Hostname: page-component-cd9895bd7-jn8rn Total loading time: 0 Render date: 2024-12-27T12:49:05.691Z Has data issue: false hasContentIssue false
  • English
  • Français

The nutritional biochemistry of wool and hair follicles

Published online by Cambridge University Press:  18 August 2016

P.I. Hynd
P.I. Hynd*
Affiliation:
Department of Animal Science, The University of Adelaide, Roseworthy Campus, Roseworthy South Australia, Australia 5371
Get access

Abstract

The rôle of various classes of nutrients (energy substrates, vitamins, minerals, amino acids) in the production of wool and hair from follicles, is considered for a variety of animal species. The wool and hair follicle have evolved a number of interesting features of carbohydrate metabolism including glutaminolysis, aerobic glycolysis, significant activity of the pentose phosphate pathway, and storage and mobilisation of glycogen. Presumably the necessity to continue to produce fibre despite fluctuations in the supply of oxygen and nutrients has resulted in some of these unique features, while others reflect the high level of DNA and protein synthesis occurring in the follicle. While it is considered that energy does not normally limit fibre growth, the relative contributions of aerobic and anerobic metabolism will greatly influence the amount of ATP available for follicle activity, such that energy availability may at times alter fibre growth. Alopecia and deficient fibre growth are consistent outcomes of deficiencies of biotin, riboflavin, pyridoxine, folate and pantothenic acid, but the precise rôles of these vitamins in follicle function await elucidation. Folate, in particular appears to play an important rôle in wool production, presumably reflecting its involvement in methionine metabolism. Cholecalciferol (vitamin D) significantly alters fibre growth in cultured follicles; vitamin D receptors are located in the outer root sheath, bulb, and dermal papilla of the follicle; and alopecia occurs in humans with defects in the vitamin D receptor. Retinol (vitamin A), too, appears to influence follicle function by altering keratinocyte proliferation and differentiation, with direct effects on the expression of keratin genes. The receptors for the retinoids are present in the keratogenous zone, the outer root sheath, the bulb, and the sebaceous glands. Vitamin A may also act indirectly on follicle function by influencing the activity of the insulin-like and epidermal growth factors and by altering vitamin D activity. At present there is little evidence implicating alpha-tocopherol (vitamin E) or phytylmenaquinone (vitamin K) in follicular events. Of the minerals, only copper and zinc have been shown to have direct effects on follicle function, independent of effects on food intake. Copper has direct effects on the activity of an unidentified enzyme on oxidation of thiol groups to form disulphide linkages. Wool produced by copper-deficient sheep lacks crimp, is weak and lustrous. Copper is also necessary for the activity of tyrosinase and the tyrosinase-related proteins involved in melanin synthesis. Zinc, like copper, is required for the normal keratinization of fibres but again, the precise rôle has yet to be elucidated. While the importance of amino acid supply for wool growth has long been established, there are still some unaswered questions such as; what are the effects of amino acids on fibre growth in animals other than sheep; what are the characteristics of the amino acid transport genes and proteins operating in the wool and hair follicle; and what are the specific rôles for amino acids in follicle function.

Keywords

biochemistryfollicleshairwool
Type
Invited paper
Information
Animal Science , Volume 70 , Issue 2 , April 2000 , pp. 181 - 195
Copyright
Copyright © British Society of Animal Science 2000

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Adachi, K. and Uno, H. 1968. Glucose metabolism of growing and resting human hair follicles. American Journal of Physiology 215: 1234-1239.Google Scholar
Allden, W. G. 1979. Feed intake, diet composition and wool growth. In Physiological and environmental limitations to wool growth (ed. Black, J. L. and Reis, P. J.), p. 61. University of New England Publishing Unit, Armidale NSW, Australia.Google Scholar
Ash, A.J. and Norton, B.W. 1987. Effect of DL-methionine supplementation on fleece growth by Australian cashmere goats. Journal of Agricultural Science, Cambridge 109: 197–199.Google Scholar
Balnave, D. 1977. Clinical symptoms of biotin deficiency in animals. American Journal of Clinical Nutrition 30: 1408–1413.Google Scholar
Bates, E. J., Hynd, P.I., Pernio, N. M. and Nancarrow, M. J. 1997. Serum-free culture of wool follicles: effects of nutrients, growth factors and hormones. British Journal of Dermatology 137: 498–505.Google Scholar
Benevenga, N. J. and Egan, A. R. 1983. Quantitative aspects of methionine metabolism. Progess in Clinical and Biological Research 125: 327–341.Google Scholar
Black, J. L. 1987. Mechanisms controlling the rate of growth, composition and morphology of wool. In Merino improvement programs in Australia (ed. McGuirk, B. J.), p. 457. Australian Wool Corporation, Melbourne, Australia.Google Scholar
Black, J. L. and Reis, P. J. 1979. Speculation on the control of nutrient partitioning between wool growth and other body functions. In Physiological and environmental limitations to wool growth (ed. Black, J. L. and Reis, P. J.), pp. 269–293. University of New England Publishing Unit, Armidale NSW, Australia.Google Scholar
Blumenberg, M., Connolly, D. M. and Freedberg, I. M. 1992. Regulation of keratin gene expression: the role of the nuclear receptors for retinole acid, thyroid hormone, and vitamin D3. Journal of Investigative Dermatology 98: 42S–49S.Google Scholar
Bond, J. J., Wynn, P.C., Brown, G.N. and Moore, G.P. 1994. Growth of wool follicles in culture. In Vitro Cellular and Developmental Biology. Animal 30A: 90–98.CrossRefGoogle ScholarPubMed
Campbell, F.R., Livingston, C.W. Jr and Watkins, T.D. 1958. Influence of vitamin A on wool growth, wool strength and reproductive performance in Rambouillet ewes. Journal of Animal Science 17: 1194 (abstr. ).Google Scholar
Catteneo, S.M., Quastler, H. and Sherman, F.G. 1961. Proliferative cycle in the growing hair follicle of the mouse. Nature 190: 923–924.Google Scholar
Chapman, R. E. and Black, J. L. 1981. Abnormal wool growth and alopecia of artificially-reared lambs. Australian Journal of Biological Sciences 34:11–26.Google Scholar
Chapman, R.E., and Ward, K. A. 1979. Histological and biochemical features of the wool fibre and follicle. In Physiological and environmental limitations to wool growth (ed. Black, J. L. and Reis, P. J.), p. 193. University of New England Publishing Unit, Armidale NSW, Australia.Google Scholar
Damak, S., Su, H., Jay, N.P. and Bullock, D. W. 1996. Improved wool production in transgenic sheep expressing insulin-like growth factor 1. Biotechnology 14:185–188.Google Scholar
Danks, D. M. 1991. Copper deficiency and the skin. In Biochemistry and physiology of the ski. (ed. Goldsmith, L.A.), pp. 1351–1361. Oxford University Press, Oxford.Google Scholar
Doncon, G.H. and Steele, P. 1988. Plasma and liver concentrations of alpha-tocopherol in weaner sheep after vitamin E supplementation. Australian Veterinary Journal 65: 210–213.Google Scholar
Downes, A.M., Lyne, A.G. and Clarke, W.H. 1962. Radioautographic studies of the incorporation of [35S]cystine into wool. Australian Journal of Biological Sciences 15: 713–719.Google Scholar
Doyle, P. and Bird, S. 1975. The influence of dietary supplements of DL-Methionine on the growth rate of wool. Australian Journal of Agricultural Research 26: 337–342.Google Scholar
Draper, H. H. and Johnson, B.C. 1952. Folie acid deficiency in the lamb. Journal of Nutrition 46:123–131.Google Scholar
Eichner, R., Kahn, M., Capetola, R.J., Gendimenico, G. J. and Mezick, J. A. 1992. Effects of topical retinoids on cytoskeletal proteins: implications for retinoid effects on epidermal differentiation. Journal of Investigative Dermatology 98:154–161.Google Scholar
Eida, K., Rubato, N., Nishigaki, T. and Kikutani, M. 1975. Harderian gland. V. Effect of dietary pantothenic acid deficiency on porphyrin synthesis in Harderian gland of rats. Chemical Pharmacology Bulletin (Tokyo) 23: 1–4.Google Scholar
Eller, M. S., Harkness, D. D., Bhawan, J. and Gilchrest, B.A. 1994. Epidermal differentiation enhances CRABP II expression in human skin. Journal of Investigative Dermatology 103: 785–790.Google Scholar
Ermolova, L. 1982. Metabolism of lipids in the rumen and productivity of fine-fleeced sheep fed on a diet containing pelleted feeds supplemented with vitamin A. Proceedings of the second Mezhdunarodnyi simpozium po fiziologii pishchevareniya zhvachnykh I ikh produktivnosti, Stara Zagora, Bulgaria, 34.Google Scholar
Eveleth, D.E., Bolin, D. W. and Goldsby, A. I. 1949. Experimental avitaminosis A in sheep. American Journal of Veterinary Research 10: 250–255.Google Scholar
Farid, M. F. A. and Ghanen, Y. S. 1982. Vitamin A deficiency and supplementation in desert sheep. II. Wool characteristics. World Review of Animal Production 18: 75–77.Google Scholar
Finzi, E., Blake, M. J., Celano, P., Skouge, J. and Diwan, R. 1992. Cellular localization of retinole acid receptor-gamma expression in normal and neoplastic skin. American Journal of Pathology 140:1463–1471.Google Scholar
Fisher, D. A. and Lakshmanan, J. 1990. Metabolism and effects of epidermal growth factor and related growth factors in mammals. Endocrine Reviews 11: 418–442.Google Scholar
Franklin, M.C., McClymont, G. L., Briggs, P. K. and Campbell, B. L. 1955. Maintenance rations for Merino sheep. II. The preformance of weaners fed daily and weekly on rations of wheat and wheaten chaff at maintenance levels and the effect of vitamin A supplements. Australian Journal of Agricultural Research 6: 324–342.Google Scholar
Freinkel, R. K. 1983. Carbohydrate metabolism of epidermis. In Biochemistry and physiology of the skin (ed. Goldsmith, L. A.), pp. 328–337. Oxford University Press, Oxford.Google Scholar
Frigg, M., Schulze, J. and Volker, L. 1989. Clinical study on the effect of biotin on skin conditions in dogs. Schweizer Archiv für Tierheilkiunde 131: 621–625.Google Scholar
Fry, J. M., McGrath, M.C., Harvey, M. and Speijers, E. J. 1996a. Vitamin E treatment of weaner sheep. II. The effect of vitamin E responsive subclinical myopathy on liveweight and wool production. Australian Journal of Agricultural Research 47: 869–876.Google Scholar
Fry, J. M., McGrath, M.C., Harvey, M., Sunderman, E., Smith, G.M. and Speijers, E.J. 1996b. Vitamin E treatment of weaner sheep. I. The effect of vitamin E supplements on plasma alpha-tocopherol concentrations, liveweight, and wool production in penned or grazing sheep. Australian Journal of Agricultural Research 47: 853–867.Google Scholar
Fuchs, E. 1990. Epidermal differentiation. Current Opinion in Cell Biologi) 2: 1028–1035.Google Scholar
Fuchs, E. and Green, H. 1981. Regulation of terminal differentiation of cultured human keratinocvtes bv vitamin A. Cell 25: 617–625.Google Scholar
Geyer, H., Pohlenz, J. and Volker, L. 1981. [Morphological and histochemical studies of skin, mucous membranes and hooves of swine in experimental biotin deficiency.] Zentralblatt für Veterinärmedizin Reihe A 28: 574–592.Google Scholar
Geyer, H., Schulze, J., Streiff, K., Tagwerker, F. and Volker, L. 1984. [Effect of experimental biotin deficiency on the morphology and histochemistry of the skin and claws in swine.] Zentralblatt für Veterinärmedizin Reihe A 31: 519–538.Google Scholar
Gillespie, J. M. 1964. The isolation and properties of some soluble proteins from wool. VIII. The proteins of copper-deficient wool. Australian Journal of Biological Sciences 17: 282–300.CrossRefGoogle Scholar
Gillespie, J.M. 1973. Keratin structure and changes with copper deficiency. Australian Journal of Dermatologi/ 15: 127–131.Google Scholar
Goldsmith, L. A. 1980. Vitamins and alopecia [editorial]. Archives of Dermatology 116:1134–1135.Google Scholar
Goos, C.M., Beaumont, A.H., Vermeesch-Markslag, A.M., Stappen, J. W. van der, Sultan, C. and Vermorken, A. J. 1987. Determination of glycogen and enzymes of glycogen metabolism in human hair follicles. Molecular Biology Reports 12: 259–264.Google Scholar
Gries, C. L. and Scott, M. L. 1972. The pathology of thiamin, riboflavin, pantothenic acid and niacin deficiencies in the chick. Journal of Nutrition 102: 1269-1285.Google Scholar
Guirgis, R. A., Ashmawy, G., El-Ezz, S. S. A. and Abu-El-Ezz, S. S. 1982. Postnatal changes in some skin and wool characteristics associated with vitamin A blood concentration. Journal of Agricultural Science, Cambridge 98: 215–219.Google Scholar
Hardy, M. H. 1992. The secret life of the hair follicle. Trends in Genetics 8: 55–61.Google Scholar
Harmon, C. S. and Nevins, T. D. 1994. Biphasic effect of 1, 25-dihydroxyvitamin D3 on human hair follicle growth and hair fibre production in whole organ cultures. Journal of Investigative Dermatology 103: 318–322.Google Scholar
Hembree, J. R., Harmon, C. S., Nevins, T. D. and Eckert, R. L. 1996. Regulation of human dermal papilla cell production of insulin-like growth factor binding protein -3 by retinole acid, glucocorticoids and insulin-like growth factor-1. Journal of Cell Physiology 167: 556–561.Google Scholar
Herrmann, S., Lange, K., and Wortmann, F. J. 1996. Characteristics of Angora rabbit fibre. 2. The influence of the methionine content in feed and of the environmental temperature on fibre and medulla diameter in Angora wool. World Rabbit Science 4:155–158.Google Scholar
Holick, M. F. 1985. Vitamin D resistance and alopecia. Archives of Dermatology 121: 601–603.Google Scholar
Holick, M. F. 1995. Noncalcaemic actions of 1, 25-dihydroxyvitamin D3 and clinical applications. Bone 17: 107S–111S.Google Scholar
Hynd, P. I. 1989. Cellular events in wool follicles. In The biology of wool and hair (ed. Rogers, G. E., Reis, P.J., Ward, K.A. and Marshall, R. C.). Chapman and Hall, London.Google Scholar
Hynd, P. I. 1994a. Follicular determinants of the length and diameter of wool fibres. I. Comparison of sheep differing in fibre length/diameter ratio at two levels of nutrition. Australian journal of Agricultural Research 45: 1137–1147.Google Scholar
Hynd, P. I. 1994b. Follicular determinants of the length and diameter of wool fibres. II. Comparison of sheep differing in thyroid hormone status. Australian journal of Agricultural Research 45: 1149–1157.Google Scholar
Hynd, P. I. and Allden, W. G. 1985. Rumen fermentation pattern, postruminal protein flow and wool growth rate of sheep on a high-barley diet. Australian Journal of Agricultural Research 36: 451–460.Google Scholar
Hynd, P. I., Hughes, A., Earl, C R. and Penno, N. M. 1997. Seasonal changes in the morphology and activity of wool follicles in Fine wool and Strongwool Merino strains at different stocking rates in southern Australia. Australian Journal of Agricultural Research 48:1087–1097.Google Scholar
Hynd, P. I. and Nancarrow, M. J. 1996. Inhibition of polyamine synthesis alters hair follicle function and fiber composition. Journal of Investigative Dermatology 106: 249–253.Google Scholar
Innis, S. M. and Allardyce, D. B. 1983. Possible biotin deficiency in adults receiving long-term parenteral nutrition. American Journal of Clinical Nutrition 37:185–187.Google Scholar
Jimbow, K., Fitzpatrick, T.B. and Wick, M. M. 1991. Biochemistry and physiology of melanin pigmentation. In Biochemistry and physiology of the skin, second edition (ed. Goldsmith, L. A.), pp. 873–909. Oxford University Press, Oxford.Google Scholar
Johnson, E. 1981. Environmental influences on the hair follicle. In Hair research (ed. Orfanos, C. E. O., Montagna, W. M. and Stuttgen, G. S.), p. 183. Springer-Verlag Berlin, Heidelberg.Google Scholar
Johnson, J. A. and Fusaro, R. M. 1972. The role of the skin in carbohydrate metabolism. Advances in Metabolic Disorders 6:1–55.Google Scholar
Kealey, T. 1983. The metabolism and hormonal responses of human eccrine sweat glands isolated by collagenase digestion. Biochemical Journal 212:143–148.Google Scholar
Kealey, T., Williams, R. and Philpott, M.P. 1991. Intermediary metabolism of the human hair follicle. Annals of the New York Academy of Sciences 642: 301–307.Google Scholar
Kealey, T., Williams, R. and Philpott, M. P. 1994. The human hair follicle engages in glutaminolysis and aerobic glycolysis: implications for skin, splanchnic and neoplastic metabolism. Skin Pharmacology 7: 41–46.Google ScholarPubMed
Kondo, S., Hozumi, Y. and Aso, K. 1990. Organ culture of human scalp hair follicles: effect of testosterone and oestrogen on hair growth. Archives of Dermatological Research 282: 442–445.Google Scholar
Krebs, H. A. 1972. The physiological role of aerobic glycolysis in various animal tissues. Essays in Biochemistry 8: 1–34.Google Scholar
Kumagai, H. and White, C. L. 1991. The effect of a supplementary vitamin and /or mineral mixture on productivity and on vitamin and glutathione status in ewes and lambs. Proceedings of the Nutrition Society of Australia 16: 208.Google Scholar
Kumagai, H. and White, C. L. 1995. The effect of supplementary minerals, retinol and alpha-tocopherol on the vitamin status and productivity of pregnant Merino ewes. Australiern Journal of Agricultural Research 46: 1159–1174.Google Scholar
Lee, H. J. 1956. The influence of copper deficiency on the fleeces of British breeds of sheep. Journal of Agricultural Science, Cambridge 47: 218–224.Google Scholar
Leng, R. A. and Stephenson, S. K. 1965. Glucose and acetate metabolism by isolated sheep wool follicles. Archives of Biochemistry and Biophysics 110: 8–15.Google Scholar
McGregor, B. A. and Hodge, R. W. 1989. Influence of energy and polymer-encapsulated methionine supplements on mohair growth and fibre diameter of Angora goats fed at maintenance. Australian Journal of Experimental Agriculture 29: 179–181.Google Scholar
Makarova, R. E. 1977. Effect of vitamin B-6 on reproductive ability of female standard mink, the growth of the young and fur quality. Krolikovodstvo i Zverovodstvo 2: 13–14.Google Scholar
Marston, H. and Lee, H. J. 1948. Nutritional factors involved in wool production by merino sheep II. The influence of copper deficiency on the rate of wool growth and on the nature of the fleece. Australian Journal of Scientific Research Series B 1: 376–387.Google Scholar
Masters, D. G., Chapman, R. E. and Vaughan, J. D. 1985. Effects of zinc deficiency on the wool growth, skin and wool follicles of pre-ruminant lambs. Australian Journal of Biological Sciences 38: 355–364.Google Scholar
Matusis, I. I., Rapoport, B. N. and Elistratova, N. B. 1976. Vitamins K. and E and functional state of connective tissues. Voprosy Pitaniya 3: 27–30.Google Scholar
Mills, C.R., Dalgarno, A. C., Williams, R. B. and Quarterman, J. 1967. Zinc deficiency and the zinc requirements of calves and lambs. British Journal of Nutrition 21: 751–768.Google Scholar
Montagna, W. 1962. The pilary system. In The structure and function of skin, second edition (ed. Montagna, W.), pp. 174–260. Academic Press, New York.Google Scholar
Moore, G. P., Du-Cros, D. L., Isaacs, K., Pisansarakit, P. and Wynn, P. C. 1991. Hair growth induction: roles of growth factors. Annals of the New York Academy of Sciences 642: 308–325.Google Scholar
National Research Council. 1978. Nutrient requirements of laboratory animals. National Academy of Sciences, Washington, DC.Google Scholar
Neldner, K.H. 1983. The biochemistry and physiology of zinc metabolism. In Biochemistry and physiology of the skin (ed. Goldsmith, L. A.), pp. 1082–1101. Oxford University Press, Oxford.Google Scholar
Nelson, D. R., Wolff, W. A., Blodgett, D. J., Luecke, B., Ely, R. W. and Zachary, J. F. 1984. Zinc deficiency in sheep and goats: three field cases. Journal of the American Veterinary Medical Association 184:1480–1485.Google Scholar
Noji, S., Yamaai, T., Koyama, E., Nohno, T., Fujimoto, W., Arata, J. and Taniguchi, S. 1989. Expression of retinoic acid receptor genes in keratinizing front of skin. FEBS Letters 259: 86–90.Google Scholar
Ohkawara, A. 1976. Glycogen metabolism and cyclic AMP in the hair follicle. In Biology and disease of the hair (ed. Toda, K. et al. ) Baltimore University Park Press, USA.Google Scholar
Ott, E.A., Smith, W.H., Stob, M., and Beeeson, W.M. 1964. Zinc deficiency syndrome in the young lamb. Journal of Nutrition 82: 41–50.Google Scholar
Ott, E.A., Smith, W.H., Stob, M., Parker, H. E., Harrington, R.B., and Beeson, W. M. 1965. Zinc requirements of the growing lamb fed a purified diet. Journal of Nutrition 87: 459–463.Google Scholar
Pegg, A. E. and McCann, P. P. 1982. Polyamine metabolism and function. American Journal of Physiologi/ 243: C212–221.Google Scholar
Philpott, M. P., Green, M. R. and Kealey, T. 1990. Human hair growth in vitro. Journal of Cell Science 97: 463–471.Google Scholar
Philpott, M. P. and Kealey, T. 1991. Metabolic studies on isolated hair follicles: hair follicles engage in aerobic glycolysis and do not demonstrate the glucose fatty acid cycle. Journal of Investigative Dermatology 96: 875–879.Google Scholar
Philpott, M. P., Sanders, D. A. and Kealey, T. 1994. Effects of insulin and insulin-like growth factors on cultured human hair follicles: IGF-1 at physiologic concentrations is an important regulator of hair follicle growth in vitro. Journal of Investigative Dermatology 102: 857–861.Google Scholar
Podshibyakin, A. E., Sapunov, A. G., Golovskoi, I. P. and Chebotarev, I. I. 1988. Use of a complex mixture of vitamin A and trace elements as a preventive against vitamin A deficiency in sheep. Doklady Vsesoyuznoï Akademii Seľskokhozyaistvennykh Nauk 2: 32–34.Google Scholar
Powell, B.C. and Rogers, G.E. 1997. The role of keratin proteins and their genes in the growth, structure and properties of hair. In Formation and structure of human hair (ed. Jolie, P., Zahn, H. and Hocker, H.), pp. 59–149. Birkhauser Verlag, Basel.Google Scholar
Purser, D. B. 1979. Minerals and wool growth. In Physiological and environmental limitations to wool growth (ed. Black, J. L. and Reis, P. J.), pp. 243–256. University of New England Publishing Unit, Armidale NSW, Australia.Google Scholar
Reichrath, J., Mittmann, M., Kamradt, J. and Muller, S. M. 1997. Expression of retinoid X receptors (-alpha, -beta, -gamma) and retinoic acid receptors (alpha, beta, gamma) in normal skin: an immunohistological evaluation. Histochemical Journal 29: 127–133.Google Scholar
Reichrath, J., Munssinger, T., Kerber, A., Rochette Egly, C., Chambon, P., Bahmer, F. A. and Baum, H. P. 1995. In situ detection of retinoid-X receptor expression in normal and psoriatic human skin. British Journal of Dermatology 133: 168–175.Google Scholar
Reichrath, J., Schilli, M., Kerber, A., Bahmer, F. A., Czarnetzki, D. M. and Paus, R. 1994. Hair follicle expression of 1, 25-dihydroxyvitamin D3 receptors during the murine hair cycle. British Journal of Dermatology 131: 477–482.Google Scholar
Reis, P.J. 1967. The growth and composition of wool. IV. The differential response of growth and of sulphur content of wool to the level of sulphur-containing amino acids given per abomasum. Australian Journal of Biological Sciences 20: 809–825.Google Scholar
Reis, P.J. 1970. The influence of abomasal supplements of some amino acids and sulphur-containing compounds on wool growth rate. Australian Journal of Biological Sciences 23: 441–446.Google Scholar
Reis, P. J. 1979. Effects of amino acids on the growth and properties of wool. In Physiological and environmental ¡imitations to wool groīuth (ed. Black, J. L. and Reis, P. J.), pp. 223–242. University of New England Publishing Unit, Annidale NSW, Australia.Google Scholar
Reis, P.J. 1982. The importance of methionine for wool growth in sheep. Proceedings of the Australian Society of Animal Production 14: 479–482.Google Scholar
Reis, P. J. 1989. The influence of absorbed nutrients on wool growth. In The biology of wool and hair (ed. Rogers, G. E., Reis, P.J., Ward, K.A. and Marshall, R.C.), pp. 185–203. Chapman and Hall, London.Google Scholar
Reis, P.J. and Colebrook, W.F. 1972. The utilization of abomasal supplements of proteins and amino acids by sheep with special reference to wool growth. Australian journal of Biological Sciences 25: 1057–1071.Google Scholar
Reis, P. J. and Hynd, P. I. 1989. The influence of difluoromethylornithine on the activity of wool follicles. Australasian journal of Animal Science 2: 204–205.Google Scholar
Reis, P. J. and Sahlu, T. 1994. The nutritional control of the growth and properties of mohair and wool fibers: a comparative review. Journal of Animal Science 72: 1899–1907.Google Scholar
Reis, P. J. and Tunks, D. A. 1976. The influence of abomasal supplements of zein and some amino acids on wool growth rate and plasma amino acids. Journal of Agricultural Science, Cambridge 86: 475–482.Google Scholar
Reis, P.J. and Tunks, D. A. 1978. Effects on wool growth of the infusion of mixtures of amino acids into the abomasum of sheep. Journal of Agricultural Science, Cambridge 90: 173–183.Google Scholar
Reis, P. J., Tunks, D. A. and Munro, S. G. 1990. Effects of the infusion of amino acids into the abomasum of sheep, with emphasis on the relative value of methionine, cysteine and homocysteine for wool growth. Journal of Agricultural Science, Cambridge 114: 59–68.Google Scholar
Reis, P. J., Tunks, D. A. and Munro, S. G. 1992. Effects of abomasal protein and energy supply on wool growth in Merino sheep. Australian Journal of Agricultural Research 43: 1353.Google Scholar
Rolinski, Z. and Duda, M. 1986. Use of biotin in the treatment of dogs and breeding foxes. Medycyna Weterynaryjna 42: 223–226.Google Scholar
Russel, A. J. F. 1992. Fibre production from sheep and goats. In Progress in sheep and goat research (ed. Speedy, A. W.), p. 235. CAB International, Wallingford, UK.Google Scholar
Russell, L. E., Easter, R. A. and Bechtel, P. J. 1985. Evaluation of erythrocyte asparate activity coefficient as an indicator of the vitamin B-6 status of postpubertal gilts. Journal of Nutrition 115: 1117-1123.Google Scholar
Ryder, M. L. 1958a. Some observations on the glycogen content of the wool follicle. Quarterly Journal of Microscopical Science 99: 221–228.Google Scholar
Ryder, M. L. 1958b. Investigations into the distribution of thiol groups in the skin follicles of mice and sheep and the entry of labelled sulphur compounds. Proceedings of the Royal Society of Edinburgh 67: 65–82.Google Scholar
Ryder, M. L. 1959. Studies of sulphur, phosphorus and some metals in skin and follicles. Journal of Histochemistry and Cytochemistry 9: 133–138.Google Scholar
Ryder, M. L. and Stephenson, S. K. 1968. Wool growth Academic Press, London.Google Scholar
Sadamoto, Y., Arase, S., Kato, S., Nakanishi, H., Fujie, K., Takeda, E. and Takeda, K. 1990. The effect of 1, 25-dihydroxyvitamin D3 on cultured human hair follicle cells from patients with vitamin D-dependent rickets type II with alopecia. Nippon Hifuka Gakkai Zasshi 100: 211–214.Google Scholar
Sahlu, T. and Fernandez, J. M. 1992. Effect of intraperitoneal administration of lysine and methionine on mohair yield and quality in Angora goats. Journal of Animal Science 70: 3188–3193.Google Scholar
Saraswat, R. C. and Arora, S.P. 1972. Zinc deficiency syndromes in lambs. Indian Veterinary Journal 49: 701–704.Google Scholar
Schlink, A. C. and Dollin, A.E. 1995. Abnormal shedding contributes to the reduced staple strength of tender wool in Western Australian Merinos. Wool Technology and Sheep Breeding 43: 268–284.Google Scholar
Schulze, A. and Ustdal, K. M. 1975. Possible etiology of alopecia in Turkish angora goats. Berliner und Munchener Tierarztlicke Wochenschrift 88: 66–70.Google Scholar
Scott, E. J.van, Ekel, T. M. and Auerbach, R. 1963. Determinants of rate and kinetics of cell division in scalp hair. Journal of Investigative Dermatology 41: 269–273.Google Scholar
Sherertz, E.F. and Goldsmith, L. A. 1991. Nutritional influences on the skin. In Physiology, biochemistry, and molecular biology of the skin (ed. Goldsmith, L. A.), p. 1315. Oxford University Press, Oxford.Google Scholar
Shipman, M., Chase, H. B. and Montagna, W. 1955. Glycogen in skin of the mouse during cycles of hair growth. Proceedings of the Society for Experimental and Biological Medicine 88: 449–451.CrossRefGoogle ScholarPubMed
Smith, R. M., Hocking Edwards, J. E. and Schlink, A. C. 1998. Sheep skin glycogen content responds to level of nutrition. Proceedings of the Australian Society of Animal Production 22: 309.Google Scholar
Smuts-Ayers, M. and Sahlu, T. 1996. The effect of various levels of lysine infused intraperitoneally on mohair quality and nitrogen metabolism in Angora goats. Small Ruminant Research 20: 113–119.Google Scholar
Spritz, R. A., Ho, L., Furumura, M. and Hearing, V. J. Jr 1997. Mutational analysis of copper binding by human tyrosinase. Journal of Investigative Dermatology 109: 207–212.Google Scholar
Steele, P., Peet, R. L., Skirrow, S., Hopkinson, W. and Masters, H. G. 1980. Low alpha-tocopherol levels in livers of weaner sheep with nutritional myopathy. Australian Veterinary Journal 56: 529–532.Google Scholar
Stein, E. D. and Diamond, J. M. 1989. Do dietary levels of pantothenic acid regulate its intestinal uptake in mice? Journal of Nutrition 119: 1973–1983.Google Scholar
Stumpf, W. E., Clark, S. A., Sar, M. and DeLuca, H. F. 1984. Topographical and developmental studies on target sites of 1, 25-(OH)2 vit D3 in skin. Cell and Tissue Research 238: 489–496.Google Scholar
Stumpf, W. E., Perez-Delgado, M. M., Li, L., Bidman, H. J. and Tuohimaa, P. 1993. Vitamin D3 (soltriol) nuclear receptors in abdominal scent gland and skin of Siberian hamster (Phodopus snngorus) localised by autoradiography and immunohistochemistry. Histochemistry 100: 115–119.Google Scholar
Sullivan, M. and Nicholls, J. 1942. Nutritional dermatoes in the rat. V. Signs and symptoms resulting from a diet containing unheated dried egg white as the source of protein. Archives of Dermatology and Syphylology 45: 295–314.Google Scholar
Sundelin, J., Busch, C., Das, K., Das, S., Eriksson, U., Jonsson, K. H., Kampe, O., Laurent, B., Liljas, A., Newcomer, M., Nilsson, M., Norlinder, H., Rask, L., Ronne, H. and Paterson, P. A. 1983. Structure and tissue distribution of some retinoid-binding proteins. Journal of Investigative Dermatology 81: 59S–63S.Google Scholar
Tripathi, R. K., Hearing, V. J., Urabe, K., Aroca, P. and Spritz, R. A. 1992. Mutational mapping of the catalytic activities of human tyrosinase. Journal of Biological Chemistry 267: 23707–23712.Google Scholar
Underwood, E. J. and Somers, M. 1969. Studies of zinc nutrition in sheep. I. The relation of zinc to growth, testicular development and spermatogenesis in young rams. Australian Journal of Agricultural Research 20: 889–897.Google Scholar
Uno, H., Adachi, K., Allegra, E. and Montagna, W. 1968. Studies of common baldness of the stump tailed macaque. II. Enzyme activities of carbohydrate metabolism in the hair follicles. Journal of Investigative Dermatology 51:11–18.Google Scholar
Uno, H., Adachi, K. and Montagna, W. 1968. Glycogen contents of primate hair follicles. Journal of Investigative Dermatology 51:197–199.Google Scholar
Vasudev, A. S., Bagga, A., Shivaram, K. S. and Srivastava, R. N. 1989. Vitamin D-dependent rickets with alopecia. Indian Pediatrics 26:1254–1258.Google Scholar
Wallace, A. L. C. 1979. The effect of hormones on wool wool growth. In Physiological and environmental limitations to wool growth (ed. Black, J. L. and Reis, P. J.), pp. 257–268. University of New England Publishing Unit, Armidale NSW, Australia.Google Scholar
Weinstein, G.D. and Mooney, K.M. 1980. Cell proliferation kinetics in the human hair root. Journal of Investigative Dermatology 74: 43–46.Google Scholar
White, C. L. and Martin, G.B. 1988. Some pathological and productivity effects of zinc deficiency in the ram. Proceedings of the Nutrition Society of Australia 13: 86.Google Scholar
White, C., Martin, G.B., Hynd, P.I. and Chapman, R.E. 1994. The effect of zinc deficiency on wool growth and skin and wool follicle histology of male Merino lambs. British Journal of Nutrition 71: 425–435.Google Scholar
Whitehead, C.C. 1978. Effect of nutrient deficiencies in animals: biotin. In Effects of nutritional disorder in animals. (CRC Handbook Series in Nutrition and Food. Section E. Nutritional disorders. Vol. II. (ed. Reicheigl, M.), pp. 65–93. CRC Press, Florida.Google Scholar
Williams, R., Philpott, M.P. and Kealey, T. 1993. Metabolism of freshly-isolated human hair follicles capable of hair elongation; a glutaminolytic, aerobic glycolytic tissue. Journal of Investigative Dermatology 100: 834–840.Google Scholar
Wilson, N., Hynd, P.I. and Tivey, D. R. 1996. Amino acid transport into wool follicles. Proceedings of the Australian Society of Animal Production 21: 438.Google Scholar
Wilson, P. A. and the lateShort, B. F. 1979. Cell proliferation and cortical cell production in relation to wool growth. Australian Journal of Biological Sciences 32: 317–327.Google Scholar
Yoshikawa, K., Adachi, K., Levine, V. and Halprin, K. M. 1975. Microdetermination of cyclic AMP levels in human epidermis, dermis and hair follicles. British Journal of Dermatology 92: 241–248.Google Scholar
Zhong, Y.A., Zhang, M. H., Ke, X. D., Bai, Y. B., Xia, C. X., Wang, Y. R., Zhang, G. D., Yang, Z. Y. and Gaonigal, . 1990. Study on the relationships between sheep alopecia and zinc status in northern China. Scientia Agricultura Sinica 23: 9–14.Google Scholar