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Due to the low degree of differentiation between the two populations, the method based on genetic differentiation may not be able to identify different genes. A combination of several selection methods may be more conducive to this research, and the DCMS method allows more precisely and filters out spurious results specific to other methods (Ma et al., 2015). We calculated DCMS statistics for each population and the overlapping genes were selected as candidate genes associated with phenotypes (Figure 3A). In our study, a total of 71 overlapping genes were obtained using the DCMS method (Figure 3B). These overlapping genes were enriched into pathways involved in hair follicle development, including positive regulation of hair follicle development, positive regulation of cytokine-mediated signaling pathway, mitotic cell cycle, positive regulation of apoptotic process, negative regulation of transforming growth factor beta receptor signaling pathway, negative regulation of cell-substrate adhesion (Figure 3C). Classic studies showed that during embryogenesis, the embryonic epidermis and mesenchyme communicated with each other to form a hair follicle (McElwee and Hoffmann, 2000). The strong selection signal of DCMS found on Chromosome 20 (20:10300001-10350000) contained the CDH1 gene, which mediated the intercellular adhesion in the mammalian epidermis and hair follicles as the adhesive component of adherens junctions (Hodivala and Watt, 1994). CDH1 was weakly expressed in the dermis, while was highly expressed in the epidermis and hair follicles (She et al., 2016). Reports showed that CDH1 played an important role in the formation of melanin in hair follicles and the adhesion of hair follicles and epidermis (Larue et al., 1994; Perl et al., 1998; Young et al., 2003; Kuphal and Bosserhoff, 2012). Previous studies also found that continuous hair follicle cycling was dependent on CDH1 (Young et al., 2003). ACOXL, a typical lipid metabolism-related gene, was strongly selected in our study (Figure 3B). This enzyme could catalyze the desaturation of acyl-CoAs to 2-trans-enoyl-CoAs in the reductive half-reaction (Brown and Baker, 2003). Festa et al. (2011) found that dermal white adipose tissue (WAT) not only provided animals with thermo-insulation but also modulated regeneration dynamics of pelage hair follicles via the production of paracrine growth factor. Regeneration of the dermal WAT periodically cycles was in synchrony with the hair cycle, undergoing the cycles of expansion and collapse (Chase et al., 1953; Donati et al., 2014). These pieces of evidence suggested that the lipogenesis and lipolysis of WAT could be influenced by the β-oxidation process, so we inferred that ACOXL may affect the metabolism of WAT to synchrony affect the hair follicle cycle in yak. One circadian rhythm-related gene (MAGEL2) was identified among the overlapping genes (Figure 3B). MAGEL2 has been found to modulate the circadian rhythm: it was primarily expressed in the suprachiasmatic nucleus where the transcription of MAGEL2 oscillated in phase with clock-controlled genes. In addition to local paracrine modulators, hair follicles are also regulated by physiological changes that take place throughout the body. For example, several results suggested the involvement of the circadian clock regulate the hair cycle and hair follicle pigmentation (Al-Nuaimi et al., 2014; Hardman et al., 2015). In the mature anagen, clock genes were prominently expressed in the hair matrix, dermal papilla, and other follicular compartments (Plikus et al., 2013). Previous research showed that mice deficient in Magel2 expression will disrupt circadian rhythm, metabolic and endocrine deficits (Kozlov et al., 2007; Mercer et al., 2013). Therefore, MAGEL2 may affect the hair growth cycle by influencing the robust rhythmicity of MAGEL2 expression, which may be one of the reasons for the different hair lengths of Tianzhu white yak. PDPK1 was involved in the negative regulation of the transforming growth factor beta receptor (TGF-β) signaling pathway. During the development process of the hair follicle, TGF-β1, TGF-β2, and their receptors were locationally and cyclically specifically expressed in hair follicles and were proved to be involved in regulating the growth and development of hair follicle through multiple signaling pathways. Studies of transgene or gene knockout of TGF-β also confirmed that TGF-β related signaling was necessary for hair follicle development. It is indicated that PDPK1 may play an important role in hair development and cycle through TGF-β (Paus et al., 1997; Foitzik et al., 1999).

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