Hostname: page-component-78c5997874-dh8gc Total loading time: 0 Render date: 2024-11-09T07:50:29.652Z Has data issue: false hasContentIssue false

Fundamental hair follicle biology and fine fibre production in animals

Published online by Cambridge University Press:  24 February 2010

H. Galbraith*
Affiliation:
Institute of Biological and Environmental Sciences, University of Aberdeen, 23 St Machar Drive, Aberdeen, AB24 3RYUK and Department of Environmental and Natural Sciences, University of Camerino, Via Pontoni 5, 62032, Camerino, Italy
*
Get access

Abstract

Hair ‘fine’ fibre is an important commercial product of farmed and certain wild animal species. The fibre is produced in follicles embedded in skin. These have properties in common with other tissues of the integument and have importance in determining yield and quality of fibre. Means of understanding and improving these characteristics are informed by knowledge of integumental and follicle biology. This paper reviews contemporary information that identifies the major fibre-producing species and their production characteristic. It surveys knowledge describing fundamental biology of the integument and considers information derived for the hair follicle from studies on a number of species including genetically modified mice. It identifies the composition of the follicle and describes components and interrelationships between epidermal hair-fibre producing epidermis and fibroblast- and connective tissue-containing dermis. The structure of different primary and secondary follicle types, and associated structures, are described. Focus is given to the alterations in anatomy and in behaviour from active to inactive state, which occurs during the hair follicle cycle. Information is provided on the anatomical substructures (hair medulla, cortex, cuticles and supporting sheaths and dermal papilla), cellular and extracellular composition, and adhesion and chemical signalling systems, which regulate development from the early embryo to post-natal state and subsequent cycling. Such signalling involves the dermis and its specialist fibroblasts, which secrete signalling molecules, which along with those from local epidermis and systemic sources, largely determine structure and function of epidermal cells. Such chemical signalling typically includes endocrine-, paracrine-, autocrine- and juxtacrine-acting molecules and interactions with their receptors located on cell membranes or intracellularly with transduction of message mediated by transcription factors at gene level. Important hormones and growth factors and inhibitors regulating morphogenic and/or mitogenic activity are identified. These mediate mechanisms associated with presence or absence in skin and development of patterning for primary or secondary follicles. Reference is made to deposition of individual keratins and keratin-associated proteins in follicle sub-structures and to fibre properties such as length, diameter, medullation, crimp and lustre. Pre- and post-natal regulation of pigmentation by melanocytes is reviewed. Brief attention is given to genomic and non-genomic variation and impact on the phenotypes expressed and the role of regulatory gene products as potential molecular markers for selection of superior animals. The importance of nutrients in providing substrates for follicular structures and enzymes and in molecules facilitating gene expression is also considered.

Type
Full Paper
Copyright
Copyright © The Animal Consortium 2010

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

Agricultural and Food Research Council (AFRC) 1993. Agricultural and Food Research Council. Energy and protein requirements of ruminants. CAB International, Wallingford, UK.Google Scholar
Allain, D, Renieri, C 2010. Genetics of fibre production and fleece characteristics in small ruminants, Angora rabbit and South American camelids. Animal 4, 14721481.CrossRefGoogle ScholarPubMed
Allain, D, Thebault, RG, Rougeot, J, Martinet, L 1994. Biology of fibre growth in mammals producing fine fibre and fur in relation to control by day length: relationship with other seasonal functions. In Hormonal control of fibre growth and shedding (ed. JP Laker and D Allain), Occasional publication no 2, pp. 2340. European Fine Fibre Network, Aberdeen, Scotland, UK.Google Scholar
Antonini, M 2010. Hair follicle characteristics and fibre production in South American camelids. Animal 4, 14601471.CrossRefGoogle ScholarPubMed
Barrett, P, Ebling, FJP, Schuhler, S, Wilson, D, Ross, AW, Warner, A, Jethwa, P, Boelen, A, Visser, TJ, Ozanne, DM, Archer, ZA, Mercer, JG, Morgan, PJ 2007. Hypothalamic thyroid hormone catabolism acts as a gatekeeper for the seasonal control of body weight and reproduction. Endocrinology 148, 36083617.CrossRefGoogle ScholarPubMed
Barsh, GS 2006. Regulation of pigment type switching by agouti, melanocortin signalling, attractin and mahoganoid. In The Pigmentary System, 2nd Edition (ed. JJ Nordlund, RE Boissy, VJ Hearing, RA King, WS Oetting and J-P Ortonne), pp. 395409. Blackwell Publishing Ltd, Oxford, UK.CrossRefGoogle Scholar
Bawden, CS, Powell, BC, Walker, SK, Rogers, GE 1998. Expression of a wool intermediate filament keratin transgene in sheep fibre alters structure. Transgenic Research 7, 273287.CrossRefGoogle ScholarPubMed
Botchkarev, VA, Botchkareva, NV, Nakamura, M, Huber, O, Funa, K, Lauster, R, Paus, R, Gilchrest, BA 2001. Noggin is required for induction of the hair follicle growth phase in postnatal skin. FASEB Journal 15, 22052214.CrossRefGoogle ScholarPubMed
Brancaccio, A, Minichiello, A, Grachtchouk, M, Antonini, D, Sheng, H, Parlato, N, Dathan, R, Dlugosz, AA, Missero, C 2004. Requirement of the forkhead gene Foxe1, a target of sonic hedgehog signaling, in hair follicle morphogenesis. Human Molecular Genetics 13, 25952606.CrossRefGoogle ScholarPubMed
Bull, JJ, Pelengaris, S, Hendrix, S, Chronnell, CMT, Khan, M, Philpott, MP 2005. Ectopic expression of c-Myc in the skin affects the hair growth cycle and causes an enlargement of the sebaceous gland. British Journal of Dermatology 152, 11251133.CrossRefGoogle ScholarPubMed
Candille, SI, Van Raamsdonk, CD, Chen, C, Kuijper, S, Chen-Tsai, Y, Russ, A, Meijlink, F, Barsh, GS 2004. Dorsoventral patterning of the mouse coat by Tbx15. PLoS Biology 2, e3.CrossRefGoogle ScholarPubMed
Caneparo, L, Huang, YL, Staudt, N, Tada, M, Ahrendt, R, Kazanskaya, O, Niehrs, C, Houart, C 2007. Dickkopf-1 regulates gastrulation movements by coordinated modulation of Wnt/βcatenin and Wnt/PCP activities, through interaction with the Dally-like homolog Knypek. Genes and Development 21, 465480.CrossRefGoogle Scholar
Chang, HY 2007. Patterning skin pigmentation via Dickkopf. Journal of Investigative Dermatology 127, 994995.CrossRefGoogle ScholarPubMed
Chintala, S, Li, W, Lamoreux, ML, Ito, S, Sviderskaya, EV, Bennett, DC, Park, Y-M, Gahl, WA, Huizing, M, Spritz, RA, Ben, S, Novak, EK, Tan, J, Swank, RT 2005. Slc7a11 gene controls production of pheomelanin pigment and proliferation of cultured cells. Proceedings of the National Academy of Sciences of the United States of America 102, 1096410969.CrossRefGoogle ScholarPubMed
Choy, VJ, Nixon, AJ, Pearson, AJ 1995. Localisation of receptors of prolactin in ovine skin. Journal of Endocrinology 144, 143151.CrossRefGoogle ScholarPubMed
Craven, AJ, Nixon, AJ, Ashby, MG, Ormandy, CJ, Blazek, K, Wilkins, RJ, Pearson, AJ 2006. Prolactin delays hair regrowth in mice. Journal of Endocrinology 191, 415425.CrossRefGoogle ScholarPubMed
Council for Scientific and Industrial Research (CSIR) 2007. CSIR eNews. CSIR to detect mohair sheen. Retrieved January 4, 2010, from http://www.csir.co.za/enews/2007_oct/msm_04.html.Google Scholar
Danks, DM 1991. Copper deficiency and the skin. In Physiology, Biochemistry and Molecular Biology of the Skin (ed. LA Goldsmith), pp. 13511361. Oxford University Press, New York, USA.Google Scholar
Dryer, JH, Marincowitz, G 1967. Some observation on the skin histology and fibre characteristics of the Angora goat (Capra hircus angoraensisi). South African Journal of Agricultural Science 10, 477500.Google Scholar
Ebling, FJG, Hale, PA, Randall, VA 1991. Hormones and hair growth. In Physiology, Biochemistry and Molecular Biology of the Skin (ed. LA Goldsmith), pp. 660696. Oxford University Press, New York, USA.Google Scholar
Ellis, T, Gambardella, L, Horcher, M, Tschanz, S, Capol, J, Bertram, P, Jochum, W, Barrandon, Y, Busslinger, M 2001. The transcriptional repressor CDP (Cutl1) is essential for epithelial cell differentiation of the lung and the hair follicle. Genes and Development 15, 23072319.CrossRefGoogle ScholarPubMed
Enshell-Seijffers, D, Lindon, C, Morgan, BA 2008. The serine protease Corin is a novel modifier of the agouti pathway. Development 135, 217225.CrossRefGoogle ScholarPubMed
Fessing, MY, Sharova, TY, Sharov, AA, Atoyan, R, Botchkarev, VA 2006. Involvement of the Edar signaling in the control of hair follicle involution (catagen). American Journal of Pathology 169, 20752084.CrossRefGoogle ScholarPubMed
Fischer, TW, Slominski, A, Tobin, DJ, Paus, R 2008. Melatonin and the hair follicle. Journal of Pineal Research 44, 115.CrossRefGoogle ScholarPubMed
Fleet, MR, Forrest, JW, Walker, SK, Rogers, GE 2004. Foetal development of melanocyte populations in Merino wool-bearing skin. Wool Technology and Sheep Breeding 52, 101123.Google Scholar
Fuchs, E, Marchuk, D 1983. Type I and type II keratins have evolved from lower eukaryotes to form the epidermal intermediate filaments in mammalian skin. Proceedings of the National Academy of Sciences of the United States of America 80, 58575861.CrossRefGoogle ScholarPubMed
Galbraith, H 1998. Nutritional and hormonal regulation of hair follicle growth and development. Proceedings of the Nutrition Society 57, 195205.CrossRefGoogle ScholarPubMed
Galbraith, H 2000. Protein and sulphur amino acid nutrition of hair fibre-producing Angora and Cashmere goats. Livestock Production Science 64, 8193.CrossRefGoogle Scholar
Galbraith, H 2006. A current perspective on the biology of fibre production in animals. In South American camelids research, vol. 1, (ed. M Gerken and C Renieri), pp. 195207. Wageningen Academic Publishers, Wageningen, The Netherlands.Google Scholar
Galbraith, H 2010. In vitro methodology, hormonal and nutritional effects and fibre production in isolated ovine and caprine anagen hair follicles. Animal 4, 14821489.CrossRefGoogle ScholarPubMed
Galbraith, H, Flannigan, S, Swan, L, Cash, P 2006. Proteomic evaluation of tissues at functionally important sites in the bovine claw. Cattle Practice 14, 127137.Google Scholar
Galbraith, H, Norton, B, Sahlu, T 2000. Recent advances in the nutritional biology of Angora and Cashmere goats. In Proceedings of the 7th International Symposium on Goats (ed. L Gruner and P Chabert), 15–21 May 2000, pp. 5965, Tours, France.Google Scholar
Galbraith, H, Scaife, JR 2008. Lameness in dairy cows: influence of nutrition on claw composition and health. In Recent advances in animal nutrition (ed. PC Garnsworthy and J Wiseman), pp. 91126. Nottingham University Press, Nottingham, UK.Google Scholar
Gerken, M 2010. Relationships between integumental characteristics and thermoregulation in South American camelids. Animal 4, 14511459.CrossRefGoogle ScholarPubMed
Gleason, BC, Crum, CP, Murphy, GF 2008. Expression patterns of MITF during human cutaneous embryogenesis: evidence for bulge epithelial expression and persistence of dermal melanoblasts. Journal of Cutaneous Pathology 35, 615622.CrossRefGoogle ScholarPubMed
Godfrey, KM 1998. Maternal regulation of fetal development and health in adult life. European Journal of Obstetrics & Gynecology and Reproductive Biology 78, 141150.CrossRefGoogle ScholarPubMed
Goldsmith, LA 1991. Physiology, biochemistry and molecular biology of the skin. Oxford University Press, Oxford, UK.Google Scholar
Gratten, J, Wilson, AJ, McRae, AF, Beraldi, D, Visscher, PM, Pemberton, JM, Slate, J 2008. A localized negative genetic correlation constrains microevolution of coat color in wild sheep. Science 319, 318320.CrossRefGoogle ScholarPubMed
Hammerschmidt, B, Schlake, T 2007. Localization of Shh expression by Wnt and Eda affects axial polarity and shape of hairs. Developmental Biology 305, 246261.CrossRefGoogle ScholarPubMed
Hanon, EA, Lincoln, GA, Fustin, JM, Dardente, H, Masson-Pevet, M, Morgan, PJ, Hazlerigg, DG 2008. Ancestral TSH mechanism signals summer in a photoperiodic mammal. Current Biology 18, 11471152.CrossRefGoogle Scholar
He, L, Eldridge, AG, Jackson, PK, Gunn, TM, Barsh, GS 2003. Accessory proteins for melanocortin signaling – attractin and mahogunin. Annals of the New York Academy of Sciences 994, 288298.CrossRefGoogle ScholarPubMed
Hébert, JM, Rosenquist, T, Götz, J, Martin, GR 1994. FGF5 as a regulator of the hair growth cycle: evidence from targeted and spontaneous mutations. Cell 78, 10171025.CrossRefGoogle ScholarPubMed
Hepburn, NL, Kinnimonth, L, Galbraith, H 2007. Pigmentation, impression hardness and the presence of melanosomes in bovine claw tissue. Journal of Agricultural Science 145, 283290.CrossRefGoogle Scholar
Hepburn, NL, Knight, CH, Wilde, CJ, Hendry, KAK, Galbraith, H 2008. L-methionine uptake, incorporation and effects on proliferative activity and protein synthesis in bovine claw tissue explants in vitro. Journal of Agricultural Science 146, 103115.CrossRefGoogle Scholar
Hynd, PI 1989. Effects of nutrition on wool follicle cell kinetics in sheep differing in efficiency of wool production. Australian Journal of Agricultural Research 40, 409417.CrossRefGoogle Scholar
Hynd, PI 2000. The nutritional biochemistry of wool and hair follicles. Animal Science 70, 181195.CrossRefGoogle Scholar
Ibraheem, M, Galbraith, H, Scaife, JR, Ewen, S 1994. Growth of secondary hair follicles of the cashmere goat in vitro and their response to prolactin and melatonin. Journal of Anatomy 185, 135142.Google ScholarPubMed
Inoue, K, Aoi, N, Yoshimura, K 2009. TGF-β2 is expressed in human dermal papilla cells and modulates folliculogenesis. Journal of Investigative Dermatology 129, S1S103, Abstract 602.Google Scholar
Ito, M, Yang, Z, Andl, T, Cui, C, Kim, N, Millar, SE, Cotsarelis, G 2007. Wnt-dependent de novo hair follicle regeneration in adult mouse skin after wounding. Nature 447, 316320.CrossRefGoogle ScholarPubMed
Jahoda, CA, Reynolds, AJ 2001. Hair follicle dermal sheath cells: unsung participants in wound healing. Lancet 358, 14451448.CrossRefGoogle ScholarPubMed
Jamora, C, Lee, P, Kocieniewski, P, Azhar, M, Hosokawa, R, Chai, Y, Fuchs, E 2005. A Signaling pathway involving TGF-β2 and Snail in hair follicle morphogenesis. PLoS Biology 3, e11.CrossRefGoogle ScholarPubMed
Kaufman, CK, Zhou, P, Pasolli, HA, Rendl, M, Bolotin, D, Lim, K-C, Dai, X, Alegre, M-L, Fuchs, E 2003. GATA-3: an unexpected regulator of cell lineage determination in skin. Genes and Development 17, 21082122.CrossRefGoogle ScholarPubMed
Kauser, S, Slominski, A, Wei, ET, Tobin, DJ 2006. Modulation of the human hair follicle pigmentary unit by corticotropin-releasing hormone and urocortin peptides. The FASEB Journal 20, 882895.CrossRefGoogle ScholarPubMed
Kawano, M, Komi-Kuramochi, A, Asada, M, Suzuki, M, Oki, J, Jiang, J, Imamura, T 2005. Comprehensive analysis of FGF and FGFR expression in skin: FGF18 is highly expressed in hair follicles and capable of inducing anagen from telogen stage hair follicles. Journal of Investigative Dermatology 124, 877885.CrossRefGoogle ScholarPubMed
Koch, PJ, Mahoney, MG, Ishikawa, H, Pulkkinen, L, Uitto, J, Shultz, L, Murphy, GF, Whitaker-Menezes, D, Stanley, JR 1997. Targeted disruption of the pemphigus vulgaris antigen (desmoglein 3) gene in mice causes loss of keratinocyte cell adhesion with a phenotype similar to pemphigus vulgaris. Journal of Cell Biology 137, 10911102.CrossRefGoogle ScholarPubMed
Kulessa, H, Turk, G, Hogan, BL 2000. Inhibition of Bmp signaling affects growth and differentiation in the anagen hair follicle. EMBO Journal 19, 66646674.CrossRefGoogle ScholarPubMed
Li, SW, Ouyang, HS, Rogers, GE, Bawden, CS 2009. Characterization of the structural and molecular defects in fibres and follicles of the merino felting lustre mutant. Experimental Dermatology 18, 134142.CrossRefGoogle ScholarPubMed
Limat, A, Breitkreutz, D, Stark, HJ, Hunziker, T, Thikoetter, G, Noser, F, Fusenig, NE 1991. Experimental modulation of the differentiated phenotype of keratinocytes from epidermis and hair follicle outer root sheath and matrix cells. Annals of the New York Academy of Sciences 642, 125146.CrossRefGoogle ScholarPubMed
Lin, KK, Geyfman, M, Kumar, V, Chudova, D, Ihler, AT, Smyth, P, Paus, R, Takahashi, JS, Andersen, B 2009. Circadian clock genes play a role in regulation of timing of anagen progression in mouse hair follicles. Journal of Investigative Dermatology 129, S1S103, Abstract 603.Google Scholar
Lincoln, GA 2002. Neuroendocrine regulation of seasonal gonadotrophin and prolactin rhythms: lessons from the Soay ram model. Reproduction Supplement 59, 131147.Google ScholarPubMed
Lincoln, GA, Baker, BI 1995. Seasonal and photoperiod-induced changes in the secretion of α-melanocyte-stimulating hormone in Soay sheep: temporal relationships with changes in β-endorphin, prolactin, follicle-stimulating hormone, activity of the gonads and growth of wool and horns. Journal of Endocrinology 144, 471481.CrossRefGoogle ScholarPubMed
Lincoln, GA, Andersson, H, Loudon, A 2003. Clock genes in calendar cells as the basis of annual timekeeping in mammals – a unifying hypothesis. Journal of Endocrinology 179, 113.CrossRefGoogle ScholarPubMed
Lodish, H, Berk, A, Zipursky, SL, Matsudaira, P, Baltimore, D, Darnell, JE 2000. Molecular cell biology, 4th edition. WH Freeman and Company, New York, USA.Google Scholar
Mak, KKL, Chan, SY 2003. Epidermal growth factor as a biologic switch in hair growth cycle. Journal of Biological Chemistry 278, 2612026126.CrossRefGoogle ScholarPubMed
Malgouries, S, Thibaut, S, Bernard, BA 2008. Proteoglycan expression patterns in human hair follicle. British Journal of Dermatology 158, 234342.CrossRefGoogle ScholarPubMed
Mastorakos, G, Zapanti, E 2004. The hypothalamic-pituitary-adrenal axis in the neuroendocrine regulation of food intake and obesity: the role of corticotropin releasing hormone. Nutritional Neuroscience 7, 271280.CrossRefGoogle ScholarPubMed
Mathers, JC 2008. Epigenomics: a basis for understanding individual differences. Proceedings of the Nutrition Society 67, 390394.CrossRefGoogle ScholarPubMed
McLaren, RJ, Rogers, GR, Davies, KP, Maddox, JF, Montgomery, GW 1997. Linkage mapping of wool keratin and keratin-associated protein genes in sheep. Mammalian Genome 8, 938940.CrossRefGoogle ScholarPubMed
Messenger, AG 1993. The control of hair growth: an overview. Journal of Investigative Dermatology 101 (Suppl. 1), 4S9S.CrossRefGoogle ScholarPubMed
Millar, SE 2002. Molecular mechanisms regulating hair follicle development. Journal of Investigative Dermatology 118, 216225.CrossRefGoogle ScholarPubMed
Mohamed, OA, Dufort, D, Clarke, HJ 2004. Expression and estradiol regulation of Wnt genes in the mouse blastocyst identify a candidate pathway for embryo-maternal signaling at implantation. Biology of Reproduction 71, 417424.CrossRefGoogle ScholarPubMed
Moore, GPM, Panaretto, BA, Carter, NB 1985. Epidermal hyperplasia and wool follicle regression in sheep infused with epidermal growth-factor. Journal of Investigative Dermatology 84, 172175.Google ScholarPubMed
Mou, C, Jackson, B, Schneider, P, Overbeek, PA, Headon, DJ 2006. Generation of the primary hair follicle pattern. Proceedings of the National Academy of Sciences of the United States of America 103, 90759080.CrossRefGoogle ScholarPubMed
Naito, A, Sato, T, Matsumoto, T, Takeyama, K, Yoshino, T, Kato, S, Ohdera, M 2008. Dihydrotestosterone inhibits murine hair growth via the androgen receptor. British Journal of Dermatology 159, 300305.CrossRefGoogle ScholarPubMed
Nagorcka, BN, Mooney, JR 1989. The reaction diffusion system as a spatial organizer during initiation and development of hair follicles and formation of fibre. In The biology of wool and hair (ed. GE Rogers, PJ Reis, KA Ward and RC Marshall), pp. 365379. Chapman and Hall, London, UK.Google Scholar
Neldner, KH 1991. The biochemistry and physiology of zinc metabolism. In Physiology, biochemistry and molecular biology of the skin (ed. LA Goldsmith), pp. 13291350. Oxford University Press, New York, USA.Google Scholar
Nilsen, TW 2007. Mechanisms of microRNA-mediated gene regulation in animal cells. Trends in Genetics 23, 243249.CrossRefGoogle ScholarPubMed
Nixon, AJ, Ford, CA, Wildermoth, JE, Craven, AJ, Ashby, MG, Pearson, AJ 2002. Regulation of prolactin receptor expression in ovine skin in relation to circulating prolactin and wool follicle growth status. Journal of Endocrinology 172, 605614.CrossRefGoogle ScholarPubMed
Nordlund, JJ, Boissy, RE, Hearing, VJ, King, RA, Oetting, WS, Ortonne, J-P 2006. The pigmentary system, 2nd Edition. Blackwell Publishing Ltd, Oxford, UK.CrossRefGoogle Scholar
Norris, BJ, Whan, VA 2008. A gene duplication affecting expression of the ovine ASIP gene is responsible for white and black sheep. Genome Research 18, 12821293.CrossRefGoogle ScholarPubMed
Odland, GF 1991. Structure of the skin. In Physiology, biochemistry and molecular biology of the skin (ed. LA Goldsmith), pp. 362. Oxford University Press, New York, USA.Google Scholar
Ohnemus, U, Uenalan, M, Inzunza, J, Gustafsson, JA, Paus, R 2006. The hair follicle as an estrogen target and source. Endocrine Reviews 27, 677706.CrossRefGoogle ScholarPubMed
O’Neill, C 2008. Phosphatidylinositol 3-kinase signaling in mammalian preimplantation embryo development. Reproduction 136, 147156.CrossRefGoogle ScholarPubMed
Orwin, DFG 1989. Variations in wool follicle morphology. In The biology of wool and hair (ed. GE Rogers, PJ Reis, KA Ward and RC Marshall), pp. 227242. Chapman and Hall, London, UK.Google Scholar
Ouji, Y, Yoshikawa, M, Moriya, K, Nishiofuku, M, Matsuda, R, Ishizaka, S 2008. Wnt-10b, uniquely among Wnts, promotes epithelial differentiation and shaft growth. Biochemical and Biophysical Research Communications 367, 299304.CrossRefGoogle ScholarPubMed
Philpott M, P, Kealey, T 1994. Effects of EGF on the morphology and patterns of DNA synthesis in isolated human hair follicles. Journal of Investigative Dermatology 102, 186191.CrossRefGoogle ScholarPubMed
Plowman, JE 2003. Review. Proteomic database of wool components. Journal of Chromatography. B, Analytical Technologies in the Biomedical and Life Sciences 787, 6376.CrossRefGoogle ScholarPubMed
Potter, CS, Peterson, RL, Barth, JL, Pruett, ND, Jacobs, DF, Kern, MJ, Argraves, WS, Sundberg, JP, Awgulewitsch, A 2006. Evidence that the satin hair mutant gene Foxq1 is among multiple and functionally diverse regulatory targets for Hoxc13 during hair follicle differentiation. Journal of Biological Chemistry 281, 2924529255.CrossRefGoogle ScholarPubMed
Powell, BC 1997. Molecular genetics of sheep. In The genetics of sheep (ed. LR Piper and A Ruvinsky), pp. 149181. CAB International, Wallingford, UK.Google Scholar
Pruche, F, Boyera, N, Bernard, BA 1996. K+ (ATP) channel openers inhibit the bradykinin-induced increase of intracellular calcium in hair follicle outer root sheath keratinocytes. In Hair research for the next millennium (ed. D Van Neste and VA Randall), pp. 463466. Elsevier, Amsterdam, The Netherlands.Google Scholar
Purvis, IW, Jeffery, N 2007. Genetics of fibre production in sheep and goats. Small Ruminant Research 70, 4247.CrossRefGoogle Scholar
Quevedo, WC, Holstein, TJ 2006. General biology of mammalian pigmentation 2006. In The pigmentary system, 2nd edition (ed. JJ Nordlund, RE Boissy, VJ Hearing, RA King, WS Oetting and J-P Ortonne), pp. 6390. Blackwell Publishing Ltd, Oxford, UK.Google Scholar
Rafik, ME, Doucet, J, Briki, F 2004. The intermediate filament architecture as determined by X-ray diffraction modeling of hard α-keratin. Biophysical Journal 86, 38933904.CrossRefGoogle Scholar
Renieri, C, Valbonesi, A, La Manna, V, Antonini, M, Lauvergne, JJ 2008. Inheritance of coat colour in Merino sheep. Small Ruminant Research 74, 2329.CrossRefGoogle Scholar
Rhind, SM, Kyle, CE 2004. Skin deiodinase profiles and associated patterns of hair follicle activity in cashmere goats of contrasting genotypes. Australian Journal of Agricultural Research 55, 443448.CrossRefGoogle Scholar
Rogers, GE 2006. Biology of the wool follicle: an excursion into a unique tissue interaction system waiting to be re-discovered. Experimental Dermatology 15, 931949.CrossRefGoogle ScholarPubMed
Rogers, MA, Schweizer, J 2005. Human KAP genes, only the half of it? Extensive size polymorphisms in hair keratin-associated protein genes. Journal of Investigative Dermatology 124, viiix.CrossRefGoogle ScholarPubMed
Schlake, T 2001. The nude gene and the skin. Experimental Dermatology 10, 293304.CrossRefGoogle ScholarPubMed
Schlake, T 2005. IGF signals specifically regulate the structure of hair shaft medulla via IGF-binding protein 5. Development (Cambridge, England) 132, 29812990.CrossRefGoogle ScholarPubMed
Schlake, T 2007. Determination of hair structure and shape. Seminars in Cell and Developmental Biology 18, 267273.CrossRefGoogle ScholarPubMed
Schmidt-Ullrich, R, Paus, R 2005. Molecular principles of hair follicle induction and morphogenesis. BioEssays 27, 247261.CrossRefGoogle ScholarPubMed
Schneider, MR, Schmidt-Ullrich, R, Paus, R 2009. The hair follicle as a dynamic miniorgan. Current Biology 19, R132R142.CrossRefGoogle ScholarPubMed
Schneider, MR, Werner, S, Paus, R, Wolf, E 2008. Beyond wavy hairs. The epidermal growth factor receptor and its ligands in skin biology and pathology. American Journal of Pathology 173, 1424.CrossRefGoogle Scholar
Schweizer, J, Bowden, PE, Coulombe, PA, Langbein, L, Lane, EB, Magin, TM, Maltais, L, Omary, MB, Parry, DAD, Rogers, MA, Wright, MW 2006. New consensus nomenclature for mammalian keratins. Journal of Cell Biology 174, 169174.CrossRefGoogle ScholarPubMed
Seto, ES, Bellen, HJ, Lloyd, TE 2002. When cell biology meets development: endocytic regulation of signaling pathways. Genes and Development 16, 13141336.CrossRefGoogle ScholarPubMed
Sherertz, EF, Goldsmith, LA 1991. Nutritional influence on the skin. In Physiology, biochemistry and molecular biology of the skin (ed. LA Goldsmith) pp. 13151328. Oxford University Press, New York, USA.Google Scholar
Slominski, A, Wortsman, J, Plonka, PM, Schallreuter, KU, Paus, R, Tobin, DJ 2005. Hair follicle pigmentation. Journal of Investigative Dermatology 124, 1321.CrossRefGoogle ScholarPubMed
Smith, SB, Zhou, BK, Orlow, SJ 1998. Expression of tyrosinase and the tyrosinase related proteins in the Mitfvit (vitiligo) mouse eye: implications for the function of the microphthalmia transcription factor (Mitf). Experimental Eye Research 66, 403410.CrossRefGoogle ScholarPubMed
Souri, M, Galbraith, H, Scaife, JR 1997. Response of isolated Mohair secondary follicles to epidermal growth factor in vitro. Proceedings of the Nutrition Society 56, 181A.Google Scholar
Stenn, KS, Paus, R 2001. Controls of hair follicle cycling. Physiological Reviews 81, 449494.CrossRefGoogle ScholarPubMed
Tahmasbi, M, Galbraith, H, Scaife, JR 2007a. The effect of biotin deficiency in the pre-ruminant and immediately post-ruminant Angora and Cashmere kids. Journal of Animal and Veterinary Advances 6, 539548.Google Scholar
Tahmasbi, M, Galbraith, H, Scaife, JR 2007b. Investigation of the role of biotin in the regulation of wool growth in sheep hair follicles cultured in vitro. Research Journal of Animal Sciences 1, 919.Google Scholar
Thisse, B, Thisse, C 2005. Functions and regulations of fibroblast growth factor signalling during embryonic development. Developmental Biology 287, 390402.CrossRefGoogle ScholarPubMed
Thomas, N, Tivey, DR, Penno, NM, Nattrass, G, Hynd, PI 2007. Characterization of transport systems for cysteine, lysine, alanine, and leucine in wool follicles of sheep. Journal of Animal Science 85, 22052213.CrossRefGoogle ScholarPubMed
Thompson, ACT, Hebart, ML, Penno, NM, Hynd, PI 2007. Perinatal wool follicle attrition coincides with elevated perinatal circulating cortisol concentration in Merino sheep. Australian Journal of Agricultural Research 58, 748752.CrossRefGoogle Scholar
Tiede, S, Klatte, JE, Wenzel, B, Lynaugh, K, Paus, R 2009. Endocrine modulation of human hair follicle epithelial progenitor cells in situ and in vitro: effects of thyroid hormones, TSH, and calcitriol. Journal of Investigative Dermatology 129, S102S102, Abstract 610.Google Scholar
Van Beek, N, Bodó, E, Kromminga, A, Gáspár, E, Meyer, K, Zmijewski, MA, Slominski, A, Wenzel, BE, Paus, R 2008. Thyroid hormones directly alter human hair follicle functions: anagen prolongation and stimulation of both hair matrix keratinocyte proliferation and hair pigmentation. The Journal of Clinical Endocrinology and Metabolism 93, 43814388.CrossRefGoogle ScholarPubMed
Vassis, P, Butcher, EC, Lee, AR, Holt, LA 2003. Reflection from natural fibres: determination of the scale angle profile. Journal of Materials Science 38, 45414549.CrossRefGoogle Scholar
Vidal, VPI, Chaboissier, MC, Lützkendorf, S, Cotsarellis, G, Mill, P, Hui, C-C, Ortonne, N, Ortonne, J-P, Schedl, A 2005. Sox9 is essential for outer root sheath differentiation and the formation of the hair stem cell compartment. Current Biology 15, 13401351.CrossRefGoogle ScholarPubMed
Weger, N, Schlake, T 2005. Igf-I signalling controls the hair growth cycle and the differentiation of hair shafts. Journal of Investigative Dermatology 125, 873882.CrossRefGoogle ScholarPubMed
Wenguang, Z, Jianghong, W, Jinquan, L, Yashizawa, M 2007. A subset of skin-expressed microRNAs with possible roles in goat and sheep hair growth based on expression profiling of mammalian microRNAs. OMICS: A Journal of Integrative Biology 11, 385396.CrossRefGoogle ScholarPubMed
Wu, D-D, Irwin, DM, Zhang, Y-P 2008. Molecular evolution of the keratin associated protein gene family in mammals, role in the evolution of mammalian hair. BMC Evolutionary Biology 8, 241.CrossRefGoogle ScholarPubMed
Yamaguchi, Y, Passeron, T, Watabe, H, Yasumoto, K, Rouzaud, F, Hoashi, T, Hearing, VJ 2007. The effects of dickkopf 1 on gene expression and Wnt signaling by melanocytes: mechanisms underlying its suppression of melanocyte function and proliferation. Journal of Investigative Dermatology 127, 12171225.CrossRefGoogle ScholarPubMed
Yi, R, O’Carroll, D, Pasolli, HA, Zhang, Z, Dietrich, FS, Tarakhovsky, A, Fuchs, E 2006. Morphogenesis in skin is governed by discrete sets of differentially expressed microRNAs. Nature Genetics 38, 356362.CrossRefGoogle ScholarPubMed
Zanello, SB, Jackson, DM, Holick, MF 2000. Expression of the circadian clock genes clock and Period1 in human skin. Journal of Investigative Dermatology 115, 757760.CrossRefGoogle ScholarPubMed