replacing 50% and 100% each of solvent extracted groundnut cake and wheat bran in the concentrate mixture, respectively. Final composition of the concentrate mixture in the L2 group was crushed maize grain 67%, groundnut cake 20% and wheat bran 13% and in the L3 group it was only crushed maize grain. The composition of ration used is presented in Table 1. The feeding trial continued for 165 days during which metabolism trials on all individual animals were conducted. In Phase II, animals in Li and L3 groups were continued on similar rations for another 210 days but leucaena was reduced in Group L, to provide 25% of CP. Basal roughage was Avena sativa (oat) hay. Mimosine content of leaves and twigs of leucaena was determined as described by Brewbaker and Kaye ( 198 1) . Jugular blood samples were obtained on Days 135,240 and 340 of the study. Serum was analysed for triiodothyronine and thyroxine using Radioimmunoassay (RIA) kits (Bhaba Atomic Research Centre, Bombay-400 094, India). Fasting heat production (FHP) of goats was determined after 72 h of fasting when methane production
203 375 234
was minimal. Air samples from respiration chamber were analysed for oxygen by a dual channel Paramagnetic Oxygen Analyser (Servomex Taylor, model OAT 184)) for carbon dioxide by a modified Sonden apparatus with a 100 ml burette and methane by Infrared gas Analyser (Analytical Development Co. Ltd., Hoddesdon, England, Model 300). Hastings Mass flow meter (Teledyne Hastings-Raydist, VA, USA) was used to record the flow rate and total volume of air coming out of the respiration chamber. Dry and wet bulb temperature of air exiting the chamber was recorded (Decibel Instruments, Chandigarh, India, sl. No. 23183). Atmospheric pressure was recorded electronically (Appleby and Ireland, sl. No. 252730). Heat production was calculated as per Brouwer’s ( 1965) equation: H= 16.175 0,+5.021 COZ-2.167 CH‘,--5.987 N where H is heat production (kJ day- ‘) , 0, is volume of oxygen consumed (1 day-‘), CO, is volume of carbon dioxide produced (1 day- ‘), CH4 is volume of methane produced (1 day-‘), and N is amount of nitrogen excreted in urine (g day-‘). Data were subjected to analysis of variance for a randomised design (Snedecor and Cochran, 1967),
3. Results Chemical composition of feeds and fodder is presented in Table 2. Leucaena leaves and twigs contained 24.24% and 29.37% CP, and 4.19% and 3.45% mimosine on DM basis in Phases I and II, respectively. Molasses has been used in both phases from the same stock. Intake of feed is presented in Table 3. Feeding of rations containing leucaena decreased serum triiodothyronine (Table 4). Decrease of triiodothyronine was significant (P ~0.05) in Group L,
N. Haque et al. /Small RuminantResearch
Table 2 Chemical composition
I9 (19%) 29-33
31
of feeds and fodders (percent on DM basis)
Particulars
Organic matter
Crude protein
Ether extract
Concentrate Mixture I (offered to L, group) Concentrate Mixture II (offered to L, group) Maize grain, crushed
93.08 (91.89) 93.95 (91.61) 95.59 (91.05) 88.54 (88.54) 92.4 1 (91.89) 93.52 (92.69)
21.71 (21.75) 17.00 (18.73) 10.18 (11.84) 8.25 (8.25) 7.69 (11.03) 24.24 (29.37)
2.67 (1.81) 3.62 ( 1.98) 4.44 (1.85) 0.12 (0.12) 1.65 (2.61) 3.27 (3.27)
Molasses Maize green, aerial part Oat hay Leucaena leaves and twigs
Crude fibre 7.21 (7.41) 5 -09 (6.11) 2.33 (2.22)
26.94 (34.14) 17.10 (15.86)
Nitrogen free extracts
Total ash
61.49 (60.92) 68.24 (64.79) 78.64 (75.14) 80.17 (80.17) 56.13 (44.11) 48.91 (44.19)
6.92 (8.11) 6.05 (8.39) 4.4 I (8.95) 11.46 (11.46) 7.59 (8.11) 6.48 (7.31)
Data in parentheses obtained in Phase Il. Table 3 Daily intake of feeds by Jamunapari
goats (percent on DM basis)
Particulars
l-2
L
SE’
l-3
L
Phase1 Maize grain, crushed (g) Groundnut cake, solvent extracted (g) Wheat bran (g ) Molasses (g ) Maize green, aerial part (g) Oat hay (g) Leucaena leaves and twigs (g) DM intake* (g) DM intake’ (g per kg W”.75)
l-3
147.9 88.7 59.1 135.8
172.3 51.4 33.4 180.6
196.4 _
213.1
235.5 105.5 778.7 56.34=
SE’
Phase II 166.1
-
_ 126.0
_ _
171.0 _
308.7
-
IRS.3
225.5 826.2 69.83
128.3 77.0 51.3 110.8
144.6 43.2 28.0 145.5
278.2
645.8 59.69
Lz
717.7 59.91
_ 38.16 1.91
644.7 52.53”
_ _
156.4 _ 278.1 204.9 835.8 61.86’
30.37 1.06
Mean
SE’
Values in row bearing different superscript letters differ (P < 0.05). L,, 0% leucaena CP in the diet; L,, 40% leucaena CP in the diet (25% Phase II); L,, 50% leucaena CP in the diet. ‘Pooled standard error. 2Statistically analyscd. Table 4 Thyroid hormone level (ng ml
‘) in serum
Particulars
L,
L,
L,
Mean
SE’
No. of animals
4
5
4
_
_
Triiodothyronine stage I Stage II Stage Ill Mean SE’
0.82a 0.84 0.76 0.81 0.07
1.oom 0.91AB 0.66A 0.86 0.05
L,
L?
L,
4
5
4
_
21.15 22.98 24.30 22.81 0.61
23.90 22.04 21.39
_
Thyroxine 1.65bn 0.96A 0.67* 1.16 0.08
1.16 0.96 0.70
0.08 0.06 0.07
‘Pooled standard error. “,bValues in row bearing diffeent superscripts differ ( P < 0.01) a.eValues in columns bearing different superscripts differ (P < 0.05 ) Stage I, Day 135 of feeding: L,, 0% leucaena CP in the diet. Stage II, Day 240 of feeding; L,, 40% leucaena CP in the diet (25% in Phase II). Stage III, Day 340 of feeding: L3, 50% leucaena CP in the diet.
24.79 19.68 16.73 20.40 158
25.76 23.46 23.15 24.12 0.95
1.01 0.78 1.42
N. Haque et al. /Small Ruminant Research 19 (19%) 29-33
32 Table 5 Gaseous exchange,
urinary nitrogen, heat production
and respiratory
quotient (R.Q.) in Jamunapari
goats during fasting
Particulars
L,
L,
L3
No. of animals Fasting live weight (kg) Oxygen consumption (1 day-‘) Carbon dioxide production (I dayMethane production (1day - ’ ) Urinary nitrogen (g day - ’ ) Heat production (kJ day - ’ ) (per kg Wo75 day-‘) Respiratory quotient (R.Q.)
4 23.5 148.9 110.4 0.76 2.59 2946
5 26.8 161.3 109.7 0.76 2.17 3145
4 25.7 151.7 102.2 0.63 4.19 2936
(276) 0.68
(269) 0.68
(262) 0.67
‘)
SE”
1.18 2.72 1.85 0.02 0.10 31.52 (9.49) 0.004
aPooled standard error.
after 240 days of feeding and after 340 days in L, group. Thyroxine level was not different among treatments or phase of the study. Average respiration quotient (R.Q.) was 0.68 f 0.006 (Table 5). The FHP in the control (L,) group was highest followed by L2 and La but there was no difference among treatment groups. Mean FHP was 269 f 9 kJ per kg w”.
4. Discussion Mimosine, a non-protein amino acid, present in leucaena leaves and twigs and its metabolite, 3-hydroxy4-( 1H) pyridone (DHP) , has a structural similarity to tyrosine which is a precursor of thyroxine and triiodothyronine. At the molecular level it has been suggested that mimosine may act as tyrosine antagonist and inhibits thyroxine synthesis (Crounseet al., 1962). During Stage I the absorption of mimosine/DHP stimulated triiodothyronine production in the L, group. Subsequently, during Stages II and III the level of triiodothyronine decreased in the L3 group resulting in similar levels in all groups. In Stage I the level of mimosine/DHP available at tissue level might have been insufficient to compete with tyrosine and caused a stimulus rather than depressing effect whereas in subsequent stages its level was such as to cause depression in the synthesis of triiodothyronine. The increase in triiodothyronine concentrations when leucaena was fed during Stage I was similar to the observations of Jones and Hegarty ( 1984) in steers fed leucaena at 20% (DM basis). Beyond this level they observed a decrease in triiodothyronine level.
However, Pessoa et al. (1988) in sheep, Rai et al. (1991) and Senani (1992) in goats did not observe any significant change in serum triiodothyronine level on leucaena feeding. Fasting heat production was not affected by feeding leucaena but FHP was lower than reported by Brody (1945), Fujihara et al. (1973), Roy-Smith (1980), Prieto et al. (1990) and Aguilera et al. (1991). The deviation may be attributed to breed difference since FHP of local nondescript goats was 302 f 7 kJ per kg w7’ day-’ (Khan et al., 1988). Shkolnik ( 1980) has reported that the Bedouin goat has a low metabolism compared with the Mediterranean goat. Vercoe and Frisch ( 1973) also reported variation in FHP of different breeds of cattle. FHP of Shorthorn X Hereford (SH) was higher than those of Brahman X SH and Africander X SH steers. In conclusion it is evident from this study that feeding 50% CP of the ration through leucaena leaves and twigs for a period of 340 days did not have significant effects on FHP and thyroxine level in serum of Jamunapari goats. Acknowledgements Suggestions on statistical analysis of the data from Dr. V.K. Dwivedi, Sr. Scientist and technical assistance from Mr. S. Paul, Laboratory Assistant are duly acknowledged. References Aguilera, J.F., Lara, L. and Prieto, C., 1991. Energy balance studies with growing Granadina goats during fasting and maintenance. Small Rumin. Res., 5: 109-l 15.
N. Hague et al. /Small Ruminant Research I9 (1994) 29-33
Brewbaker, J.L. and Kaye, S., 1981. Mimosine variations in species of the genus leucaena. Leucaena Res. Rep., 2: 66-67. Brody, S., 1945. Bioenergeticsand Growth. Hafner Press, New York, 1023 pp. Brouwer, E., 1965. Report of Sub-Committee on Constants and Factors. Proc. 3rd EAAP Symposium on Energy Metabolism, Publ. No. 11, Academic Press, London, pp. 441443. Crounse, R.C., Maxwell, J.O. and Blank, R., 1962. Inhibition of growth of hair by mimosine. Nature, 194: 694-695. Falvey, L., 1976. The effects of Leucaena leucocephda on cattle in the Northern territory. Aust. Vet. J., 52: 243. Fujihara, T., Tasaki, I. and Furuhashi, T., 1973. Energetic utilization of starch introduced into abomasum of goats. Proc. 6th EAAP Symposium on Energy Metabolism, Publ. No. 14, UniversitatsDokumentationsstelle, Hohenheim, pp. 67-70. Jones, R.J. and Hegarty. M.P., 1984. Effect of different proportions of Leucaena feucocephala in the diet of cattle on growth. feed intake, thyroid function and urinary excretion of 3-Hydroxy4( lH)-pyridone. Aust. J. Agric. Res., 35: 317-325. Jones, R.J., Blunt, C.G. and Holmes, J.H.G., 1976. Enlarged thyroid glands in cattle grazing leucaena pastures. Trop. Grassl., 10: 113116. Kailas, M.M., 1991. Studies on the use of Leucaena leucocephala (Subabul) in the feeding of cattle. Ph.D. Thesis. IVRI, Izatnagar, Bareilly (U.P), pp. 78-82. Khan, M.Y., Murari L., Kishan, J. and Joshi, DC., 1988. Fasting heat production of Indian livestock. Proc. 3rd Symp. on Latest Research Trends in Livestock and Poultry Nutrition (Abstract No. 1). Banaras Hindu University. Varanasi, (U.P.) India, p. 1. NRC, 1981. Nutrient requirements of domestic animals. Goats. Publ. No. 15. National Academy Press, Washington, DC, pp. 1612.
33
Pessoa, J.M., Cardoso, W.M., Velez, C.E.S. and Rodroguez, N.M., 1988. Serum levels of thyroxine and tri-iodothyronine determined by radio-immune assay in sheep fed Leucaena leucocephuh Arq. Bras. Med. Vet. Zootech., 40: 431436. Prieto, C., Aguilera, J.F., Lara, L. and Fonolla, J., 1990. Protein and energy requirements for maintenance of indigenous Granadina goats. Br. J. Nutr., 63: 155-163. Rai, S.N., Kaur, H., Harika, A.S. and Walli, T.K., 1991. Influence of substituting groundnut cake with leucaena leaf meal in concentrate mixture on live weight gain and thyroxine profile in crossbred kids (Abstract No. 175). Proc. First International Animal Nutrition Workers conference for Asia and Pacific, 23-28 September 1991, Bangalore (India), 108 pp. Roy-Smith, F., 1980. The fasting metabolism and relative energy intake of goats compared with sheep. Anim. Prod., 30: 491. Senani, S.. 1992. Studies on the utilization of Leucaena leucocephala (Subabul) as a protein source in the ration of goats. Ph.D. Thesis, IVRI, Izatnagar, (U.P), India, pp. 78-82. Shkolnik. A., 1980. Energy metabolism in hedgehogs: primitive strategies? In: K. Schmidt-Nielsen, L. Bobs and CR. Taylor (Editors), Comparative Physiology: Primitive Mammals. Cambridge University Press, Cambridge, pp. 148-154. Snedecor, G.W. and Cochran, W.G., 1967. Statistical Methods. Sixth Edn. Oxford and IBH Publishing Co., Calcutta, pp. 258-296. Vercoe, J.E. and Frisch, J.E., 1973. Fasting metabolism, liveweight and voluntary feed intake of different breeds of cattle. Proc. 6th Symposium on Energy Metabolism of Farm Animals, Hohenheim. Germany, pp. 131-134. Wahyuni, S., Yulianli,E.S, Komara, W., Yates, N.G., Obst, J.M. and Lowry, J.B., 1982. The performance of Ongole cattle offered either grass, sun dried L. leucocephala or varying proportions of each. Trop. Anim. Prod., 7: 275-282.
Small Ruminant Research 19 (1996) 29-33
Effect of feeding Leucaena leucocephala supplemented rations on thyroid hormones and fasting heat production in Jamunapari goats N. Haque”, V.P. Varshney’, M.Y. Khan, M. La1 Division of Animul Nutrition. Indian Veterinary Research Institute, Izahzagar-243 12.2, Etareilly (U.P.), India Accepted 25 January 1995
Abstract Fifteen intact male Jamunapari goats, average BW 22.0 kg, were divided into three groups to investigate the effects of longterm consumption of leucaena on fasting heat production and thyroid hormones. Animals were offered rations containing leucaena leaves and twigs to provide about O%, 40% and 50% CP in L, (control), L2 and L3 groups, respectively, in Phase I ( 165 days). In the second phase (210 days) L, and L3 were similar to the first phase whereas in L2 the proportion of leucaena was reduced to provide 25% CP. Basal roughage was Zea mays (maize), aerial part, in Phase 1 and Aoena saliva (oat) hay in Phase II. Blood samples were collected at loo-day intervals. The mean value of triiodothyronine was 0.82 ng ml-‘, 1.00 ng ml-‘, 1.65 ng ml-‘; 0.84 ng ml-‘, 0.91 ng ml-‘, 0.96 ng ml-’ and 0.76 ng ml-‘, 0.66 ng ml-’ and 0.67 ng ml-’ in L,, L2 and L, groups, respectively, on Day 135 (Stage I), Day 240 (Stage II) and Day 340 (Stage III) of feeding. Triiodothyronine was higher (P< 0.01) for the L3 group in Stage I with no differences at later samplings. Thyroxine level was not affected in this study. Fasting heat production (FHP) was not affected by diet (269 + 9 kJ per kg W”.75 day- ‘). It is concluded that feeding leucaena up to a level to provide 50% CP in the rations to Jamunapari goats had no effect on FHP and serum thyroxine level. Ke.vword.r Leucaena leucucephala; Jamunapari goat; Fasting heat production; Triiodothyronine; Thyroxine
1. Introduction Feeding of Leucaena leucocephala to farm livestock has generally been associated with thyroid gland enlargement. Cattle grazing on mixed grass-leucaena pasture had enlarged thyroid glands with low serum thyroxine level (Jones et al., 1976). Non-significant elevation of thyroxine was observed along with hyperactivity and clinical symptoms of leucaena toxicity in cattle consuming daily 1.3 kg leucaena DM (Falvey, 1976). No significant change in thyroid hormones has * Corresponding author. ’ Present address: Division of Physiology Veterinary India.
Research
and Climatology, Indian Institute, Izatnagigar-243 122, Bareilly (U.P.),
0921.4488/96/$15.00 0 1996 Elsevier Science B.V. All rights reserved .SSDfO921-4488(95)00736-9
also been reported in sheep (Pessoa et al., 1988)) goat (Rai et al., 1991; Senani, 1992), cattle (Wahyuni et al., 1982; Kailas, 1991) consuming leucaena. The objective of this experiment was to study the effect of long-term feeding of leucaena to goat on fasting heat production (FHP) and serum thyroid hormones level.
2. Materials
and methods
Fifteen intact male goats of the Jamunapari breed, 12-15 months old and weighing 22 kg were divided into three groups of five animals in a completely randomised design.
30 Table 1 Ingredients
N. Haque et al. /Small Ruminant Research 19 (1996) 29-33
(g kg-‘)
in the diets (DM basis)
Ingredient
L,
LZ
L,
Maize grain, crushed Groundnut cake, solvent extracted Wheat bran Molasses Maize green, aerial part Leucaena leaves and twigs
207 124 83 172 414
153 45 30 187 390 195
188
The animals on the control treatment (Li ) received a conventional ration consisting of concentrate mixture (crushed maize grain 50%, solvent extracted groundnut cake 30% and wheat bran 20%) and Zea mays (maize) green aerial part, to provide nutrients as per recommendations of the NRC ( 198 1) . The concentrate mixture was supplemented with 2% mineral mixture (contained moisture max. 5%, calcium min. 28%, phosphorus min. 12%, iodine (as KI) 0.02&0.130%, copper 0.077-O. 130%, fluorine max. 0.04%) and 1% common salt. Molasses was added to meet energy requirements, Animals in L2 and L, groups received similar rations except that leucaena leaves and twigs provided about 40% and 50% CP of total ration, respectively, by replacing 50% and 100% each of solvent extracted groundnut cake and wheat bran in the concentrate mixture, respectively. Final composition of the concentrate mixture in the L2 group was crushed maize grain 67%, groundnut cake 20% and wheat bran 13% and in the L3 group it was only crushed maize grain. The composition of ration used is presented in Table 1. The feeding trial continued for 165 days during which metabolism trials on all individual animals were conducted. In Phase II, animals in Li and L3 groups were continued on similar rations for another 210 days but leucaena was reduced in Group L, to provide 25% of CP. Basal roughage was Avena sativa (oat) hay. Mimosine content of leaves and twigs of leucaena was determined as described by Brewbaker and Kaye ( 198 1) . Jugular blood samples were obtained on Days 135,240 and 340 of the study. Serum was analysed for triiodothyronine and thyroxine using Radioimmunoassay (RIA) kits (Bhaba Atomic Research Centre, Bombay-400 094, India). Fasting heat production (FHP) of goats was determined after 72 h of fasting when methane production
203 375 234
was minimal. Air samples from respiration chamber were analysed for oxygen by a dual channel Paramagnetic Oxygen Analyser (Servomex Taylor, model OAT 184)) for carbon dioxide by a modified Sonden apparatus with a 100 ml burette and methane by Infrared gas Analyser (Analytical Development Co. Ltd., Hoddesdon, England, Model 300). Hastings Mass flow meter (Teledyne Hastings-Raydist, VA, USA) was used to record the flow rate and total volume of air coming out of the respiration chamber. Dry and wet bulb temperature of air exiting the chamber was recorded (Decibel Instruments, Chandigarh, India, sl. No. 23183). Atmospheric pressure was recorded electronically (Appleby and Ireland, sl. No. 252730). Heat production was calculated as per Brouwer’s ( 1965) equation: H= 16.175 0,+5.021 COZ-2.167 CH‘,--5.987 N where H is heat production (kJ day- ‘) , 0, is volume of oxygen consumed (1 day-‘), CO, is volume of carbon dioxide produced (1 day- ‘), CH4 is volume of methane produced (1 day-‘), and N is amount of nitrogen excreted in urine (g day-‘). Data were subjected to analysis of variance for a randomised design (Snedecor and Cochran, 1967),
3. Results Chemical composition of feeds and fodder is presented in Table 2. Leucaena leaves and twigs contained 24.24% and 29.37% CP, and 4.19% and 3.45% mimosine on DM basis in Phases I and II, respectively. Molasses has been used in both phases from the same stock. Intake of feed is presented in Table 3. Feeding of rations containing leucaena decreased serum triiodothyronine (Table 4). Decrease of triiodothyronine was significant (P ~0.05) in Group L,
N. Haque et al. /Small RuminantResearch
Table 2 Chemical composition
I9 (19%) 29-33
31
of feeds and fodders (percent on DM basis)
Particulars
Organic matter
Crude protein
Ether extract
Concentrate Mixture I (offered to L, group) Concentrate Mixture II (offered to L, group) Maize grain, crushed
93.08 (91.89) 93.95 (91.61) 95.59 (91.05) 88.54 (88.54) 92.4 1 (91.89) 93.52 (92.69)
21.71 (21.75) 17.00 (18.73) 10.18 (11.84) 8.25 (8.25) 7.69 (11.03) 24.24 (29.37)
2.67 (1.81) 3.62 ( 1.98) 4.44 (1.85) 0.12 (0.12) 1.65 (2.61) 3.27 (3.27)
Molasses Maize green, aerial part Oat hay Leucaena leaves and twigs
Crude fibre 7.21 (7.41) 5 -09 (6.11) 2.33 (2.22)
26.94 (34.14) 17.10 (15.86)
Nitrogen free extracts
Total ash
61.49 (60.92) 68.24 (64.79) 78.64 (75.14) 80.17 (80.17) 56.13 (44.11) 48.91 (44.19)
6.92 (8.11) 6.05 (8.39) 4.4 I (8.95) 11.46 (11.46) 7.59 (8.11) 6.48 (7.31)
Data in parentheses obtained in Phase Il. Table 3 Daily intake of feeds by Jamunapari
goats (percent on DM basis)
Particulars
l-2
L
SE’
l-3
L
Phase1 Maize grain, crushed (g) Groundnut cake, solvent extracted (g) Wheat bran (g ) Molasses (g ) Maize green, aerial part (g) Oat hay (g) Leucaena leaves and twigs (g) DM intake* (g) DM intake’ (g per kg W”.75)
l-3
147.9 88.7 59.1 135.8
172.3 51.4 33.4 180.6
196.4 _
213.1
235.5 105.5 778.7 56.34=
SE’
Phase II 166.1
-
_ 126.0
_ _
171.0 _
308.7
-
IRS.3
225.5 826.2 69.83
128.3 77.0 51.3 110.8
144.6 43.2 28.0 145.5
278.2
645.8 59.69
Lz
717.7 59.91
_ 38.16 1.91
644.7 52.53”
_ _
156.4 _ 278.1 204.9 835.8 61.86’
30.37 1.06
Mean
SE’
Values in row bearing different superscript letters differ (P < 0.05). L,, 0% leucaena CP in the diet; L,, 40% leucaena CP in the diet (25% Phase II); L,, 50% leucaena CP in the diet. ‘Pooled standard error. 2Statistically analyscd. Table 4 Thyroid hormone level (ng ml
‘) in serum
Particulars
L,
L,
L,
Mean
SE’
No. of animals
4
5
4
_
_
Triiodothyronine stage I Stage II Stage Ill Mean SE’
0.82a 0.84 0.76 0.81 0.07
1.oom 0.91AB 0.66A 0.86 0.05
L,
L?
L,
4
5
4
_
21.15 22.98 24.30 22.81 0.61
23.90 22.04 21.39
_
Thyroxine 1.65bn 0.96A 0.67* 1.16 0.08
1.16 0.96 0.70
0.08 0.06 0.07
‘Pooled standard error. “,bValues in row bearing diffeent superscripts differ ( P < 0.01) a.eValues in columns bearing different superscripts differ (P < 0.05 ) Stage I, Day 135 of feeding: L,, 0% leucaena CP in the diet. Stage II, Day 240 of feeding; L,, 40% leucaena CP in the diet (25% in Phase II). Stage III, Day 340 of feeding: L3, 50% leucaena CP in the diet.
24.79 19.68 16.73 20.40 158
25.76 23.46 23.15 24.12 0.95
1.01 0.78 1.42
N. Haque et al. /Small Ruminant Research 19 (19%) 29-33
32 Table 5 Gaseous exchange,
urinary nitrogen, heat production
and respiratory
quotient (R.Q.) in Jamunapari
goats during fasting
Particulars
L,
L,
L3
No. of animals Fasting live weight (kg) Oxygen consumption (1 day-‘) Carbon dioxide production (I dayMethane production (1day - ’ ) Urinary nitrogen (g day - ’ ) Heat production (kJ day - ’ ) (per kg Wo75 day-‘) Respiratory quotient (R.Q.)
4 23.5 148.9 110.4 0.76 2.59 2946
5 26.8 161.3 109.7 0.76 2.17 3145
4 25.7 151.7 102.2 0.63 4.19 2936
(276) 0.68
(269) 0.68
(262) 0.67
‘)
SE”
1.18 2.72 1.85 0.02 0.10 31.52 (9.49) 0.004
aPooled standard error.
after 240 days of feeding and after 340 days in L, group. Thyroxine level was not different among treatments or phase of the study. Average respiration quotient (R.Q.) was 0.68 f 0.006 (Table 5). The FHP in the control (L,) group was highest followed by L2 and La but there was no difference among treatment groups. Mean FHP was 269 f 9 kJ per kg w”.
4. Discussion Mimosine, a non-protein amino acid, present in leucaena leaves and twigs and its metabolite, 3-hydroxy4-( 1H) pyridone (DHP) , has a structural similarity to tyrosine which is a precursor of thyroxine and triiodothyronine. At the molecular level it has been suggested that mimosine may act as tyrosine antagonist and inhibits thyroxine synthesis (Crounseet al., 1962). During Stage I the absorption of mimosine/DHP stimulated triiodothyronine production in the L, group. Subsequently, during Stages II and III the level of triiodothyronine decreased in the L3 group resulting in similar levels in all groups. In Stage I the level of mimosine/DHP available at tissue level might have been insufficient to compete with tyrosine and caused a stimulus rather than depressing effect whereas in subsequent stages its level was such as to cause depression in the synthesis of triiodothyronine. The increase in triiodothyronine concentrations when leucaena was fed during Stage I was similar to the observations of Jones and Hegarty ( 1984) in steers fed leucaena at 20% (DM basis). Beyond this level they observed a decrease in triiodothyronine level.
However, Pessoa et al. (1988) in sheep, Rai et al. (1991) and Senani (1992) in goats did not observe any significant change in serum triiodothyronine level on leucaena feeding. Fasting heat production was not affected by feeding leucaena but FHP was lower than reported by Brody (1945), Fujihara et al. (1973), Roy-Smith (1980), Prieto et al. (1990) and Aguilera et al. (1991). The deviation may be attributed to breed difference since FHP of local nondescript goats was 302 f 7 kJ per kg w7’ day-’ (Khan et al., 1988). Shkolnik ( 1980) has reported that the Bedouin goat has a low metabolism compared with the Mediterranean goat. Vercoe and Frisch ( 1973) also reported variation in FHP of different breeds of cattle. FHP of Shorthorn X Hereford (SH) was higher than those of Brahman X SH and Africander X SH steers. In conclusion it is evident from this study that feeding 50% CP of the ration through leucaena leaves and twigs for a period of 340 days did not have significant effects on FHP and thyroxine level in serum of Jamunapari goats. Acknowledgements Suggestions on statistical analysis of the data from Dr. V.K. Dwivedi, Sr. Scientist and technical assistance from Mr. S. Paul, Laboratory Assistant are duly acknowledged. References Aguilera, J.F., Lara, L. and Prieto, C., 1991. Energy balance studies with growing Granadina goats during fasting and maintenance. Small Rumin. Res., 5: 109-l 15.
N. Hague et al. /Small Ruminant Research I9 (1994) 29-33
Brewbaker, J.L. and Kaye, S., 1981. Mimosine variations in species of the genus leucaena. Leucaena Res. Rep., 2: 66-67. Brody, S., 1945. Bioenergeticsand Growth. Hafner Press, New York, 1023 pp. Brouwer, E., 1965. Report of Sub-Committee on Constants and Factors. Proc. 3rd EAAP Symposium on Energy Metabolism, Publ. No. 11, Academic Press, London, pp. 441443. Crounse, R.C., Maxwell, J.O. and Blank, R., 1962. Inhibition of growth of hair by mimosine. Nature, 194: 694-695. Falvey, L., 1976. The effects of Leucaena leucocephda on cattle in the Northern territory. Aust. Vet. J., 52: 243. Fujihara, T., Tasaki, I. and Furuhashi, T., 1973. Energetic utilization of starch introduced into abomasum of goats. Proc. 6th EAAP Symposium on Energy Metabolism, Publ. No. 14, UniversitatsDokumentationsstelle, Hohenheim, pp. 67-70. Jones, R.J. and Hegarty. M.P., 1984. Effect of different proportions of Leucaena feucocephala in the diet of cattle on growth. feed intake, thyroid function and urinary excretion of 3-Hydroxy4( lH)-pyridone. Aust. J. Agric. Res., 35: 317-325. Jones, R.J., Blunt, C.G. and Holmes, J.H.G., 1976. Enlarged thyroid glands in cattle grazing leucaena pastures. Trop. Grassl., 10: 113116. Kailas, M.M., 1991. Studies on the use of Leucaena leucocephala (Subabul) in the feeding of cattle. Ph.D. Thesis. IVRI, Izatnagar, Bareilly (U.P), pp. 78-82. Khan, M.Y., Murari L., Kishan, J. and Joshi, DC., 1988. Fasting heat production of Indian livestock. Proc. 3rd Symp. on Latest Research Trends in Livestock and Poultry Nutrition (Abstract No. 1). Banaras Hindu University. Varanasi, (U.P.) India, p. 1. NRC, 1981. Nutrient requirements of domestic animals. Goats. Publ. No. 15. National Academy Press, Washington, DC, pp. 1612.
33
Pessoa, J.M., Cardoso, W.M., Velez, C.E.S. and Rodroguez, N.M., 1988. Serum levels of thyroxine and tri-iodothyronine determined by radio-immune assay in sheep fed Leucaena leucocephuh Arq. Bras. Med. Vet. Zootech., 40: 431436. Prieto, C., Aguilera, J.F., Lara, L. and Fonolla, J., 1990. Protein and energy requirements for maintenance of indigenous Granadina goats. Br. J. Nutr., 63: 155-163. Rai, S.N., Kaur, H., Harika, A.S. and Walli, T.K., 1991. Influence of substituting groundnut cake with leucaena leaf meal in concentrate mixture on live weight gain and thyroxine profile in crossbred kids (Abstract No. 175). Proc. First International Animal Nutrition Workers conference for Asia and Pacific, 23-28 September 1991, Bangalore (India), 108 pp. Roy-Smith, F., 1980. The fasting metabolism and relative energy intake of goats compared with sheep. Anim. Prod., 30: 491. Senani, S.. 1992. Studies on the utilization of Leucaena leucocephala (Subabul) as a protein source in the ration of goats. Ph.D. Thesis, IVRI, Izatnagar, (U.P), India, pp. 78-82. Shkolnik. A., 1980. Energy metabolism in hedgehogs: primitive strategies? In: K. Schmidt-Nielsen, L. Bobs and CR. Taylor (Editors), Comparative Physiology: Primitive Mammals. Cambridge University Press, Cambridge, pp. 148-154. Snedecor, G.W. and Cochran, W.G., 1967. Statistical Methods. Sixth Edn. Oxford and IBH Publishing Co., Calcutta, pp. 258-296. Vercoe, J.E. and Frisch, J.E., 1973. Fasting metabolism, liveweight and voluntary feed intake of different breeds of cattle. Proc. 6th Symposium on Energy Metabolism of Farm Animals, Hohenheim. Germany, pp. 131-134. Wahyuni, S., Yulianli,E.S, Komara, W., Yates, N.G., Obst, J.M. and Lowry, J.B., 1982. The performance of Ongole cattle offered either grass, sun dried L. leucocephala or varying proportions of each. Trop. Anim. Prod., 7: 275-282.