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L-carnitine and its functional effects in poultry nutrition

Published online by Cambridge University Press:  20 June 2011

SH. GOLZAR ADABI ,
R.G. COOPER ,
N. CEYLAN  and
M. CORDUK
SH. GOLZAR ADABI*
Affiliation:
Faculty of Agriculture, Department of Animal Science, Ankara University, Diskapi, Ankara, 06110-Turkey
R.G. COOPER
Affiliation:
Physiologist, (formerly University of Zimbabwe), 22 Kimble Grove, Pype Hayes, Birmingham B24 0RW, UK
N. CEYLAN
Affiliation:
Faculty of Agriculture, Department of Animal Science, Ankara University, Diskapi, Ankara, 06110-Turkey
M. CORDUK
Affiliation:
Faculty of Agriculture, Department of Animal Science, Ankara University, Diskapi, Ankara, 06110-Turkey
*
Corresponding author: adabishahram@gmail.com
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Abstract

L-carnitine has indispensable functions in intermediary metabolism and is received by endogenous synthesis and from exogenous sources. It plays an obligate role in fatty acid metabolism by directing fatty acids into the mitochondrial oxidative pathway through the action of specialised acyltransferases. In poultry production, L-carnitine has a multi functional purpose, which includes: growth promotion, strengthening the immune system, antioxidant effects and improving semen quality. The concentration of L-carnitine in animals varies widely across species, tissue type, and nutritional status of the animal. It has been suggested that the L-carnitine requirement may be increased under certain circumstances such as via higher performance, various stress conditions and where the diet is deficient in animal protein sources. The review of L-carnitine functions uniquely includes the main aspects of this consequential feed supplementary inclusion in poultry nutrition.

Keywords

chickenL-carnitineperformanceimmune responseantioxidationfertility

Information

Type
Review Article
Information
World's Poultry Science Journal , Volume 67 , Issue 2 , June 2011 , pp. 277 - 296
Copyright
Copyright © World's Poultry Science Association 2011

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References

AGARWAL, A. and SAID, T.M. (2004) L-Carnitines and male infertility. Reproductive Biomedicine Online 8: 376-384.CrossRefGoogle Scholar
AGARWAL, A., GUPTA, S. and SHARMA, R.K. (2005) Role of oxidative stress in female reproduction. Reproductive Biology and Endocrinology 3: 28.Google Scholar
AITKEN, R.J., HARKISS, D. and BUCKINGHAM, D.W. (1993) Analysis of lipid peroxidation mechanisms in human spermatozoa. Molecular Reproduction and Development 35: 302-315.Google Scholar
AITKEN, R.J., DE IULIIS, G.N., FINNIE, J.M., HEDGES, A. and MCLACHLAN, R.I. (2010) Analysis of the relationships between oxidative stress, DNA damage and sperm vitality in a patient population. Development of Diagnostic Criteria 00: 1-12.Google Scholar
AMENDOLA, R., CORDELLI, E., MAURO, F. and SPANO, M. (1991) Effects of L-acetylcarnitine (LAC) on the post-injury recovery of mouse spermatogenesis monitored by flow cytometry 2. Recovery after hyperthermic treatment. Andrologia 23: 135-140.Google Scholar
ALVAREZ, J.G. and STOREY, B.T. (1989) Role of glutathione peroxidase in protecting mammalian spermatozoa from loss of motility caused by spontaneous lipid peroxidation. Gamete Research 23: 77-90.CrossRefGoogle ScholarPubMed
ARDUINI, A. (1992) Carnitine and its acyl esters as secondary antioxidants? American Heart Journal 123: 1726-1727.CrossRefGoogle ScholarPubMed
ARSLAN, C., CITIL, M. and SAATCI, M. (2003) Effects of L-carnitine administration on growth performance, carcass traits, blood serum parameters and abdominal fatty acid composition of ducks. Archives of Animal Nutrition 57(5): 381-388.CrossRefGoogle Scholar
ARSLAN, C., CITIL, M. and SAATCI, M. (2004a) Effects of L-carnitine administration on growth performance, carcass traits and some serum components of Japanese quail (Coturnix cot. japonica). A rchiv für Geflügelkunde 68: 111-114.Google Scholar
ARSLAN, C., CITIL, M. and SAATCI, M. (2004b) Effects of L-carnitine administration on growth performance, carcass traits, serum lipids and abdominal fatty acid compositions of geese. Revue de Médecine Vétérinaire 155: 315-320.Google Scholar
ATROSHI, F., RIZZO, A., BIESE, L., VEIJALAINEN, P., SALONIEMI, H., SANKARI, S. and ANDERSSON, K. (1999) Fumonisin B1-induced DNA damage in rat liver and spleen: Effect of pre-treatment with coenzyme Q10, L-carnitine, α-tocopherol and selenium. Pharmacological Research 40: 459-467.Google Scholar
BALERCIA, G., REGOLI, F., ARMENI, T., KOVERECH, A., MANTERO, F. and BOSCARO, M. (2005) Placebo-controlled double-blind randomized trial on the use of l-carnitine, l-acetylcarnitine, or combined L-carnitine and l-acetylcarnitine in men with idiopathic asthenozoospermia. Fertility and Sterility 84: 662-671.CrossRefGoogle ScholarPubMed
BARBATO, G.F. (1999) Genetic relationships between selection for growth and reproductive effectiveness. Poultry Science 78: 444-452.CrossRefGoogle ScholarPubMed
BARKER, D.L. and SELL, J.L. (1994) Dietary carnitine did not influence performance and carcass composition of broiler chickens and young turkey fed low- or high- fat diets. Poultry Science 73: 281-287.CrossRefGoogle ScholarPubMed
BAYRAKTAR, N., PASAOGLU, H., PASAOGLU, A., KAYMAZ, M., BIBEROGLU, G. and KULAKSIZOGLU, S. (2008) The relationship between carnitine levels and lipid peroxidation in glial brain tumors. Turkish Journal of Medical Science 38: 293-299.Google Scholar
BECCAVIN, C., CHEVALIER, B., COGBURN, L.A., SIMON, J. and DUCLOS, M.J. (2001) Insulin-like growth factors and body growth in chickens divergently selected for high or low growth rate. Journal of Endocrinology 168: 297-306.Google Scholar
BENEVENGA, N.J., STEINMAN-GOLDWORTHY, J.K., CRENSHAW, V. and ODLE, J. (1989) Utilization of medium-chain triglycerides by neonatal piglets: I. Effects on milk consumption and body fuel utilization. Journal of Animal Science 67: 3331-3339.CrossRefGoogle ScholarPubMed
BIEBER, L.L. (1988) Carnitine. Annual Review of Biochemistry 57: 261-283.CrossRefGoogle ScholarPubMed
BORUM, P.R. (1983) L-Carnitine. Annual Review of Nutrition 3: 233-259.Google Scholar
BORUM, P.R. (1987) Role of carnitine in lipid metabolism, in: HORISBERGER, M. & BRACCO, U. (Eds) Lipids in modern nutrition, pp. 51-58 (New York, Nestle Vevey/Raven Press).Google Scholar
BOTTJE, W., TANG, Z.X., IQBAL, M., CAWTHON, D., OKIMOTO, R., WING, T. and COOPER, M. (2002) Association of mitochondrial function with feed efficiency within a single genetic line of male broilers. Poultry Science 81: 546-555.Google Scholar
BREMER, J. (1962) L-Carnitine in intermediary metabolism. The metabolism of fatty acid esters of carntine by mitochondria. Journal of Biological Chemistry 237(12): 3628-3632.CrossRefGoogle Scholar
BREMER, J. (1983) Carnitine-metabolism, functions. Physiological Reviews 63: 1420-1480.CrossRefGoogle ScholarPubMed
BROWN, G.C. (1999) Nitric oxide and mitochondrial respiration, Biochimica and Biophysica Acta 1411: 351-369.CrossRefGoogle ScholarPubMed
BUYSE, J., JANSSENS, G.P.J. and DECUYPERE, E. (2001) The effects of dietary L-carnitine supplementation on the performance, organ weights and circulating hormone and metabolite concentrations of broiler chickens reared under abnormal or low temperature schedule. British Poultry Science 42: 230-241.CrossRefGoogle ScholarPubMed
BUYSE, J., QUIRINE, Q., NIEWOLD, T.A., KLASING, K.C., JANSSENS, G.P.J., BAUMGARTNER, M. and GODDEERIS, B.M. (2007) Dietary L-carnitine supplementation enhances the lipopolysaccharide-induced acute phase protein response in broiler chickens. Veterinary Immunology and Immunopathology 118: 154-159.CrossRefGoogle ScholarPubMed
CARTWRIGHT, A.L. (1986) Effect of carnitine and dietary energy concentration on body weight and body lipid of growing broilers. Poultry Science 65: 21-29.Google Scholar
CASILLAS, E.R. and NEWBURGH, R.W. (1969) L-carnitine and derivatives in embryonic chick tissue. Biochimica et Biophysica Acta. 184: 566-577.CrossRefGoogle ScholarPubMed
CAVIGLIA, D., SCARABELLI, L. and PALMERO, S. (2004) Effects of carnitines on rat sertoli cell protein metabolism. Hormone and Metabolic Research 36: 221-225.Google Scholar
CELIK, L. and OZTURKCAN, O. (2003) Effects of dietary supplemental L-carnitine and ascorbic acid on performance, carcass composition and plasma L-carnitine concentration of broiler chicks reared under different temperature. Archive of Animal Nutrition 57: 27-38.CrossRefGoogle ScholarPubMed
CEVIK, A.F. and CEYLAN, N. (2005) Etlik piliç yemlerine L-karnitin ilavesinin performans ve karkas özelliklerine etkileri (Effects of dietary L-carnitine supplementation on performance and carcass traits of broiler chickens). 3rd Ulusal Hayvan Besleme Kongresi, Adana, pp. 391-396.Google Scholar
CHIODI, P., CIANI, B., KENTROTI, S., MACCARI, F., VERNADAKIS, A., ANGELUCCI, L. and RAMACCI, M.T. (1994) L-carnitine and derivatives in the central nervous system of chick embryo. International Journal of Biochemistry 26: 711-720.CrossRefGoogle ScholarPubMed
CITIL, M., GUNES, V., ATAKISI, O., OZCAN, A., TUZCU, M. and DOGAN, A. (2005) Protective effect of L-carnitine against oxidative damage caused by experimental chronic aflatoxicosis in quail (Coturnix coturnix). Acta Veterinaria Hungarica 53: 319-324.Google Scholar
COOPER, T.G., GUDERMANN, T.W. and YEUNG, C.H. (1986a) Characteristics of the transport of carnitine into the cauda epididymis of the rat as ascertained by luminal perfusion in vitro. International Journal of Andrology 9: 348-358.Google Scholar
COOPER, T.G., YEUNG, C.H. and WEINBAUER, G.F. (1986b) Transport of carnitine by the epididymis of the cynomolgus macaque (Macaca fascicularis). Journal of Reproduction and Fertility 77: 297-301.Google Scholar
CORDUK, M., CEYLAN, N. and ILDIZ, F. (2007) Effects of dietary energy density and L-carnitine supplementation on growth performance, carcass traits and blood parameters of broiler chickens. South African Journal of Animal Science 37: 65-73.Google Scholar
CORDUK, M., SARICA, S., CALIKOGLU, E. and KIRALAN, M. (2008) Effects of L-carnitine supplementation to diets with different fat sources and energy levels on fatty acid composition of egg yolk of laying hens. Journal of the Science of Food and Agriculture 88: 2244-2252.CrossRefGoogle Scholar
CORRIER, D.E., MOLLENHAUER, H.H., CLARK, D.E., HARE, M.F. and ELISSALDE, M.H. (1985) Testicular degeneration and necrosis induced by dietary cobalt. Veterinary Pathology 22: 610-616.Google Scholar
CYR, D.M., EGAN, S.G., BRINI, C.M. and TREMBLAY, G.C. (1991) On the mechanism of inhibition of gluconeogenesis and ureagenesis by sodium benzoate. Biochemical Pharmacology 42: 645-654.CrossRefGoogle ScholarPubMed
DASKIRAN, M. and TEETER, R.G. (2001) Effect of dietary Lcarnitine (carniking) supplementation on overall performance and carcass characteristics of seven- week-old broiler chickens. Publ. Oklahoma Agric. Exp. Sta, Oklahoma State University, Stillwater, Ok, pp. 986.Google Scholar
DASKIRAN, M. ONOLl, A.G., , CENGIZ, O., TATLI, O. and SARI, M. (2009) Effects of dietary methionine levels and L-carnitine supplementation on performance and egg quality parameters of layers. Journal of Animal and Feed Sciences 18: 650-661.CrossRefGoogle Scholar
DAYANANDAN, A., KUMAR, P. and PANNEER, S.C. (2001) Protective role of L-carnitine on liver and heart lipid peroxidation in atherosclerotic rats. The Journal of Nutritional Biochemistry 12: 254-257.CrossRefGoogle ScholarPubMed
DENG, K., WONG, C.W. and NOLAN, J.V. (2006) Long-term effects of early-life dietary L-carnitine on lymphoid organs and immune responses in Leghorn-type chickens. Journal of Animal Physiology and Animal Nutrition 90: 81-86.Google Scholar
DE SIMONE, C., FERRARI, M., LOZZI, A., MELI, D., RICA, D. and SORICE, F. (1982) Vitamins and immunity: II. Influence of L-carnitine on the immune system. Acta Vitaminology and Enzymology 4: 135-140.Google Scholar
DE SIMONE, C., FAMULARO, G., TZANTZOGLOU, S., TRINCHIERI, V., MORETTI, S. and SORICE, F. (1994) Carnitine depletion in peripheral blood mononuclear cells from patients with AIDS: effect of oral L-carnitine. AIDS 8: 655-660.CrossRefGoogle ScholarPubMed
DI GIACOMO, C., LATTERI, F., FICHERA, C., SORRENTI, V., CAMPISI, A., CASTORINA, C., RUSSO, A., PINTURO, R. and VANELLA, A. (1993) Effect of acetyl-L- carnitine on lipid peroxidation and xanthine oxidase activity in rat skeletal muscle. Neurochemical Research 18: 1157-1162.Google Scholar
ENOMOTO, A., WEMPE, M.F., TSUCHIDA, H., SHIN, H.J., CHA, S.H., ANZAI, N., GOTO, A., SAKAMOTO, A., NIWA, T., KANAI, Y., ANDERS, M.W. and ENDOU, H. (2002) Molecular identification of a novel carnitine transporter specific to human testis. The Journal of Biological Chemistry 277: 36262-36271.Google Scholar
FAMULARO, G. and DE SIMONE, C. (1995) A new era for carnitine? Immunology today 16: 211-213.Google Scholar
FELLENBERG, M.A. and SPEISKY, H. (2006) Antioxidants: their effects on broiler oxidative stress and its meat oxidative stability. World's Poultry Science Journal 62: 53-70.CrossRefGoogle Scholar
FISCHER, M., VARADY, J., HIRCHE, F., KLUGE, H. and EDER, K. (2009) Supplementation of L-carnitine in pigs: Absorption of carnitine and effect on plasma and tissue carnitine concentrations. Archives of Animal Nutrition 63: 1-15.CrossRefGoogle ScholarPubMed
FUJIHARA, N. and KOGA, O. (1984) Prevention of the production of lipid peroxide in rooster spermatozoa. Animal Reproduction Science 7: 385-390.CrossRefGoogle Scholar
GARCIA-MIRANDA, P., DURAN, J.M., PERAL, M.J. and ILUNDAIN, A.A. (2005) Developmental maturation and segmental distribution of rat small intestinal L-carnitine uptake. Journal of Membrane Biology 206: 9-16.Google Scholar
GENG, A., LI, B. and GUO, Y. (2007) Effects of dietary L-carnitine and coenzyme Q10 at different supplemental ages on growth performance and some immune response in ascites-susceptible broilers. Archives of Animal Nutrition 61: 50-60.CrossRefGoogle ScholarPubMed
GHODS-ALAVI, B., SAMIE, H. and JAHANIAN, R. (2010) Effects of supplementary dietary L-carnitine on performance and egg quality of laying hens fed diets different in fat level. Italian Journal of Animal Science 9: e3CrossRefGoogle Scholar
GOLZAR ADABI, SH., MOGHADDAM, G., TAGHIZADEH, A., NEMATOLLAHI, A. and FARAHVASH, T. (2006a) Effect of two dietary levels of L-carnitine and vegetable fat powder on broiler breeder fertility, hatchability, egg yolk and serum cholesterol and triglyceride. International Journal of Poultry Science 5: 970-974.Google Scholar
GOLZAR ADABI, SH., RAHIMI, SH. and MOGHADDAM, G. (2006b) . Effect of dietary L-Carnitine supplementation on performance and immune system of broiler chickens. Scientific Journal of Faculty of Agricultural Science of Tabriz University 16: 218-227 (with English abstract).Google Scholar
GOLZAR ADABI, SH., RAHIMI, SH., KAMALI, M.A. and KARIMI TORSHIZI, M.A. (2007) The effects of two dietary levels of L-carnitine and vegetable fat powder on quality of cockerels sperm, and fertility and hatchability in broiler breeders. Journal of Veterinary Research 62: 107-114 (with English abstract).Google Scholar
GOLZAR ADABI, SH., HAJIBABAEI, A., FARAHVASH, T., LOTFOLLAHIAN, H. and MOSLEMI POUR, F. (2008) L-carnitine effect on quantity and quality of African black neck ostrich sperm. Journal of Animal and Veterinary Advances 3: 369-374.Google Scholar
GUERIN, P., EL MOUATASSIM, S. and MENEZO, Y. (2001) Oxidative stress and protection against reactive oxygen species in the pre-implantation embryo and its surroundings. Human Reproduction Update 7: 175-189.CrossRefGoogle ScholarPubMed
GULCIN, I. (2006) Antioxidant and antiradical activities of L-carnitine. Life Sciences 78: 803-811.Google Scholar
HABER, F. and WEISS, J. (1934) The catalytic decomposition of hydrogen peroxide by iron salts. Proceedings of the Royal Society of London. Series A, Mathematical and Physical Sciences 147: 332-351.Google Scholar
HAJIBABAEI, A., SHIVAZAD, M., GOLZAR ADABI, SH., LAVAF, A. And EILA and N., (2008) L-carnitine improves egg production in black neck ostriches. Australian Journal of Experimental Agriculture 48: 1341-1344.Google Scholar
HARPAZ, S., BECKER, K. and BLUM, R. (1999) The effect of dietary L-carnitine supplementation on cold tolerance and growth of the ornamental cichlid fish pelvicachromis pulcher-preliminary results. Journal of Thermal Biology 24: 57-62.Google Scholar
HARPAZ, S. (2005) L-carnitine and its attributed functions in fish culture and nutrition. Aquaculture 249: 3-21.CrossRefGoogle Scholar
HATANAKA, T., HARAMURA, M., FEI, Y.J., MIYAUCHI, S., BRIDGES, C.C., GANAPATHY, P.S., SMITH, S.B., GANAPATHY, V. and GANAPATHY, M.E. (2004) Transport of amino acid-based prodrugs by the Na+- and Cl--coupled amino acid transporter ATB0,+ and expression of the transporter in tissues amenable for drug delivery. Journal of Pharmacology and Experimental Therapeutics 308: 1138-1147.CrossRefGoogle ScholarPubMed
HOPPEL, C. (2003) The Role of carnitine in normal and altered fatty acid metabolism. American Journal of Kidney Diseases 41: S4-S12.Google Scholar
JANSSENS, G.P.J. and DE WILDE, R.O. (1994) Carnitine in racing pigeons. Proceedings of the 4th ESVIM Annual Congress, Brussels, pp. 18-19.Google Scholar
JANSSENS, G.P.J., DE RYCKE, H., HESTA, M. and De WILDE, R.O. (1999) Analysis of carnitine, betaine, gamma-butyrobetaine, and separate short-chain acylcarnitines in pigeon plasma, crop milk and tissues by HPLC coupled with UV-detection. Biotechnology Techniques 13: 231-234.Google Scholar
JANSSENS, G.P.J., HESTA, M., DEBAL, V., DEBRAEKELEER, J. and DE WILDE, R.O. (2000a) L-Carnitine supplementation in breeding pigeons: impact on zootechnical performance and carnitine metabolism. Reproduction Nutrition Development 40: 535-548.Google Scholar
JANSSENS, G.P.J., MAST, J., GODDEERIS, B.M., COX, E., HESTA, M. and DE WILDE, R.O. (2000b) Enhanced specific antibody response to bovine serum albumin in pigeons due to L-carnitine supplementation. British Poultry Science 41: 448-453.CrossRefGoogle ScholarPubMed
JEULIN, C. and Lewin, L.M. (1996) Role of free L-carnitine and acetyl-L-carnitine in post-gonadal maturation of mammalian spermatozoa. Human Reproduction Update 2: 87-102.Google Scholar
KALAISELVI, T. and PANNEERSELVAM, C. (1998) Effect of L-carnitine on the status of lipid peroxidation and antioxidants in aging rats. The Journal of Nutritional Biochemistry 9: 575-581.CrossRefGoogle Scholar
KATO, Y., SUGIURA, M., SUGIURA, T., WAKAYAMA, T., KUBO, Y., KOBAYASHI, D., SAI, Y., TAMAI, I., ISEKI, S. and TSUJI, A. (2006) Organic cation/carnitine transporter OCTN2 (Slc22a5) is responsible for carnitine transport across apical membranes of small intestinal epithelial cells in mouse. Molecular Pharmacology 70: 829-837.Google Scholar
KELSO, K.A., CEROLINI, S., SPEAKE, B.K., CAVALCHINI, L.G. and NOBLE, R.G. (1997) Effects of dietary supplementation with α-linolenic acid on the phospholipid fatty acid composition and quality of spermatozoa in cockerel from 24 to 72 weeks of age. Journal of Reproduction and Fertility 110: 53-59.Google Scholar
KERALAPURATH, M.M., KERIS, R.W., CORZO, A., BENNETT, L.W., PULIKANTI, R. and PEEBLES, E.D. (2010a) Effects of in ovo injection of L-carnitine on subsequent broiler chick tissue nutrient profiles. Poultry Science 89: 335-341.Google Scholar
KERALAPURATH, M.M., CORZO, A., PULIKANTI, R., ZHAI, W. and PEEBLES, E.D. (2010b) Effects of in ovo injection of L-carnitine on hatchability and subsequent broiler performance and slaughter yield. Poultry Science 89: 1497-1501.CrossRefGoogle ScholarPubMed
KHAIRALLAH, E.A. and WOLF, G. (1965) Growth-promoting and lipotropic effect of carnitine in rats fed diets limited in protein and methionine. Journal of Nutrition 87: 469-476.CrossRefGoogle ScholarPubMed
KIDD, M.T., MCDANIEL, C.D., PEEBLES, E.D., BARBER, S.J., CORZO, A., BRANTON, A.L. and WOODWORTH, J.C. (2005) Breeder hen dietary L-carnitine affects progeny carcase traits. British Poultry Science 46: 97-103.Google Scholar
KIDD M.T., , GILBERT, J., CORZO, A., PAGE, C., VIRDEN, W.S. and WOODWORTH, J.C. (2009) . Dietary L-carnitine influences broiler thigh yield. Asian-Australasian Journal of Animal Sciences 22: 681-685.Google Scholar
KITA, K., KATO, S., YAMAN, M.A., OKUMURA, J. and YOKOTA, H. (2002) Dietary L-carnitine increases plasma insulin-like growth factor-I concentration in chicks fed a diet with adequate dietary protein level. British Poultry Science 43: 117-121.CrossRefGoogle ScholarPubMed
KITA, K., NAKAJIMA, S. and NAKAGAWA, J. (2005) Dietary L-carnitine supplementation improves albumen quality of laying hens. The Journal of Poultry Science 42: 79-83.CrossRefGoogle Scholar
KNUTTEL-GUSTAVSEN, S. and HARMEYER, J. (2007) The determination of L-carnitine in several food samples. Food Chemistry 105: 793-804.Google Scholar
KOBAYASHI, D., IROKAWA, M., MAEDA, T., TSUJI, A. and TAMAI, I. (2005) Carnitine/organic cation transporter OCTN2-mediated transport of carnitine in primary-cultured epididymal epithelial cells. Reproduction 130: 931-937.CrossRefGoogle ScholarPubMed
KOGUT, M.H. (2009) Impact of nutrition on the innate immune response to infection in poultry. Journal of Applied Poultry Research 18: 111-124.CrossRefGoogle Scholar
KREMSER, K., STANGL, H., PAHAN, K. and SINGH, I. (1995) Nitric oxide regulates peroxisomal enzyme activities. European Journal of Clinic Chemistry and Clinic Biochemistry 33: 763-774.Google Scholar
LA COUNT, D.W., DRACKLEY, J.K. and WEIGEL, D.J. (1995) Responses of dairy cows during early lactation to ruminal or abdominal administration of L-carnitine. Journal of Dairy Science 78: 1824-1836.Google Scholar
LEIBETSEDER, J. (1995) Studies on effects of L-carnitine in poultry. Archives of Animal Nutrition 48: 97-108.Google ScholarPubMed
LENZI, A., PICARDO, M., GANDINI, L. and DONDERO, F. (1996) Lipids of the sperm plasma membrane: from polyunsaturated fatty acids considered as markers of sperm function to possible scavenger therapy. Human Reproduction Update 2: 246-256.Google Scholar
LENZI, A., LOMBARDO, F., SGRO, P., SALACONE, P., CAPONECCHIA, L., DONDERO, F. and GANDINI, L. (2003) Use of carnitine therapy in selected cases of male factor infertility: a double-blind crossover trial. Fertility and Sterility 79: 292-300.Google Scholar
LENZI, A., SGRO, P., SALACONE, P., PAOLI, D., GILIO, B., LOMBARDO, F., SANTULLI, M., AGARWAL, A. and GANDINI, L. (2004) A placebo-controlled double-blind randomized trial of the use of combined L-carnitine and l-acetyl-carnitine treatment in men with asthenozoospermia. Fertility and Sterility 81: 1578-1584.Google Scholar
LETTNER, V.F., ZOLLITSCH, W. and HALBMAYER, E. (1992) Use of L-carnitine in the broiler ration. Bodenkultur 43: 161-167.Google Scholar
LIEDTKE, A.J., NELLIS, S.H., WHITESELL, L.F. and MAHAR, C.Q. (1982) Metabolic and mechanical effects using L- and D-carnitine in working swine hearts. Heart and Circulatory Physiology 243: H691-H697.Google Scholar
LIEN, T.F. and HORNG, Y.M. (2001) The effect of supplementary dietary L-carnitine on the growth performance, serum components, carcass traits and enzyme activities in relation to fatty acid beta-oxidation of broiler chickens. British Poultry Science 42: 92-95.CrossRefGoogle Scholar
LIN, D.S., CONNOR, W.E. and ANDERSON, G.J. (1991) The incorporation of n-3 and n-6 essential fatty acids into the chick embryo from egg yolks having vastly different fatty acid compositions. Pediatric Research 29: 601-605.CrossRefGoogle ScholarPubMed
LIU, J., HEAD, E., KURATSUNE, H., COTMAN, C.W. and AMES, B.N. (2004) Comparison of the effects of L-carnitine and acetyl-l-carnitine on carnitine levels, ambulatory activity, and oxidative stress biomarkers in the brain of old rats. Annals of the New York Academy of Sciences 1033: 117-131.Google Scholar
MARDONES, C., RIOS, A., VALCARCEL, M. and CICCIARELLI, R. (1999) Enantiomeric separation of D- and L-carnitine by integrating on-line derivatization with capillary zone electrophoresis. Journal of Chromatography A 849: 609-616.Google Scholar
MARSH, J.A. (1994) Neuroendocrine-immune interactions. Poultry Science 73: 1049-1061.Google Scholar
MAST, J., BUYSE, J. and GODDEERIS, B.M. (2000) Dietary L-carnitine supplementation increases antigen-specific Ig G production in broiler chickens. British Journal of Nutrition 83: 161-166.Google Scholar
MATSUDA, K., YUASA, H. and WATANABE, J. (1998) Fractional absorption of L-carnitine after oral administration in rats: evaluation of absorption site and dose dependency. Biological and Pharmaceutical Bulletin 21: 752-755.CrossRefGoogle ScholarPubMed
METZLER, D.E. and METZLER, C.M. (2003) Biochemistry: The chemical reactions of living cells. 2nd Edition, Vol. 1 and 2, pp. 944-945 (New York, Academic Press).Google Scholar
MILES, E.A. and CALDER, P.C. (1998) Modulation of immune function by dietary fatty acids. Proceedings of the Nutrition Society 57: 277-292.Google Scholar
MISRA, K.B., JOHNSON, W.A. and MEDEIROS, D.M. (2004) Stimulation by carnitine of branched-chain alfa keto acid dehydrogenase in intact heart mitochondria of rats. Nutrition Research 24: 647-657.CrossRefGoogle Scholar
MORAN, E.T. Jr. (2007) Nutrition of the developing embryo and hatchling. Poultry Science 86: 1043-1049.Google Scholar
MORETTI, S., ALESSE, E., DI MARZIO, L., ZAZZERONI, F., RUGGERI, B., MARCELLINI, S., FAMULARO, G., STEINBERG, S.M., BOSCHINI, A., CIFONE, M.G. and DE SIMONE, C. (1998) Effect of L-carnitine on human immuno deficiency virus-1 infection-associated apoptosis: a pilot study. Blood 91: 3817-3824.CrossRefGoogle Scholar
MURRAY, R.K., GRANNER, D.K., MAYES, P.A. and RODWELL, V.W. (2003) Harper's illustrated biochemistry. 26th Edition. pp: 181 (New York, Lange Medical Books/McGraw-Hill).Google Scholar
MUSSER, R.E., GOOD BAND, R.D., TOKACH, M.D., OWEN, K.Q., NELSSEN, J.L., BLUM, S.A., DRITZ, S.S. and CIVIS, C.A. (1999) Effects of L-carnitine fed during gestation and lactation on sow and litter performance. Journal of Animal Science 77: 3289-3295.Google Scholar
NAKANISHI, T., HATANAKA, T., HUANG, W., PRASAD, P.D., LEIBACH, F.H., GANAPATHY, M.E. and GANAPATHY, V. (2001) Na+ and Cl--coupled active transport of carnitine by the amino acid transporter ATB(0,+) from mouse colon expressed in HRPE cells and Xenopus oocytes. The Journal of Physiology 532: 297-304.Google Scholar
NEUMAN, S.L., Lin, T.L. and Hester, P.Y. (2002) The effect of dietary carnitine on semen traits of White Leghorn roosters. Poultry Science 81: 495-503.CrossRefGoogle ScholarPubMed
NOBLE, R.C. and COCCHI, M. (1990) Lipid metabolism and the neonatal chicken. Progress in Lipid Research 29: 107-140.CrossRefGoogle ScholarPubMed
NOUBOUKPO, K.E., TONA, K., AGBONON, A., GBEASSOR, M., BUYSE, J. and DECUYPERE, E. (2010) Effects of L-carnitine supplementation in drinking water on layer-type chick juvenile performance. Archiv für Geflügelkunde 74(2): S. 116-120.Google Scholar
OWEN, K.Q., NELSSEN, J.L., GOODBAND, R.D., WEEDEN, T.L. and BLUM, S.A. (1996) Effect of L-carnitine and soy-bean oil on growth performance and body composition of early-weaned pigs. Journal of Animal Science 74: 1612-1619.Google Scholar
OWEN, K.Q., JIT, H., MAXWELL, C.V., NELSSEN, J.L., GOODBAND, R.D., TOKACH, M.D., TREMBLAY, G.C. and KOO, S.I. (2001) Dietary L-carnitine suppresses mitochondrial branched chain keto acid dehydrogenase activity and enhances protein accretion and carcass characteristics of swine. Journal of Animal Science 79: 3104-3112.Google Scholar
PALMERO, S., LEONE, M., PRATI, M., COSTA, M., MESSENI LEONE, M., FUGASSA, E. and DE CECCO, L. (1990) The effect of l-acetylcarnitine on some reproductive functions in the oligoasthenospermic rat. Hormone and Metabolic Research 22: 622-626.Google Scholar
PALMERO, S., BOTTAZZI, C., COSTA, M., MESSENI LEONE, M. and FUGASSA, E. (2000) Metabolic effects of L-carnitine on prepubertal rat sertoli cells. Hormone and Metabolic Research 32: 87-90.Google Scholar
PAUL, H.S. and ADIBI, S.A. (1978) Effect of carnitine on branched-chain amino acid oxidation by liver and skeletal muscle. American Journal of Physiology - Endocrinology and Metabolism 234: E494-E499.Google Scholar
PEEBLES, E.D., KIDD, M.T., MCDANIEL, C.D., TANKSLEY, J.P., PARKER, H.M., CORZO, A. and WOOD WORTH, J.C. (2007) Effects of breeder hen age and dietary L-carnitine on progeny embryogenesis. British Poultry Science 48: 299-307.Google Scholar
RABIE M.H., , SZILAGYI, M. and GIPPERT, T. (1997a) Effects of dietary L-carnitine supplementation and protein level on performance and degree of meatiness and fatness of broilers. Acta Biologica Hungarica 48: 221-229.Google Scholar
RABIE, M.H., SZILAGYI, M., GIPPERT, T., VOTISKY, E. and GERENDAI, D. (1997b) Influence of dietary L-carnitine on performance and carcass quality of broiler chickens. Acta Biologica Hungarica 48: 241-252.CrossRefGoogle ScholarPubMed
RABIE, M.H., SZILAGYI, M. and GIPPER, T. (1997c) Effects of dietary L-carnitine on the performance and egg quality of laying hens from 65-73 weeks of age. British Journal of Nutrition 78: 615-623.Google Scholar
RABIE, M.H. and SZILAGYI, M. (1998) Effects of Lcarnitine supplementation of diets differing in energy levels on performance, abdominal fat content and yield and composition of edible meat of broilers. British Journal of Nutrition 80: 391-400.Google Scholar
RAMADAN, L.A., ABD-ALLAH, A.R., ALY, H.A.A. and SAAS-EL-DIN, A.A. (2002) Testicular toxicity effects of magnetic field exposure and prophylactic role of coenzyme Q10 and L-carnitine in mice. Pharmacological Research 46: 363-370.Google Scholar
RANI, P.J. and PANNEERSELVAM, C. (2002) Effect of carnitine on brain lipid peroxidation and antioxidant enzymes in old rats. Journal of Gerontology 57: B134E137.Google ScholarPubMed
REBOUCHE, C.J. and PAULSON, D.J. (1986) Carnitine metabolism and function in humans. Annual Review of Biochemistry 6: 41-66.Google Scholar
REBOUCHE, C.J. (1992) L-carnitine function and requirements during the life cycle. Journal of the Federation of American Societies for Experimental Biology 6: 3379-3386.Google Scholar
REDDY, R.P.K. and SAJADI, A. (1990) Selection for growth and semen traits in the poultry industry: What can we expect in the future? in: Control of Fertility in Domestic Birds. INRA, Paris, pp. 47-59.Google Scholar
REZAEI, M., DEHGHANI, S., GHAFFARI, J. and HAGHNAZAR, A. (2008) The effects of different levels of L-carnitine and fat on performance and egg quality of laying hens. Agricultural and Food Science 17: 360-366.Google Scholar
ROBINSON, D.S. (1987) The chemical basis of albumen quality. in: WELLS, R.G. & BELYAVIN, C.G. (Eds) Egg quality-current problems and recent advances, pp. 179-191 (London, Butterworths Ltd).Google Scholar
RODEHUTSCORD, M., TIMMLER, R. and DIECKMANN, A. (2002) Effect of L-carnitine supplementation on utilisation of energy and protein in broiler chicken fed different dietary fat levels. Archives of Animal Nutrition 56: 431-441.Google Scholar
SANOCKA, D. and KURPISZ, M. (2004) Reactive oxygen species and sperm cells. Reproductive Biology and Endocrinology 2: 12.CrossRefGoogle ScholarPubMed
SARICA, S., CORDUK, M. and KILINC, K. (2005) The effect of dietary L-carnitine supplementation on growth performance, carcass traits and composition of edible meat in Japanese quail (Coturnix coturnix japonica). Journal of Applied Poultry Research 14: 709-715.CrossRefGoogle Scholar
SARICA, S., CORDUK, M., ENSOY, U., BASMACIOGLU H., and KARATAS, U. (2007) Effects of dietary supplementation of L-carnitine on performance, carcass and meat characteristics of quails. South African Journal of Animal Science 37: 189-201.Google Scholar
SHAW R.D., , LI, B.U., HAMILTON, J.W., SHUG, A.L. and OLSEN, W.A. (1983) Carnitine transport in rat small intestine. American Journal of Physiology 245: G376-G381.Google Scholar
SHUG, A.L. and GRAVENSTEIN, S. (1996) Method of stimulating antibody formation. United States Patent 5,569,457.Google Scholar
SIDDIQUE, R.A., TANDON, M., AMBWANI, T., RAI, S.N. and and ATREJA, S.K. (2009) Nutrigenomics: nutrient-gene interactions. Food Reviews International 25: 326-345.Google Scholar
SMITH, C., MARKS, A. and LIEBERMAN, M.A. (2004) Marks’ basic medical biochemistry: a clinical approach, 2nd edition, pp. 418-430 (Lippincott Williams & Wilkins).Google Scholar
SONAIYA, E.B. (1985) Abdominal fat weight and thickness as predictors of total body fat in broilers. British Poultry Science 26: 453-458.Google Scholar
SPEAKE, B.K., MURRAY, A.M. and NOBLE, R.C. (1998) Transport and transformations of yolk lipids during development of the avian embryo. Progress in Lipid Research 37: 1-32.CrossRefGoogle ScholarPubMed
SUCHY, P., STRAKOVA, E. and VITULA, F. (2008) The effect of a diet supplemented with L-carnitine on egg production in pheasant (Phasianus colchicus). Czech Journal Animal Science 53(1): 31-35.Google Scholar
SUR, J.H., DOSTER, U., CHRISTIAN, J.S., GALEOTA, J.A., WILLS, R.W., ZIMMERMAN, J.J. and OSORIO, F.A. (1997) Porcine reproductive and respiratory syndrome virus replicates in testicular germ cells, alters spermatogenesis and induces germ cell death by apoptosis. The Journal of Virology 71: 9170-9179.Google Scholar
SURAI, P.F., BLESBOIS, E., GRASSEAU, I., CHALAH, T., BRILLARD, J.P., WISHART, G.J., CEROLINI, S. and SPARKS, N.H.C. (1998) Fatty acid composition, glutathione peroxidase, and SOD activity and total antioxidant activity of avian semen. Comparative Biochemistry and Physiology - Part B 120: 527-533.Google Scholar
SURAI, P.F. (1999) Tissue-specific changes in the activities of antioxidant enzymes during the development of the chicken embryo. British Poultry Science 40: 397-405.Google Scholar
SURAI, P.F. (2007) Natural antioxidants in poultry nutrition: new developments. Proceedings of the 16th European Symposium on Poultry Nutrition. Strasbourg , France, pp. 26-30.Google Scholar
THIEMEL, J. and JELINEK, P. (2004) The effect of carnitine on hatching rate end metabolic profile of blood in breeding layers. Czech Journal Animal Science 49: 517-523.Google Scholar
TOMITA, M. and SENDJU, Y. (1927) Über die oxyaminoverbindungen, welche die Biuretreaktion zeigen. III. Spaltung der g-Amino-b-oxy-buttersäure in die optisch-aktiven Komponenten. Hoppe-Seyler's Zeitschrift fürPhysiologische Chemie 169: 263-277.Google Scholar
VAN HINSBERGH, V.W., VEERKAMP, J.H. and CORDEWENER, J.H. (1980) Effect of carnitine and branched-chain acylcarnitines on the 2-oxo acid dehydrogenase activity in intact mitochondria of rat muscle. International Journal of Biochemistry 12: 559-565.CrossRefGoogle ScholarPubMed
WALTER, P. and SCHAFFHAUSER, A.O. (2000) L-Carnitine a vitamin-like substance for functional food. Annals of Nutrition and Metabolism 44: 75-96.Google Scholar
XU, Z.R., WANG, M.Q., MAO, H.X., ZHAN, X.A. and HU, C.H. (2003) Effects of L- carnitine in growth performance, carcass composition, and metabolism of lipids in male broilers. Poultry Science 82: 408-413.Google Scholar
ZEYNER, A. and HARMEYER, J. (1999) Metabolic functions of L-carnitine and its effects as feed additive in horses. Archives of Animal Nutrition 52: 115-138.Google Scholar
ZHAI, W., NEUMAN, S., LATOUR, M.A. and HESTER, P.Y. (2007) The effect of dietary L-carnitine on semen traits of white leghorns. Poultry Science 86: 2228-2235.Google Scholar
ZHAI, W., NEUMAN, S., LATOUR, M.A. and HESTER, P.Y. (2008a) The effect of in ovo injection of L-carnitine on hatchability of white leghorns. Poultry Science 87: 569-572.CrossRefGoogle ScholarPubMed
ZHAI, W., NEUMAN, S., LATOUR, M.A. and HESTER, P.Y. (2008b) The effect of male and female supplementation of L-carnitine on reproductive traits of white leghorns. Poultry Science 87: 1171-1181.Google Scholar