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Living at a high altitude involves many environmental challenges. The combined effects of hypoxia and cold stress impose severe physiological challenges on endothermic animals. The yak is integral to the livelihood of the people occupying the vast, inhospitable Qinghai–Tibetan plateau and the surrounding mountainous region. Due to long-term selection, the yak exhibits stable and unique genetic characteristics which enable physiological, biochemical, and morphological adaptations to a high altitude. Thus, the yak is a representative model for mammalian plateau-adaptability studies. Understanding coping mechanisms provides unique insights into adaptive evolution, thus informing the breeding of domestic yaks. This review provides an overview of genetic adaptations in Bos grunniens to high-altitude environmental stress. Combined genomics and theoretical advances have informed the genetic basis of high-altitude adaptations.

Yak has been subject to natural selection, human domestication and interspecific introgression during its evolution. However, genetic variants favored by each of these processes have not been distinguished previously. We constructed a graph-genome for 47 genomes of 7 cross-fertile bovine species. This allowed detection of 57,432 high-resolution structural variants (SVs) within and across the species, which were genotyped in 386 individuals. We distinguished the evolutionary origins of diverse SVs in domestic yaks by phylogenetic analyses. We further identified 334 genes overlapping with SVs in domestic yaks that bore potential signals of selection from wild yaks, plus an additional 686 genes introgressed from cattle. Nearly 90% of the domestic yaks were introgressed by cattle. Introgression of an SV spanning the KIT gene triggered the breeding of white domestic yaks. We validated a significant association of the selected stratified SVs with gene expression, which contributes to phenotypic variations. Our results highlight that SVs of different origins contribute to the phenotypic diversity of domestic yaks. Yaks have been subject to natural selection, human domestication and interspecific introgression during their evolution. Here, the authors have identified genomic structural variations and the linked genes involved in these processes in domestic yaks, to reveal new insight into genetic basis of phenotypic diversity.

Background and aim Mitochondrial genome has aseries of characteristics such as simple structure, no recombination, maternalinheritance, stable structure, fast evolution rate, and high copy number. Moreover, it is easy to be sequenced,contains high-resolution phylogenetic information, and exists in a wide rangeof taxa. Therefore, it is widely used in the study of biological phylogeny. Atpresent, phylogenetic studies focus mainly on D-loop region, cytochrome b gene,and protein-coding sequence. Phylogenetic studies using the mitochondrialcomplete sequence are rarely reported in yak. Therefore, the present studyaimed to construct phylogenetic tree using yak mitochondrial complete sequenceand compare the subsequent results with previous findings obtained usingpartial sequences. Results Complete mitochondrial sequences of five yakpopulations from Qinghai and Xinjiang were obtained. The mitotype diversity ofthe five populations was Xueduo yak (0.992 ± 0.015), Pamir yak (0.990 ± 0.014),Yushu yak (0.963 ± 0.033), Qilian yak (0.948 ± 0.036), and Huanhu yak (0.905 ±0.048), which showed a higher mitotype diversity compared with other breeds fromthe previous reports, including Jiulong yak, Maiwa yak, Zhongdian yak, andTianzhu yak. A total of 78 mitotypes were obtained from 111 individuals. Amongthese, Yushu yak, Huanhu yak, Xueduo yak, and Qilian yak all shared mitotypes,but the Pamir yak did not share mitotypes with these four populations.Phylogenetic analysis showed that yak populations were separable into threedistinct branches. The analysis identified a new phylogenetic branch containingboth wild and domestic yaks. The 155 mitotypes found in 206 individuals weredivided into 3 haplogroups by mitotype clustering. Thehaplogroup was not associated with the geographical distribution of yaks. Theyaks in the same population or the same ecological environment were distributedin different haplogroups. Among the threehaplogroups, haplogroup A and haplogroup B showed a star-shaped distribution ofmitotypes. The central mitotypes were widely distributed and had a highfrequency. Conclusions Thegenetic diversity of yaks in Qinghai was high. Both domestic and wild yaks clusteredinto three branches.

Background Cattle have evolved genetic adaptations to a diverse range of agroecological zones, such as plateaus and arid zones. However, little is known about its genetic basis of adaptation to harsh environments within a short period of time after domestication. Here, we analyzed whole-genome sequence data from three indigenous cattle breeds (Anxi, Qaidam and Zhangmu) in northwest China and five worldwide cattle breeds (Angus, Holstein, Jersey, Gir and N’Dama) to explore their genetic composition and identify selective sweeps in the Chinese cattle breeds. Results Analyses of phylogenetic and population structure revealed that three indigenous cattle breeds share genomic components from Bos taurus and Bos indicus . A novel set of candidate genes was identified through comparative genomic analyses of cattle from contrasting environments based on SNP and copy number variation (CNV) data. These candidate genes are potentially associated with adaptive phenotypes, including high-altitude adaptability (e.g., ANGPT1, PPARGC1A, RORA), cold climate adaptation (e.g., TSHR, PRKG, OXCT1), and dryland adaptation (e.g., PLEKHA7, NFATC1, PLCB1). Conclusions This study unravels the unique adaptive diversity of three Chinese indigenous cattle breeds, providing a valuable resource for future research on sustainable livestock breeding strategies to response to climate change.

We investigated the expression profiles and functions of CircRNAs in the longissimus dorsi muscle of Datong yaks at different developmental stages, with the aim of clarifying their regulatory roles in skeletal muscle development. Samples of longissimus dorsi muscle were collected from Datong yaks at three developmental stages: 90-day-old fetuses, 6-month-old juveniles, and 3-year-old adults. High-throughput RNA sequencing was performed to identify CircRNAs. Differential expression analysis, along with GO and KEGG enrichment analyses, was conducted. A competing endogenous RNA (ceRNA) regulatory network was subsequently constructed to screen for core CircRNAs. A total of 17,027 CircRNAs were identified, with 6821 being differentially expressed. These differentially expressed CircRNAs showed significant enrichment in skeletal muscle development-related functions, such as sarcomere and calcium ion homeostasis, and were involved in key pathways, including the FoxO (Forkhead box O) and calcium signaling pathways. Construction of the ceRNA network revealed 20 core CircRNAs (e.g., CircRNA_10402 and CircRNA_15445), which may modulate the expression of 84 mRNAs by competing for binding with 42 miRNAs. This study preliminarily reveals the dynamic regulatory network of CircRNAs during skeletal muscle development in Datong yaks, providing new theoretical insights for understanding the molecular mechanisms underlying yak muscle development and for molecular breeding.

Fast-twitch and slow-twitch muscle fibers not only differ in metabolic characteristics and physiological functions but also significantly influence the texture of livestock meat. RNA editing represents an important post-transcriptional regulatory process that can influence both gene expression and the resulting protein function. However, studies on RNA editing events in yak muscle remain limited. This study systematically identified RNA editing events in yak biceps femoris (BF, n = 3) and obliquus externus abdominis (OEA, n = 3) using transcriptomic data, discovering 17,713 unique editing sites, most located in non-coding regions. Within coding regions, 3350 sites were detected, with 1195 resulting in non-synonymous amino acid substitutions. Further analysis revealed that 785 sites potentially affected miRNA binding sites, suggesting RNA editing may participate in miRNA-mediated gene regulation. Tukey’s post hoc test (p < 0.05) identified 242 sites (involving 170 genes) with significantly different editing levels between BF and OEA. KEGG pathway analysis indicated that genes with differential RNA editing were predominantly associated with pathways involved in muscle fiber type transitions, including the MAPK and calcium signaling pathways. Collectively, this study maps the RNA editing landscape in yak muscle tissue and identifies distinct, fiber-type-specific RNA editing patterns between oxidative and glycolytic muscle fibers, including differences in editing levels and site distributions, supporting a potential association between RNA editing and muscle fiber type transformation.

Background The hair follicle development process is regulated by sophisticated genes and signaling networks, and the hair grows from the hair follicle. The Tianzhu white yak population exhibits differences in hair length, especially on the forehead and shoulder region. However, the genetic mechanism is still unclear. Isoform sequencing (Iso-seq) technology with advantages in long reads sequencing. Hence, we combined the Iso-seq and RNA-seq methods to investigate the transcript complexity and difference between long-haired yak (LHY) and normal-haired yak (NHY). Results The hair length measurement result showed a significant difference between LHY and NHY on the forehead and the shoulder ( P-value  < 0.001). The skin samples from the forehead and the shoulder of LHY and NHY were pooled for isoform sequencing (Iso-seq). We obtained numerous long transcripts, including novel isoforms, long non-coding RNA, alternative splicing events, and alternative polyadenylation events. Combined with RNA-seq data, we performed differential isoforms (DEIs) analysis between LHY and NHY. We found that some hair follicle and skin development-related DEIs, like BMP4 , KRT2 , IGF2R , and COL1A2 in the forehead skin; BMP1 , KRT1 , FGF5 , COL2A1 , and IGFBP5 in the shoulder skin. Enrichment analysis revealed that DEIs in both two comparable groups significantly participated in skin and hair follicle development-related pathways, like ECM-receptor interaction, focal adhesion, and PI3K-Akt signaling pathways. The results indicated that the hair follicle development of Tianzhu white yak may influence the hair length difference. Besides, the protein-protein interaction (PPI) network of DEIs showed COL2A1 and COL3A1 exhibited a high degree of centrality, and these two genes were suggested as potential candidates for the hair length growth of Tianzhu white yak. Conclusions The results provided a comprehensive analysis of the transcriptome complexity and identified differential transcripts that enhance our understanding of the molecular mechanisms underlying the variation in hair length growth in Tianzhu white yak.

Animal growth and development are regulated by the growth hormone receptor (GHR). To elucidate the transcriptional regulation mechanism of the yak GHR gene, we cloned its proximal promoter region and performed bioinformatic analyses. Deletion constructs were generated to produce fragments of the proximal promoter, and the fragment exhibiting the highest activity was identified using a dual-luciferase reporter assay system. Moreover, the functionality of the transcription factor binding sites was confirmed via site-directed mutagenesis and electrophoretic mobility shift assay (EMSA). Bioinformatics analysis revealed two transcriptional initiation sites were located in the upstream regulation region at positions − 277/ − 327 and − 1734/ − 1784 bp, and CpG islands were located at − 552/ − 654 and − 82/ − 289 bp upstream. We identified the − 160 to + 27 bp region as the minimal core promoter of the yak GHR gene. The transcription factor KLF5, which is involved in muscle growth and development, was predicted to bind the proximal minimum core promoter region of the GHR gene. Using site-directed mutagenesis and EMSA, we demonstrated that KLF5 is a critical transcription regulator of the GHR gene. Our findings establish a theoretical basis for investigating the regulatory mechanisms of GHR gene expression.

Background There is no consensus as to the origin of the domestic yak ( Bos grunniens ). Previous studies on yak mitochondria mainly focused on mitochondrial displacement loop (D-loop), a region with low phylogenetic resolution. Here, we analyzed the entire mitochondrial genomes of 509 yaks to obtain greater phylogenetic resolution and a comprehensive picture of geographical diversity. Results A total of 278 haplotypes were defined in 509 yaks from 21 yak breeds. Among them, 28 haplotypes were shared by different varieties, and 250 haplotypes were unique to specific varieties. The overall haplotype diversity and nucleotide diversity of yak were 0.979 ± 0.0039 and 0.00237 ± 0.00076, respectively. Phylogenetic tree and network analysis showed that yak had three highly differentiated genetic branches with high support rate. The differentiation time of clades I and II were about 0.4328 Ma, and the differentiation time of clades (I and II) and III were 0.5654 Ma. Yushu yak is shared by all haplogroups. Most (94.70%) of the genetic variation occurred within populations, and only 5.30% of the genetic variation occurred between populations. The classification showed that yaks and wild yaks were first clustered together, and yaks were clustered with American bison as a whole. Altitude had the highest impact on the distribution of yaks. Conclusions Yaks have high genetic diversity and yak populations have experienced population expansion and lack obvious phylogeographic structure. During the glacial period, yaks had at least three or more glacial refugia.

The current study aimed to investigate the damage of long-term high concentrate diet feeding pattern on Yak jejunal structure, physiological function and protein composition during cold season. Twelve Datong male Yak ( Bos grunniens ) with the same age from cold season were randomly selected and slaughtered to determine Yak jejunal digestive enzyme activity, morphology and protein composition on different feeding patterns in Tibetan Plateau. The results showed that Yak jejunum digestive enzyme activity and morphology of grazing reared group were better than those in the intensively reared group. A total of 96 differentially expressed proteins were identified by label-free Mass Spectrometry (MS), which could be concluded to two predominant themes: protein structure and inflammatory response. Nine differentially expressed proteins were correlated in Yak jejunum damage in different feeding patterns. According to this research, we found that feeding pattern resulted the differences in Yak jejunum physiological function, morphology and protein composition. This fact was confirmed long-term high dietary concentrate feeding could damage the jejunum epithelial morphology and function.