高原家养动物环境适应性的研究进展

doi:10.11843/j.issn.0366-6964.2020.07.001

摘要/Abstract

摘要： 青藏高原环境主要体现为低氧低压、寒冷干燥、紫外线强、食物匮乏等，而高原家养动物经历了长期的选择和培育后能够生活在此环境下，其机体已经形成了独特的高原适应性特征。由于高原畜禽机体适应进化体系的复杂性，全面系统的适应性分子机制解析尚未完善。随着分子生物学和生物信息学的发展，高原家养动物基因组组装和功能注释完成，然后逐步开展了基因组、转录组和蛋白质组等组学层面的工作，挖掘到一系列高原动物环境适应性关键候选基因，为解析高原家养动物环境适应性分子机制研究提供了强有力支撑。本文以世居在青藏高原的藏鸡（Gallus gallus）、藏猪（Sus scrofa）、牦牛（Bos grunniens）、藏山羊（Capra hircus）、藏绵羊（Ovis aries）、藏马（Equus caballus）和藏獒（Canis lupus familiaris）等高原土著畜禽资源作为分类单元，分别从组织器官的解剖学结构、血液生理生化指标和分子遗传机制解析3个方面进行论述，并对高原适应性进化的研究趋势进行展望，以期为下一步培育高原高寒低氧地区新品种奠定基础。

关键词: 高原适应性, 组织器官, 血液生理指标, 分子机制

Abstract: The environment of the Qinghai-Tibet Plateau is mainly manifested in low oxygen and low pressure, cold and dry, strong ultraviolet rays and food shortage. The plateau domestic animals can live in this environment after long-term selection and cultivation, and their organisms have formed unique adaptive characteristics to the plateau. Due to complicated characteristics of plateau domestic animals adaptive evolutionary system, the comprehensive and systematic analysis of adaptive molecular mechanism has not been completed. With the development of molecular biology and bioinformatics, the genome assembly and functional annotation of the plateau domestic animals have been accomplished, and then the work of genome, transcriptome and proteome has been gradually carried out. Since then, a series of key candidate genes for plateau animals environmental adaptability have been identified. These provided further support for the analysis of the molecular mechanisms in the environment adaptation of plateau domestic animals. This review takes Tibetan chicken (Gallus gallus), Tibetan pig(Sus scrofa), yak(Bos grunniens), Tibetan goat(Capra hircus), Tibetan sheep(Ovis aries), Tibetan horse(Equus caballus) and Tibetan mastiff(Canis lupus familiaris) as the classification unit, their anatomical structure of tissues and organs, blood physiological and biochemical indexes and molecular genetic mechanism are discussed, respectively, and the future research of plateau adaptive evolution are envisaged, which will lay a theoretical foundation for the new variety breeding in the plateau high-cold and hypoxic regions.

Key words: high altitude adaptability, tissues and organs, blood physiological index, molecular mechanism

中图分类号:

Q951.3

张天留, 高雪, 徐凌洋, 陈燕, 张路培, 朱波, 高会江, 李俊雅. 高原家养动物环境适应性的研究进展[J]. 畜牧兽医学报, 2020, 51(7): 1475-1487.

ZHANG Tianliu, GAO Xue, XU Lingyang, CHEN Yan, ZHANG Lupei, ZHU Bo, GAO Huijiang, LI Junya. Research Progress on Environment Adaptation of Plateau Domestic Animals[J]. Acta Veterinaria et Zootechnica Sinica, 2020, 51(7): 1475-1487.

参考文献 75

[1]	BEALL C M.Two routes to functional adaptation:Tibetan and Andean high-altitude natives[J].Proc Natl Acad Sci U S A,2007,104(S1):8655-8660.
[2]	XU Z X,GONG T L,LI J Y.Decadal trend of climate in the Tibetan Plateau-regional temperature and precipitation[J]. Hydrol Processes,2008,22(16):3056-3065.
[3]	何俊峰,余四九,崔燕.不同年龄高原牦牛肺脏的组织结构特征[J].畜牧兽医学报,2009,40(5):748-755.HE J F,YU S J,CUI Y.Characteristics of lung structure in different age plateau yak[J].Acta Veterinaria et Zootechnica Sinica,2009,40(5):748-755.(in Chinese)
[4]	许永华,张东辉,许琴,等.西藏小型猪心脏、呼吸系统组织学观察[J].中国比较医学杂志,2009,19(8):61-62.XU Y H,ZHANG D H,XU Q,et al.Histological observation of heart and respiratory system in Tibet Minipigs[J].Chinese Journal of Comparative Medicine,2009,19(8):61-62.(in Chinese)
[5]	ANAND I S,HARRIS E,FERRARI R,et al.Pulmonary haemodynamics of the yak,cattle,and cross breeds at high altitude[J]. Thorax,1986,41(9):696-700.
[6]	SONG S,YAO N,YANG M,et al.Exome sequencing reveals genetic differentiation due to high-altitude adaptation in the Tibetan cashmere goat (Capra hircus)[J].BMC Genomics,2016,17(1):122.
[7]	卢晓丽,赵彦玲,吴征王,等.西藏色瓦藏绵羊高原适应性的血液生理学特性研究[J].西南农业学报,2019,32(6):1443-1447.LU X L,ZHAO Y L,WU Z W,et al.Study on blood physiological characteristics of plateau adaptation in Tibetan sawa sheep[J].Southwest China Journal of Agricultural Sciences,2019,32(6):1443-1447.(in Chinese)
[8]	WANG H F,WANG S S,ZHENG M,et al.Hypoxia promotes vasculogenic mimicry formation by vascular endothelial growth factor A mediating epithelial-mesenchymal transition in salivary adenoid cysticcarcinoma[J].Cell Proliferation, 2019, 52(3):e12600.
[9]	BAUMANN F,BAUER M S,REES M,et al.Increasing evidence of mechanical force as a functional regulator in smooth muscle myosin light chain kinase[J].Elife,2017,6(11):e26473.
[10]	WANG Y X,ZHENG Y M.ROS-dependent signaling mechanisms for hypoxic Ca2+ responses in pulmonary artery myocytes[J]. Antioxid Redox Signal,2010,12(5):611-623.
[11]	ADHIKARI A S,TRIVEDI D V,SARKAR S S,et al.β-Cardiac myosin hypertrophic cardiomyopathy mutations release sequestered heads and increaseenzymatic activity[J].Nat Commun,2019,10(1):2685.
[12]	CHANG Y W,LI L,ZHANG L P,et al.Plexin-B1 indirectly affects glioma invasiveness and angiogenesis by regulating the RhoA/αvβ3 signaling pathway and SRPK1[J].Tumor Biol,2016,37(8):11225-11236.
[13]	TASHI T,READING N S,WUREN T,et al.Gain-of-function EGLN1 prolyl hydroxylase (PHD2 D4E:C127S) in combination with EPAS1(HIF-2α) polymorphism lowers hemoglobin concentration in Tibetan highlanders[J].J Mol Med,2017,95(6):665-670.
[14]	WEIDEMANN A,JOHNSON R S.Biology of HIF-1α[J].Cell Death Differ,2008,15(4):621-627.
[15]	SASAKI R,MASUDA S,NAGAO M.Pleiotropicfunctions and tissue-specific expression of erythropoietin[J].News Physiol Sci,2001,16:110-113.
[16]	AGGARWAL S,NEGI S,JHA P,et al.EGLN1 involvement in high-altitude adaptation revealed through genetic analysis of extreme constitution types defined in Ayurveda[J].Proc Natl Acad Sci U S A,2010,107(44):18961-18966.
[17]	ROVNY R,MARKO M,KATINA S,et al.Association between genetic variability of neuronal nitric oxide synthase and sensorimotor gating in humans[J].Nitric Oxide,2018,80:32-36.
[18]	MATEJÁK M,KULHÁNEK T,MATOUČEK S.Adair-based hemoglobin equilibrium with oxygen,carbon dioxide and hydrogen ion activity[J].Scand J Clin Lab Invest,2015,75(2):113-120.
[19]	ZHANG Q,GOU W Y,WANG X T,et al.Genome resequencing identifies unique adaptations of Tibetan chickens to hypoxia and high-dose ultraviolet radiation in high-altitude environments[J].Genome Biol Evol,2016,8(3):765-776.
[20]	WANG M S,LI Y,PENG M S,et al.Genomic analyses reveal potential independent adaptation to high altitude in Tibetan chickens[J].Mol Biol Evol,2015,32(7):1880-1889.
[21]	LI M Z,TIAN S L,JIN L,et al.Genomic analyses identify distinct patterns of selection in domesticated pigs and Tibetan wild boars[J].Nat Genet,2013,45(12):1431-1438.
[22]	QIU Q,WANG L Z,WANG K,et al.Yak whole-genome resequencing reveals domestication signaturesand prehistoric population expansions[J].Nat Commun,2015,6(1):10283.
[23]	WEI C H,WANG H H,LIU G,et al.Genome-wide analysis reveals adaptation to high altitudes in Tibetan sheep[J].Sci Rep,2016,6(1):26770.
[24]	LIU X X,ZHANG Y L,LI Y F,et al.EPAS1 gain-of-function mutation contributes to high-altitude adaptation in Tibetan horses[J].Mol Biol Evol,2019,36(11):2591-2603.
[25]	GOU X,WANG Z,LI N,et al.Whole-genome sequencing of six dog breeds from continuous altitudes reveals adaptation to high-altitude hypoxia[J].Genome Res,2014,24(8):1308-1315.
[26]	张浩,吴常信,强巴央宗,等.高海拔孵化鸡胚死亡曲线分析[J].中国农业大学学报,2005,10(4):109-114.ZHANG H,WU C X,CHAMBA Y Z,et al.Curve analysis of embryonic mortality in chickens incubation at high altitude[J]. Journal of China Agricultural University,2005,10(4):109-114.(in Chinese)
[27]	杜志强,曲鲁江,李显耀,等.藏鸡群体遗传多样性研究[J].遗传,2004,26(2):167-171.DU Z Q,QU L J,LI X Y,et al.Genetic diversity in Tibetan chicken[J].Hereditas,2004,26(2):167-171.(in Chinese)
[28]	XIANG H,GAO J Q,YU B Q,et al.Early Holocene chicken domestication in northern China[J].Proc Natl Acad Sci U S A,2014,111(49):17564-17569.
[29]	ZHANG H,WU C X,CHAMBA Y,et al.Blood characteristics for high altitude adaptation in Tibetan chickens[J].Poult Sci, 2007, 86(7):1384-1389.
[30]	ZHANG H,WANG X T,CHAMBA Y,et al.Inf-luencesof hypoxia on hatching performance in chickens with different genetic adaptation to high altitude[J].Poult Sci,2008,87(10):2112-2116.
[31]	WEI Z H,ZHANG H,JIA C L,et al.Blood gas,hemoglobin,and growth of Tibetan chicken embryos incubated at high altitude[J]. Poult Sci,2007,86(5):904-908.
[32]	ZHANG Y W,GOU W Y,ZHANG Y,et al.Insights into hypoxic adaptation in Tibetan chicken embryos from comparative proteomics[J].Comp BiochemPhysiol Part D Genomics Proteomics,2019,31:100602.
[33]	LIU Y P,SHENG L,MA M H,et al.Proteome-based identification of chicken egg yolk proteins associated with antioxidant activity on the Qinghai-Tibetan Plateau[J].Int J Biol Macromol,2020,150:1093-1103.
[34]	ZHANG Z R,DU H R,BAI L J,et al.Whole genome bisulfite sequencing reveals unique adaptations to high-altitude environments in Tibetan chickens[J].PLoS One,2018,13(3):e0193597.
[35]	CHENG P.Livestock breeds of China[R].Food and Agriculture Organization of the United Nations,1985.
[36]	周琳,王蜀金.藏猪生理学研究进展[J].家畜生态学报,2014,35(7):7-12.ZHOU L,WANG S J.Research progress on Tibetan swine physiology[J].Acta Ecologae Animalis Domastici,2014, 35(7):7-12.(in Chinese)
[37]	强巴央宗,张浩,白玛央宗,等.高原环境中藏猪血液生理指标测定与比较[J].西南农业学报,2011,24(6):2382-2384.CHAMBA Y Z,ZHANG H,BAINA Y Z,et al.Determinationof blood physiological parameters in Tibet pig at high altitude[J]. Southwest China Journal of Agricultural Sciences,2011,24(6):2382-2384.(in Chinese)
[38]	AI H,HUANG L,REN J.Genetic diversity,linkage disequilibrium and selection signatures in Chinese and Western pigs revealed by genome-wide SNP markers[J].PLoS One,2013,8(2):e56001.
[39]	PHAM C G,BUBICI C,ZAZZERONI F,et al.Ferritin heavy chain upregulation by NF-κB inhibits TNFα-induced apoptosis by suppressing reactive oxygen species[J].Cell,2004,119(4):529-542.
[40]	AI H S,YANG B,LI J,et al.Population history and genomic signatures for high-altitude adaptation in Tibetan pigs[J].BMC Genomics,2014,15(1):834.
[41]	ZHANG B,QIANGBA Y Z,SHANG P,et al.Gene expression of vascular endothelial growth factor A and hypoxic adaptation in Tibetan pig[J].J Anim Sci Biotechnol,2016,7(1):21.
[42]	ZHANG B,CHAMBA Y,SHANG P,et al.Comparative transcriptomic and proteomic analyses provide insights into the key genes involved in high-altitude adaptation in the Tibetan pig[J].Sci Rep,2017,7(1):3654.
[43]	JIA C L,KONG X Y,KOLTES J E,et al.Genecoexpression network analysis unraveling transcriptional regulation of high-altitude adaptation of Tibetan pig[J].PLoS One,2016,11(12):e0168161.
[44]	魏青,俞红贤.180日龄高原牦牛和平原黄牛肺泡组织结构的比较研究[J].青海大学学报:自然科学版,2008,26(4):36-39.WEI Q,YU H X.Comparison of histological structure of pulmonary alveoli between 180 days old yak and plain cattle[J]. Journal of Qinghai University:Nature Science,2008,26(4):36-39.(in Chinese)
[45]	DING X Z,LIANG C N,GUO X,et al.Physiological insight into the high-altitude adaptations in domesticated yaks(Bos grunniens) along the Qinghai-Tibetan Plateau altitudinal gradient[J].Livestock Sci,2014,162:233-239.
[46]	QIU Q,ZHANG G J,MA T,et al.The yak genome and adaptation to life at high altitude[J].Nat Genet,2012,44(8):946-949.
[47]	CHEN N B,CAI Y D,CHEN Q M,et al.Whole-genome resequencing reveals world-wide ancestry and adaptive introgression events of domesticated cattle in East Asia[J].Nat Commun,2018,9(1):2337.
[48]	WU D D,DING X D,WANG S,et al.Pervasive introgression facilitated domestication and adaptation in the Bos species complex[J].Nat Ecol Evol,2018,2(7):1139-1145.
[49]	ZHANG X,WANG K,WANG L Z,et al.Genome-wide patterns of copy number variation in the Chinese yak genome[J].BMC Genomics,2016,17(1):379.
[50]	QI X B,ZHANG Q,HE Y X,et al.The transcriptomic landscape of yaks reveals molecular pathways for high altitude adaptation[J].Genome Biol Evol,2019,11(1):72-85.
[51]	XIN J W,CHAI Z X,ZHANG C F,et al.Transcriptome profiles revealed the mechanisms underlying the adaptation of yak to high-altitude environments[J].Sci Rep,2019,9(1):7558.
[52]	GUAN J Q,LONG K,MA J D,et al.Comparative analysis of the microRNA transcriptome between yak and cattle provides insight into high-altitude adaptation[J].PeerJ,2017,5:e3959.
[53]	XIONG X R,FU M,LAN D L,et al.Yak response to high-altitude hypoxic stress by altering mRNA expression and DNA methylation of hypoxia-inducible factors[J].Anim Biotechnol,2015,26(3):222-229.
[54]	江家椿,何玛利,嘎玛仁增,等.不同海拔高度西藏高原山羊若干血液生理特性的对比分析[J].西南农业学报,1992,5(1):79-83.JIANG J C,HE M L,GAMA R Z,et al.Comparison on several hematological values of goats in tibet plateau at different altitude[J].Southwest China Journal of Agricultural Sciences,1992,5(1):79-83.(in Chinese)
[55]	欧阳熙,王杰,王永,等.藏山羊血液生理、生化指标的季节性变化[J].西南民族学院学报:自然科学版,1992,18(3):284-288.OUYANG X,WANG J,WANG Y,et al.Seasonal variety of blood physiological and biochemical targets of Tibetan Goat[J]. Journal of SouthwestNationalities College:Natural Science,1992,18(3):284-288.(in Chinese)
[56]	DONG Y,XIE M,JIANG Y,et al.Sequencing and automated whole-genome optical mapping of the genome of a domestic goat (Capra hircus)[J].Nat Biotechnol,2013,31(2):135-141.
[57]	DONG Y,ZHANG X L,XIE M,et al.Reference genome of wild goat (Capra aegagrus) and sequencing of goat breeds provide insight into genic basis ofgoat domestication[J].BMC Genomics,2015,16(1):431.
[58]	ZHANG W P,FAN Z X,HAN E,et al.Hypoxia adaptations in the grey wolf (Canis lupus chanco) fromQinghai-Tibet Plateau[J].PLoS Genet,2014,10(7):e1004466.
[59]	LORENZO F R,HUFF C,MYLLYMÄKI M,et al.A genetic mechanism for Tibetan high-altitude adaptation[J].Nat Genet, 2014,46(9):951-956.
[60]	WANG X L,LIU J,ZHOU G X,et al.Whole-genome sequencing of eight goat populations for the detection of selection signatures underlying production and adaptive traits[J].Sci Rep,2016,6:38932.
[61]	GUO J Z,TAO H X,LI P F,et al.Whole-genome sequencing reveals selection signatures associated with important traits in six goat breeds[J].Sci Rep,2018,8(1):10405.
[62]	DENG J,FENG J,LI L,et al.Polymorphisms,diffe-rentiation,and phylogeny of 10 Tibetan goat populations inferred from mitochondrial D-loop sequences[J].Mitochondrial DNA Part A,2018,29(3):439-445.
[63]	LI X Y,WANG Y,GUO J Z,et al.Identification and expression patterns of adipokine genes during adipocyte differentiation in the Tibetan goat (Capra hircus)[J].Gene,2018,643:17-25.
[64]	WEI C H,WANG H H,LIU G,et al.Genome-wide analysis reveals population structure and selection in Chinese indigenous sheep breeds[J].BMC Genomics,2015,16(1):194.
[65]	YANG J,LI W R,LV F H,et al.Whole-genome sequencing of native sheep provides insights into rapid adaptations to extreme environments[J].Mol Biol Evol,2016,33(10):2576-2592.
[66]	HU X J,YANG J,XIE X L,et al.The genome landscape of Tibetan sheep reveals adaptive introgression from argali and the history of early human settlements on the Qinghai-Tibetan Plateau[J].Mol Bio.Evol,2018,36(2):283-303.
[67]	WANG W M,ZHANG X X,ZHOU X,et al.Deep genome resequencing reveals artificial and natural selection for visual deterioration,plateau adaptability and high prolificacy in Chinese domestic sheep[J].Front Genet,2019,10:300.
[68]	赵雪,魏雪锋,连林生,等.藏马低氧适应的血液生理指标研究[J].云南农业大学学报,2014,29(5):684-688.ZHAO X,WEI X F,LIAN L S,et al.Study on blood physiological indicators of adaptation to hypoxia in Tibet Horse[J].Journal of Yunnan Agricultural University,2014,29(5):684-688.(in Chinese)
[69]	XU S Q,LUOSANG J B,HUA S,et al.High altitude adaptation and phylogenetic analysis of Tibetan horse based on the mitochondrial genome[J].J Genet Genomics,2007,34(8):720-729.
[70]	YANG L,KONG X Y,YANG S L,et al.Haplotype diversity in mitochondrial DNA reveals the multiple origins of Tibetan horse[J].PLoS One,2018,13(7):e0201564.
[71]	周大鹏,刘建国,王芳,等.藏獒肺组织对高原低氧环境的适应特性[J].甘肃农业大学学报,2009,44(4):25-28.ZHOU D P,LIU J G,WANG F,et al.Pulmonary tissue adaptation to high altitude of Tibetan Mastiff[J].Journal of Gansu Agricultural University,2009,44(4):25-28.(in Chinese)
[72]	LI Y,WU D D,BOYKO A R,et al.Population variation revealed high-altitude adaptation of Tibetan mastiffs[J].Mol Biol Evol,2014,31(5):1200-1205.
[73]	MIAO B P,WANG Z,LI Y X.Genomic analysis reveals hypoxia adaptation in the Tibetan mastiff by introgression of the gray wolf from the Tibetan plateau[J].Mol Biol Evol,2017,34(3):734-743.
[74]	WU H,LIU Y H,WANG G D,et al.Identifying molecular signatures of hypoxia adaptation from sex chromosomes:a case for Tibetan Mastiff based on analyses of X chromosome[J].Sci Rep,2016,6(1):35004.
[75]	SHENDURE J,JI H.Next-generation DNA sequencing[J].Nat Biotechnol,2008,26(10):1135-1145.