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Sheep show characteristics of phenotypic diversity and adaptation to diverse climatic regions. Previous studies indicated associations between copy number variations (CNVs) and climate-driven adaptive evolution in humans and other domestic animals. Here, we constructed a genomic landscape of CNVs ( n = 39,145) in 47 old autochthonous populations genotyped at a set of high-density (600 K) SNPs to detect environment-driven signatures of CNVs using a multivariate regression model. We found 136 deletions and 52 duplications that were significantly ( P adj. < 0.05) associated with climatic variables. These climate-mediated selective CNVs are involved in functional candidate genes for heat stress and cold climate adaptation (e.g., B3GNTL1 , UBE2L3 , and TRAF2 ), coat and wool-related traits (e.g., TMEM9 , STRA6 , RASGRP2 , and PLA2G3 ), repairing damaged DNA (e.g., HTT ), GTPase activity (e.g., COPG ), fast metabolism (e.g., LMF2 and LPIN3 ), fertility and reproduction (e.g., SLC19A1 and CCDC155 ), growth-related traits (e.g., ADRM1 and IGFALS ), and immune response (e.g., BEGAIN ) in sheep. In particular, we identified significant ( RNF121 ) in sheep. In particular, we identified significant ( P < 0.05) enriched gene ontology terms and pathways related to functions such as nucleotide, protein complex, and GTPase activity. Additionally, we observed overlapping between the CNVs and 140 known sheep QTLs. Our findings imply that CNVs can serve as genomic markers for the selection of sheep adapted to specific climatic conditions.adj. < 0.05) associations between probes in deleted/duplicated CNVs and solar radiation. Enrichment analysis of the gene sets among all the CNVs revealed significant ( P adj. < 0.05) enriched gene ontology terms and pathways related to functions such as nucleotide, protein complex, and GTPase activity. Additionally, we observed overlapping between the CNVs and 140 known sheep QTLs. Our findings imply that CNVs can serve as genomic markers for the selection of sheep adapted to specific climatic conditions.

The domestication and subsequent development of sheep are crucial events in the history of human civilization and the agricultural revolution. However, the impact of interspecific introgression on the genomic regions under domestication and subsequent selection remains unclear. Here, we analyze the whole genomes of domestic sheep and their wild relative species. We found introgression from wild sheep such as the snow sheep and its American relatives (bighorn and thinhorn sheep) into urial, Asiatic and European mouflons. We observed independent events of adaptive introgression from wild sheep into the Asiatic and European mouflons, as well as shared introgressed regions from both snow sheep and argali into Asiatic mouflon before or during the domestication process. We revealed European mouflons might arise through hybridization events between a now extinct sheep in Europe and feral domesticated sheep around 6000-5000 years BP. We also unveiled later introgressions from wild sheep to their sympatric domestic sheep after domestication. Several of the introgression events contain loci with candidate domestication genes (e.g., PAPPA2, NR6A1, SH3GL3, RFX3 and CAMK4), associated with morphological, immune, reproduction or production traits (wool/meat/milk). We also detected introgression events that introduced genes related to nervous response (NEURL1), neurogenesis (PRUNE2), hearing ability (USH2A), and placental viability (PAG11 and PAG3) into domestic sheep and their ancestral wild species from other wild species., (© 2021. The Author(s).)

How animals, particularly livestock, adapt to various climates and environments over short evolutionary time is of fundamental biological interest. Further, understanding the genetic mechanisms of adaptation in indigenous livestock populations is important for designing appropriate breeding programs to cope with the impacts of changing climate. Here, we conducted a comprehensive genomic analysis of diversity, interspecies introgression, and climate-mediated selective signatures in a global sample of sheep and their wild relatives. By examining 600K and 50K genome-wide single nucleotide polymorphism data from 3,447 samples representing 111 domestic sheep populations and 403 samples from all their seven wild relatives (argali, Asiatic mouflon, European mouflon, urial, snow sheep, bighorn, and thinhorn sheep), coupled with 88 whole-genome sequences, we detected clear signals of common introgression from wild relatives into sympatric domestic populations, thereby increasing their genomic diversities. The introgressions provided beneficial genetic variants in native populations, which were significantly associated with local climatic adaptation. We observed common introgression signals of alleles in olfactory-related genes (e.g., ADCY3 and TRPV1) and the PADI gene family including in particular PADI2, which is associated with antibacterial innate immunity. Further analyses of whole-genome sequences showed that the introgressed alleles in a specific region of PADI2 (chr2: 248,302,667-248,306,614) correlate with resistance to pneumonia. We conclude that wild introgression enhanced climatic adaptation and resistance to pneumonia in sheep. This has enabled them to adapt to varying climatic and environmental conditions after domestication., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

Genetic structure and genetic diversity levels of indigenous Iranian sheep breeds are not clear, despite the interest this region has in itself as an important center for domestication of livestock. Early population genetic studies have reported high levels of diversity among Iranian sheep breeds until recently, when high admixture levels and genetic homogeneity have been detected. The rapid reduction of diversity observed in Iranian breeds might be due to an increasing trend of intensive crossbreeding practices or even total replacement of native breeds by highly specialized and productive ones. From a conservative perspective, this situation is highly concerning; thus, it might be wise to consider a conservation program in Iran to preserve the original genetic diversity in native sheep breeds. In this study, a total of 1065 animals with the purest morphological features representing 24 Iranian indigenous sheep breeds were sampled, corresponding to ancestral breed diversity. These samples were genotyped for 17 microsatellite loci in order to (1) determine the native ancestral diversity of Iranian breeds, (2) define the degree of genetic relationship among studied breeds, and (3) assess conservation priorities among defined groups. Our results showed no recent loss of diversity, but high genetic diversity levels for indigenous sheep breeds in Iran. Indeed, the analysis of conservation priorities pointed out the importance of 8 breeds for maintaining Iranian sheep breeds' maximum genetic diversity. Thus, under a genetic perspective, these 8 breeds should be the ones included into conservation programs for restocking endangered areas.

The immune systems are fundamentally vital for evolution and survival of species; as such, selection patterns in innate immune loci are of special interest in molecular evolutionary research. The interferon regulatory factor (IRF) gene family control many different aspects of the innate and adaptive immune responses in vertebrates. Among these, IRF3 is known to take active part in very many biological processes. We assembled and evaluated 1356 base pairs of the IRF3 gene coding region in domesticated goats from Africa (Nigeria, Ethiopia and South Africa) and Asia (Iran and China) and the wild goat (Capra aegagrus). Five segregating sites with θ value of 0.0009 for this gene demonstrated a low diversity across the goats' populations. Fu and Li tests were significantly positive but Tajima's D test was significantly negative, suggesting its deviation from neutrality. Neighbor joining tree of IRF3 gene in domesticated goats, wild goat and sheep showed that all domesticated goats have a closer relationship than with the wild goat and sheep. Maximum likelihood tree of the gene showed that different domesticated goats share a common ancestor and suggest single origin. Four unique haplotypes were observed across all the sequences, of which, one was particularly common to African goats (MOCH-K14-0425, Poitou and WAD). In assessing the evolution mode of the gene, we found that the codon model dN/dS ratio for all goats was greater than one. Phylogenetic Analysis by Maximum Likelihood (PAML) gave a ω0 (dN/dS) value of 0.067 with LnL value of -6900.3 for the first Model (M1) while ω2 = 1.667 in model M2 with LnL value of -6900.3 with positive selection inferred in 3 codon sites. Mechanistic empirical combination (MEC) model for evaluating adaptive selection pressure on particular codons also confirmed adaptive selection pressure in three codons (207, 358 and 408) in IRF3 gene. Positive diversifying selection inferred with recent evolutionary changes in domesticated goat IRF3 led us to conclude that the gene evolution may have been influenced by domestication processes in goats., Competing Interests: The authors have declared that no competing interests exist.

A crossbreeding experiment was carried out to determine the sire breed effects on fattening performance of fat-tailed lambs. Rams from four Iranian fat-tailed sheep breeds, namely Kurdi (K), Afshari (A), Chaal (C) and Sanjabi (S) were mated to Kurdi ewes producing 454 progeny of four genotype groups (KK, AK, CK and SK). A total of 80 lambs comprising ten female and ten male lambs from each genotype group were used in a fattening experiment during a period of 88 days. Genotype of the lamb had significant effect on average dry matter intake (ADMI) (P < 0.05). SK lambs had the lowest ADMI. Average daily gain (ADG) and food conversion ratio (FCR) were not influenced by genotype of the lamb. Sex of the lamb had significant effect on both ADG and FCR (P < 0.01). Slaughter weight (SLW), hot carcass weight (HCW), cold carcass weight (CCW) and daily carcass gain (DCG) were not affected by lamb's genotype. However, genotype had significant effect on dressing percentage, carcass loss percentage (P < 0.01), carcass length (CL), leg circumference (LC), carcass compactness (CC) (P < 0.01) and carcass density (CD) (P < 0.05). Carcass width (CW) and leg length (LL) were not affected by lamb's genotype. Sex of the lamb had significant effects on CL, CW, LC, CC, CD (P < 0.01) and LL (P < 0.05). The results indicated the existence of sire breed difference for daily dry matter intake and carcass conformation in the fat-tailed sheep.

BRCA1 gene plays an important role in DNA damage repair, cell cycle, and transcription process regulation; hence it's called gate keeper. The current research aims to perform bioinformatics analyzes of the BRCA1 gene of different breeds of domestic and wild sheep from 49 breeds in 14 countries using the NCBI genome database. The desired sequences were aligned using MEGA11 software and a phylogenetic tree was drawn by Neighbor-Joining method. The number of mutations, nucleotide diversity, and haploid diversity were also analyzed using Dnaspv5 software. The analyses showed 296 polymorphisms, which led to the creation of 45 different haplotypes with a haplotype diversity of 0.035. Nucleotide diversity and average nucleotide differences among breeds were estimated as 0.259 and 0.052, respectively. The average genetic distance within the population of countries was calculated as 0.052. The amount of sequence conservation in this research was 0.313 on average, which indicates the high polymorphism of this gene and the emergence of new proteins. Tajima's D value in Tajima's neutrality test was −2.421, which was significant (p < 0.05). One of the reasons for the high genetic diversity in Iran's wild sheep population is the existence of forests and open environments, which prevent genetic drift and reduce inbreeding. [ABSTRACT FROM AUTHOR]

It's well-documented that most economic traits have a complex genetic structure that is controlled by additive and non-additive gene actions. Hence, knowledge of the underlying genetic architecture of such complex traits could aid in understanding how these traits respond to the selection in breeding and mating programs. Computing and having estimates of the non-additive effect for economic traits in sheep using genome-wide information can be important because; non-additive genes play an important role in the prediction accuracy of genomic breeding values and the genetic response to the selection. This study aimed to assess the impact of non-additive effects (dominance and epistasis) on the estimation of genetic parameters for body weight traits in sheep. This study used phenotypic and genotypic belonging to 752 Scottish Blackface lambs. Three live weight traits considered in this study were included in body weight at 16, 20, and 24 weeks). Three genetic models including additive (AM), additive + dominance (ADM), and additive + dominance + epistasis (ADEM), were used. The narrow sense heritability for weight at 16 weeks of age (BW16) were 0.39, 0.35, and 0.23, for 20 weeks of age (BW20) were 0.55, 0.54, and 0.42, and finally for 24 weeks of age (BW24) were 0.16, 0.12, and 0.02, using the AM, ADM, and ADEM models, respectively. The additive genetic model significantly outperformed the non-additive genetic model (p < 0.01). The dominance variance of the BW16, BW20, and BW24 accounted for 38, 6, and 30% of the total phenotypic, respectively. Moreover, the epistatic variance accounted for 39, 0.39, and 47% of the total phenotypic variances of these traits, respectively. In addition, our results indicated that the most important SNPs for live weight traits are on chromosomes 3 (three SNPS including s12606.1, OAR3_221188082.1, and OAR3_4106875.1), 8 (OAR8_16468019.1, OAR8_18067475.1, and OAR8_18043643.1), and 19 (OAR19_18010247.1), according to the genome-wide association analysis using additive and non-additive genetic model. The results emphasized that the non-additive genetic effects play an important role in controlling body weight variation at the age of 16–24 weeks in Scottish Blackface lambs. It is expected that using a high-density SNP panel and the joint modeling of both additive and non-additive effects can lead to better estimation and prediction of genetic parameters. [ABSTRACT FROM AUTHOR]

Genomic assessment of local sheep breeds is relevant to the reconstruction of post-domestication migrations and to filling in gaps in the developmental history and contemporary phylogeographic-differentiation patterns in Eurasia. In this study, we aimed to reveal possible genetic relationships between local sheep breeds in Russia and the Persian Highlands (Iran) based on high-density SNP genotypes. All samples (n = 395) from 11 Iranian and 10 Russian sheep breeds were genotyped by using the Ovine Infinium HD BeadChip (Illumina, San Diego, CA, USA). Principal Component Analysis, maximum-likelihood assessment, and Neighbor-Net graph analysis demonstrated a clear differentiation between Russian sheep breeds of European ancestry from the Iranian local cluster and Russian breeds of Asian origin. Admixture analysis revealed a shared ancestral background, which was detected in several Iranian-local and Russian-local fat-tailed coarse-wool sheep breeds. Our findings contribute to a better understanding of the pattern of historic admixture, which is present in the genomes of many Eurasian sheep breeds. [ABSTRACT FROM AUTHOR]

Background: Artificial and natural selection for important economic traits and genetic adaptation of the populations to specific environments have led to the changes on the sheep genome. Recent advances in genome sequencing methods have made it possible to use comparative genomics tools to identify genes under selection for traits of economic interest in domestic animals. Objectives: In this study, we compared the genomes of Assaf and Awassi sheep breeds with those of the Cambridge, Romanov and British du cher sheep breeds to explore positive selection signatures for milk traits using nucleotide diversity (Pi) and FST statistical methods. Methods: Genome sequences from fourteen sheep with a mean sequence depth of 9.32X per sample were analysed, and a total of 23 million single nucleotide polymorphisms (SNPs) were called and applied for this study. Genomic clustering of breeds was identified using ADMIXTURE software. The FST and Pi values for each SNP were computed between population A (Assaf and Awassi) and population B (Cambridge, British du cher, and Romanov). Results: The results of the PCA grouped two classes for these five dairy sheep breeds. The selection signatures analysis displayed 735 and 515 genes from FST and nucleotide diversity (Pi) statistical methods, respectively. Among all these, 12 genes were shared between the two approaches. The most conspicuous genes were related to milk traits, including ST3GAL1 (the synthesis of oligosacáridos), CSN1S1 (milk protein), CSN2 (milk protein), OSBPL8 (fatty acid traits), SLC35A3 (milk fat and protein percentage), VPS13B (total milk production, fat yield, and protein yield), DPY19L1 (peak yield), CCDC152 (lactation persistency and somatic cell count), NT5DC1 (lactation persistency), P4HTM (test day protein), CYTH4 (FAT Production) and METRNL (somatic cell), U1 (milk traits), U6 (milk traits) and 5S_RRNA (milk traits). Conclusions: The findings provide new insight into the genetic basis of sheep milk properties and can play a role in designing sheep breeding programs incorporating genomic information. [ABSTRACT FROM AUTHOR]

Runs of homozygosity (ROH) stretches are continuous homozygous fragments of the genome, which are more suitable for calculating the genomic inbreeding, identifying footprints involved in economic traits and understanding the population history in livestock species. In this study, using a dataset of Ovine SNP50 BeadChip genotypes, the distribution of ROH in nine sheep populations (Soay n= 110, Australian Poll Dorset n=108, Australian Suffolk n=109, Finn sheep n= 99, Scottish Texel n= 80, Scottish Blackface n= 56, Galway n= 49, Border Leicester n= 48, German Texel n=46) from Europe were investigated. ROHs were detected by using PLINK v1.09 with the minimal number of SNPs in ROH set at 40; the maximal gap between the adjacent SNPs was set to 1Mb; the minimum SNP density per ROH was set to 1/100 kb and no heterozygote less than 16 Mb was allowed. The detected ROHs were specified to four categories, based on their length: 1–4 Mb, 4–8 Mb, 8–16 Mb and above 16 Mb. A total of 22,204 ROHs were identified, in which ~ 92 – 98 % were less than 16 Mb in length, covering 4.6% to 12.9% of the entire genome. The inbreeding coefficient based on ROH (FROH) varied among populations (ranging from 0.05 to 0.14). The highest inbreeding rate was found in Border Leicester and Soay breeds. In addition, we detected 90 possible ROH Islands that overlapped with candidate genes associated with different economic traits such as the body weight, meat production, fat deposition, horn-less, and coat color in sheep. Our results suggested that although genetic selection for meat and wool traits in these breeds have been extensively carried out during the last decades, the autozygotic proportion of the genome is still considerably low, and it could lead to an acceptable response to selection in breeding schemes. [ABSTRACT FROM AUTHOR]

Copy number variations (CNVs) are a major source of structural variation in mammalian genomes. Here, we characterized the genome-wide CNV in 2059 sheep from 67 populations all over the world using the Ovine Infinium HD (600K) SNP BeadChip. We tested their associations with distinct phenotypic traits by conducting multiple independent genome-wide tests. In total, we detected 7547 unique CNVs and 18,152 CNV events in 1217 non-redundant CNV regions (CNVRs), covering 245 Mb (∼10%) of the whole sheep genome. We identified seven CNVRs with frequencies correlating to geographical origins and 107 CNVRs overlapping 53 known quantitative trait loci (QTLs). Gene ontology and pathway enrichment analyses of CNV-overlapping genes revealed their common involvement in energy metabolism, endocrine regulation, nervous system development, cell proliferation, immune, and reproduction. For the phenotypic traits, we detected significantly associated (adjusted P < 0.05) CNVRs harboring functional candidate genes, such as SBNO2 for polycerate; PPP1R11 and GABBR1 for tail weight; AKT1 for supernumerary nipple; CSRP1 , WNT7B , HMX1 , and FGFR3 for ear size; and NOS3 and FILIP1 in Wadi sheep; SNRPD3 , KHDRBS2 , and SDCCAG3 in Hu sheep; NOS3 , BMP1 , and SLC19A1 in Icelandic; CDK2 in Finnsheep; MICA in Romanov; and REEP4 in Texel sheep for litter size. These CNVs and associated genes are important markers for molecular breeding of sheep and other livestock species. [ABSTRACT FROM AUTHOR]

Investigation of genetic relationship among populations has been traditionally based on the analysis of allele frequencies at different loci. The prime objective of this research was to measure the genetic polymorphism of five microsatellite markers (McMA2, BM6444, McMA26, HSC, and OarHH35) and study genetic diversity of 14 sheep types in Iran. Genomic DNA was extracted from blood samples of 565 individuals using an optimized salting-out DNA extraction procedure. The polymerase chain reaction (PCR) was successfully performed with the specific primers. Some locus-population combinations were not at Hardy-Weinberg equilibrium (P < 0:05). The microsatellite analysis revealed high allelic and gene diversity in all 14 breeds. Pakistani and Arabi breeds showed the highest mean number of alleles (11.8 and 11 respectively), while the highest value for polymorphic information content was observed for the Arabi breed (0.88). A UPGMA (unweighted pair group method with arithmetic mean) dendrogram based on the Nei's standard genetic distance among studied breeds showed a separate cluster for Arabi and Pakistani breeds and another cluster for other breeds. The Shannon index (H0) for McMA2, BM6444, McMA26, HSC, and OarHH35 was 2.31, 2.17, 2.27, 2.04 and 2.18, respectively, and polymorphic information content (PIC) values were 0.88, 0.92, 0.87, 0.84, and 0.86 for McMA2, BM6444, McMA26, HSC, and OarHH35, respectively. The high degree of variability demonstrated within the studied sheep types implies that these populations are rich reservoirs of genetic diversity that must be preserved. [ABSTRACT FROM AUTHOR]

The mammary gland experiences vast changes between the onset of lactation and pregnancy. This remodeling involves different functions such as lactation that is controlled by innumerable regulators and various gene networks which are still not completely understood. MicroRNAs (miRNAs) are one of the important non-coding gene regulators which control an extensive range of biological processes. Thus, exploring miRNAs functions is important for solving gene regulation complexity. The main purpose in the present study is to identify the various gene regulative integrated networks involved in lactation progress in mammary gland. We analyzed ovine mammary tissue data sets which included expression profiles of mRNA (genes) and miRNAs related to six ewes in different days of lactation and nutritional treatments. We combined two different types of information: the network that is module inference by mRNAs (RNA-seq data), miRNAs and transcription factors (TFs) expression matrix and prediction of targets via computational methods. To discover the miRNAs regulatory function, 134 modules were predicted by using gene expression data and 14 TFs and 20 miRNAs were allocated to these predicted modules. By applying this integrated computation-based method, 38 miRNA-modules and 35 TF-module interactions were identified from ovine mammary tissue data during lactogenesis. A lot of these modules were involved in lipid and protein metabolism, as well as steroids and vitamin biosynthesis, which would play key roles in mammary tissue and lactation development. These results present new information about the regulatory procedures at the miRNAs and TF levels throughout lactation. • The regulatory networks involved in sheep lactogenesis was constructed by analysis of mammary tissue transcriptomes. • The module inference network by TFs, miRNAs, and mRNA expression profiles was integrated into computational target predictions. • A total of 38 miRNA-modules and 35 TF-module interactions were identified by the integrated computation-based method. • The GO analysis results emphasized the key role of fatty acids, lipids, steroids, and vitamin in the milk secretory cells. [ABSTRACT FROM AUTHOR]

Archaeological and genetic evidence show that sheep were originally domesticated in area around the North of Zagros mountains, North-west of Iran. The Persian plateau exhibits a variety of native sheep breeds with a common characteristic of coarse-wool production. Therefore, knowledge about the genetic structure and diversity of Iranian sheep and genetic connections with other sheep breeds is of great interest. To this end, we genotyped 154 samples from 11 sheep breeds distributed across Iran with the Ovine Infinium HD SNP 600 K BeadChip array, and analyzed this dataset combined with the retrieved data of 558 samples from 19 worldwide coarse-wool sheep breeds. The average genetic diversity ranged from 0.315 to 0.354, while the F ST values ranged from 0.016 to 0.177 indicating a low differentiation of Iranian sheep. Analysis of molecular variance showed that 90.21 and 9.79% of the source of variation were related to differences within and between populations, respectively. Our results indicated that the coarse-wool sheep from Europe were clearly different from those of the Asia. Accordingly, the Asiatic mouflon was positioned between Asian and European countries. In addition, we found that the genetic background of Iranian sheep is present in sheep from China and Kyrgyzstan, as well as India. The revealed admixture patterns of the Iranian sheep and other coarse-wool sheep breeds probably resulted from the expansion of nomads and through the Silk Road trade network. • Low genetic differentiation was detected for the Iranian sheep populations. • There is a high genetic diversity within Iranian native sheep breeds. • The effective population size is declining in Iranian native breeds. • There is a common ancestry between the Iranian and Asian coarse-wool sheep. • There are admixture patterns for the Iranian sheep and other coarse-wool sheep breeds. [ABSTRACT FROM AUTHOR]

Genome-wide pattern of runs of homozygosity (ROH) across ovine genome can provide a useful resource for studying diversity and demography history in sheep. We analyzed 50 k SNPs chip data of 2536 animals to identify pattern, distribution and level of ROHs in 68 global sheep populations. A total of 60,301 ROHs were detected in all breeds. The majority of the detected ROHs were <16 Mb and the average total number of ROHs per individual was 23.8 ± 13.8. The ROHs greater than 1 Mb covered on average 8.2% of the sheep autosomes, 1% of which was related to the ROHs with 1–4 Mb of length. The mean sum of ROH length in two-thirds of the populations was less than 250 Mb ranging from 21.7 to near 570 Mb. The level of genomic inbreeding was relatively low. The average of the inbreeding coefficients based on ROH (F ROH) was 0.09 ± 0.05. It was rising in a stepwise manner with distance from Southwest Asia and maximum values were detected in North European breeds. A total of 465 ROH hotspots were detected in 25 different autosomes which partially surrounding 257 Refseq genes across the genome. Most of the detected genes were related to growth, body weight, meat production and quality, wool production and pigmentation. In conclusion, our analysis showed that the sheep genome, compared with other livestock species such as cattle and pig, displays low levels of homozygosity and appropriate genetic diversity for selection response and genetic merit gain. • The genome-wide pattern of Runs of Homozygosity (ROH) across world-wide sheep populations was evaluated. •.The sheep genome compared with other livestock such as cattle and pigs displays low homozygosity • A total of 465 ROH hotspots were detected surrounding 257 genes across the genome. [ABSTRACT FROM AUTHOR]