Chinese Journal of Animal Nutrition >

2025 , Vol. 37 >Issue 11: 7264 - 7275

DOI: https://doi.org/10.12418/CJAN2025.591

REVIEW

Research Progress on Influencing Factors and Molecular Mechanisms of Fattening Performance in Chinese Yellow Cattle

  • ZHI Li , 1, 2 ,
  • GUO Xiaodan 1 ,
  • YU Ying 3 ,
  • MAO Huaming , 1, *
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  • 1 College of Animal Science and Technology, Yunnan Agricultural University, Kunming 650201, China
  • 2 Faculty of Animal Science, Xichang University, Xichang 615000, China
  • 3 College of Animal Science and Technology, China Agricultural University, Beijing 100091, China
*professor, E-mail:

Received date: 2025-04-19

  Online published: 2025-11-14

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Abstract

With the continuous improvement of living standards, consumer demand for both the quantity and quality of beef has been steadily increasing. Chinese yellow cattle are highly favored by consumers due to their superior eating quality. However, their low meat production performance has hindered the development of the industry. This paper systematically reviews the main factors influencing the fattening performance of Chinese yellow cattle, including breed and region, nutritional levels and feeding methods. It further compares carcass trait changes within the same breed across different growth periods. Finally, the molecular mechanisms underlying fattening performance are explored. The aim is to provide a theoretical basis for the conservation of Chinese yellow cattle genetic resources, efficient breeding practices, and genetic selection programs.

Key words： Chinese yellow cattle; fattening performance; influencing factors; carcass traits; molecular mechanisms

Cite this article

ZHI Li , GUO Xiaodan , YU Ying , MAO Huaming . Research Progress on Influencing Factors and Molecular Mechanisms of Fattening Performance in Chinese Yellow Cattle[J]. Chinese Journal of Animal Nutrition, 2025 , 37(11) : 7264 -7275 . DOI: 10.12418/CJAN2025.591

我国广袤的土地上分布着丰富的牛遗传资源,不同的地理环境加上多样化的适应性选择共同促成了我国黄牛的遗传和表型上的多样性[1],根据《国家畜禽遗传资源品种名录(2024年版)》统计,我国现有84个普通牛品种,包括58个地方品种、11个培育品种和15个引入品种,这些品种资源构成了我国肉牛产业发展的基础,但各品种表现出不同的育肥性能、胴体性状和环境适应性,相较于引入品种,中国黄牛以其较强的抗病性和抗应激能力著称,尽管生长效率普遍相对较低,影响了产肉量,但凭借其优越的食用品质,依然深受消费者喜爱[2],因此提高中国黄牛的育肥性能具有重要意义。
育肥性能受到多种因素影响,本文系统综述了品种和地域、营养水平和饲喂方式对中国黄牛育肥性能的影响,并对比了同品种不同时期中国黄牛胴体性状的变化,最后探讨了影响中国黄牛育肥性能的分子机理,旨在为中国黄牛遗传资源保护、高效养殖以及遗传选育方面提供理论依据。

1 中国黄牛育肥性能影响因素

1.1 品种和地域

表1列举了部分中国黄牛(地方品种和培育品种)的屠宰性能,包括宰前活重、胴体重、屠宰率和瘦肉率。Ge等[2]对9个品种中国黄牛(新疆褐牛、平凉红牛、云岭牛、晋南牛、鲁西牛、渤海黑牛、郏县红牛、夷陵牛、文山牛)的屠宰性能进行对比研究,每个品种均选取28~36月龄肉牛并饲喂相同的饲粮(代谢能水平为11.78 MJ/kg,粗蛋白质水平为8.65%),育肥12个月后进行屠宰,结果表明与地方品种相比,培育品种云岭牛、新疆褐牛和平凉红牛表现出更优越的屠宰性能,其中云岭牛在宰前活重、胴体重和净肉重方面分别超过9个品种平均值的34.58%、37.32%和36.71%;另外,南方地方品种(文山牛、夷陵牛)屠宰性能低于北方地方品种(晋南牛、鲁西牛、渤海黑牛和郏县红牛),其中夷陵牛在宰前活重(378.67 kg)、胴体重(223.40 kg)和净肉重(203.70 kg)与其他中国黄牛相比表现出最低值。另外,从表1可以看出,培育品种的屠宰性能普遍高于地方品种,从地域上可以看到从北到南,中国黄牛屠宰性能呈现“高→低”的变化趋势。造成这种差异的主要原因是由于成年牛体重的不同,而牛的体重受到饲料条件、环境气候、历史文化以及种质交流等因素的综合影响。中国黄牛地方品种来源于普通牛(Bos taurus)和瘤牛(Bos taurus indicus)2个祖先[3],普通牛和瘤牛在向全球扩散的过程中会在交汇区域发生种群混合,导致它们的血统比例在地理区域上呈现差异化,具体而言,由南到北普通牛血统呈现逐渐上升而瘤牛血统呈现逐渐下降的趋势[4]。因此,相比于北方地方品种,南方地方品种因含有更高比例的瘤牛血统,并在饲料条件、环境气候等因素共同作用下形成了体型较小、耐湿热的特征。另外,培育品种通过长期杂交选育而获得优良的生产性能,而地方品种虽然体型较小,但具有更强的生长潜力。Meng等[5]关于12和18月龄云岭牛和文山牛的体尺、体重对比研究显示,云岭牛的体重、体高等均显著高于文山牛,但值得注意的是,文山牛在12和18月龄的生长速度显著高于云岭牛。
表1 中国黄牛屠宰性能对比

Table 1 Comparisons on slaughter performance of Chinese yellow cattle

发表年份
Publication
year		 品种
Breed		 省份
Province		 屠宰月龄
Slaughter
month of age		 饲粮组成
Diet
composition		 宰前活重
Slaughter
weight/kg		 胴体重
Carcass
weight/kg		 屠宰率
Dressing
percentage/%		 净肉率
Lean meat
percentage/%
2023 新疆褐牛[2]
(培育品种)		 新疆 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 692.00±36.07 418.47±27.80 60.48±2.86 53.81±1.41
2023 平凉红牛[2]
(培育品种)		 甘肃 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 638.33±48.69 376.33±24.01 59.01±1.82 56.14±2.14
2023 中国西门
塔尔牛[6]
(培育品种)		 内蒙 — — 577.33±46.36 315.67±26.99 54.66±0.46 —
2023 华西牛[7]
(培育品种)		 内蒙 18 — 516.37±93.33 313.80±63.01 60.42±4.63 51.69±4.37
2023 云岭牛[2]
(培育品种)		 云南 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 826.00±42.46 507.00±23.07 61.40±0.85 55.54±0.52
2023 延边牛[8]
(地方品种)		 吉林 — 玉米秸秆48.0%、玉米37.0%、豆粕7.0%、干酒糟及
其可溶物2.6%、玉米胚芽粕2.0%、碳酸氢钠1.0%、
谷氨酸钠蛋白1.0%、盐0.4%、预混料1.0%		 588.60 342.34 57.98 —
2023 晋南牛[2]
(地方品种)		 山西 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 536.00±39.15 337.40±45.37 62.76±4.15 56.77±4.11
2023 鲁西牛[2]
(地方品种)		 山东 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 656.10±39.97 384.97±35.11 58.60±1.75 53.40±1.51
2023 渤海黑牛[2]
(地方品种)		 山东 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 715.00±13.23 413.67±18.45 57.89±3.30 52.78±3.63
2023 郏县红牛[2]
(地方品种)		 河南 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 562.67±43.92 348.53±40.72 61.83±3.05 55.68±3.20
2023 夷陵牛[2]
(地方品种)		 湖北 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 378.67±20.65 223.40±19.62 58.92±2.04 53.79±1.07
2021 湘西黄牛[9]
(地方品种)		 湖南 30 青贮玉米65.0%、玉米12.0%、棉籽饼6.5%、麦麸
13.0%、预混料1.0%、石粉0.5%、食盐0.3%、
碳酸氢钠0.5%、磷酸氢钙1.2%		 222.75±16.62 — 48.98±1.23 —
2019 锦江牛[10]
(地方品种)		 江西 — 青贮料43.36%、玉米36.18%、小麦麸14.15%、豆粕
3.38%、小苏打0.22%、食盐0.11%、预混料2.60%		 350.30 195.80 55.86 —
2022 秦川牛[11]
(地方品种)		 陕西 22 精料:玉米55%、麸皮12%、豆粕25%、食盐1%、
碳酸氢钙2%、预混料5%
粗料:全株玉米青贮、干草		 391.00 238.17 60.65 48.74
2020 三江牛[12]
(地方品种)		 四川 48 — 309.55 162.44 52.48 43.85
2021 宣汉黄牛[13]
(地方品种)		 四川 29.6 — 478.54±9.44 260.67±4.50 54.63±0.12 —
2023 威宁黄牛[14]
(地方品种)		 贵州 27±1 黑麦草30.00%、青贮玉米35.00%、玉米12.00%、麦麸
12.00%、豆粕7.90%、碳酸钙0.55%、碳酸氢钙0.45%、
食盐0.10%、碳酸氢钠1.00%、预混料1.00%		 249.71±12.88 133.52±8.77 53.47±3.19 44.81±3.92
2021 务川黑牛[15]
(地方品种)		 贵州 18 玉米青贮83.23%、玉米10.06%、麦麸2.35%、豆粕
2.60%、玉米脱水酒精糟1.21%、碳酸钙0.13%、
碳酸氢钙0.13%、碳酸氢钠0.08%、食盐0.13%、预混料0.06%		 260.44 142.36 54.66 46.04
2025 思南牛[16]
(地方品种)		 贵州 36 — 357.55±32.62 182.81±53.84 55.98±3.00 47.25±3.04
2023 文山牛[2]
(地方品种)		 云南 28~36 玉米青贮39.96%、玉米37.07%、麸皮17.62%、棉籽粕
5.00%、矿物质饲料2.81%、食盐0.54%、预混料2.00%		 519.00±23.07 313.13±16.02 60.32±0.61 53.84±1.12
2022 红河黄牛[17]
(地方品种)		 云南 26 玉米青贮58%、蛋白粉2%、脂肪粉4%、精料36%
(精料:豆粕12%、玉米蛋白粉9%、酒糟5%、
玉米63%、麸皮6%、磷酸氢钙3%、食盐1%、预混料1%)		 271.83 151.38 54.66 45.96
2020 江城牛[18]
(地方品种)		 云南 34.8 精料:豆粕12%、玉米蛋白粉9%、酒糟5%、
玉米63%、麸皮6%、磷酸氢钙3%、食盐1%、预混料1%
粗料:全株玉米青贮		 199.85±51.12 — 54.45±2.01 41.02±0.30
2024 滇中牛[19]
(地方品种)		 云南 24±2 精料:豆粕、菜籽粕、玉米、氯化钠、
碳酸氢钙、矿物元素及多种维生素、抗氧化剂
粗料:全株玉米青贮		 270.53±25.77 157.60±16.76 58.22±1.96 48.43±2.16

1.2 营养水平和饲喂方式

前人大量的研究探究了饲粮中不同营养水平对中国黄牛胴体性状的影响。邵三勤等[20]研究了3个不同能量水平[前期综合净能(NEmf)水平分别为6.21、6.52、6.83 MJ/kg,粗蛋白质水平为12%;中期NEmf水平分别为7.14、7.45、7.76 MJ/kg,粗蛋白质水平为11.6%;末期NEmf水平分别为8.07、8.38、8.69 MJ/kg,粗蛋白质水平为11%]玉米秸秆青贮饲料对郏县红牛[体重(363±22) kg]胴体性状的影响,饲喂127 d后屠宰,结果表明随着能量水平的提高,郏县红牛屠宰率、胴体产肉率、背膘厚度和眼肌面积有所增加,除了高能组背膘厚度显著高于低能组,其他指标均无显著差异。柏峻等[10]对50头锦江牛[体重(301.7±30.1) kg]随机分为5个组,A组饲喂基础饲粮,B~E组饲粮的NEmf水平在A组的基础上依次增加6%、12%、18%和24%,5个组饲粮粗蛋白质水平约为14%,饲喂116 d后5个组的胴体重、屠宰率、背膘厚度和眼肌面积均无显著差异,但背膘厚度随着NEmf水平的提高呈现先下降后上升的趋势,其中C组(NEmf水平为6.74 MJ/kg)背膘厚度最低(2.01 cm),眼肌面积随着NEmf水平的提高逐渐增加,E组(NEmf水平为7.46 MJ/kg)眼肌面积最高(81.04 cm2)。Lin等[21]将15头红河黄牛[体重(259.42±23.87) kg]分为3个组,分别为低能组(3.72 MJ/kg)、中能组(4.52 MJ/kg)和高能组(5.32 MJ/kg),3个组饲粮粗蛋白质水平均为11.7%,饲喂70 d后屠宰,结果显示胴体重、屠宰率、净肉重、肉骨比和背膘厚度随着饲粮能量水平的增加而逐渐提高,但各组差异均不显著;另外,中能组和高能组间的眼肌面积差异不显著,但均显著高于低能组。靳光等[22]研究了不同粗饲料与玉米青贮组合对晋南牛胴体性状的影响,将45头晋南牛[体重(400±30) kg]随机分为3个组:麦秸30%+玉米青贮20%+精料50%(麦秸组,NEmf水平为6.42 MJ/kg,粗蛋白质水平为9.32%)、谷草30%+玉米青贮20%+精料50%(谷草组,NEmf水平为6.74 MJ/kg,粗蛋白质水平为9.86%)、苜蓿干草30%+玉米青贮20%+精料50%(苜蓿干草组,NEmf水平为6.78 MJ/kg,粗蛋白质水平为11.64%),饲喂187 d后屠宰,结果表明谷草组和苜蓿干草组的胴体重显著高于麦秸组,但3个组屠宰率、净肉率和肉骨比均无显著差异。上述研究表明,提高饲粮营养水平不能显著改善中国黄牛的胴体性状(如屠宰率、净肉率等),而饲粮营养水平对胴体性状的影响需协同育肥初始体重、品种遗传特性以及饲喂周期长短等因素。
和世春等[18]研究对比了放牧和强度育肥后江城牛的胴体性状,将30头江城牛随机分为3个组,分别饲喂精料补充料(2 kg/d,粗蛋白质水平为18.0%)、浓缩料(1 kg/d,粗蛋白质水平为35.0%)以及在人工草地(以臂形草为主)上放牧,精料组和浓缩料组自由采食全株玉米青贮,饲喂100 d后屠宰,研究显示精料组的屠宰率、净肉率和胴体产肉率均最高,放牧组最低,且精料组和浓缩料组的屠宰率和净肉率显著高于放牧组,3个组的胴体产肉率无显著差异,表明江城牛经过强度育肥后有更高的产肉性能。
栓系和散栏饲养对不同品种中国黄牛胴体性状和育肥性能的影响有所不同。苏楠等[23]研究了栓系和散栏2种饲养方式对新疆褐牛(30月龄)胴体性状的影响,结果显示栓系组净肉重显著低于散栏组,屠宰率、净肉率和肉骨比极显著低于散栏组。
张涛[24]统计了2019—2022年期间栓系饲养和散栏饲养新疆褐牛(栓系饲养7 539~7 611头,散栏饲养1 088~1 091头)和中国西门塔尔牛(栓系饲养2 914~2 907头,散栏饲养351~348头)的日增重和总增重,结果表明散栏饲养的新疆褐牛和中国西门塔尔牛的日增重、总增重相较于栓系饲养均有所提高,但新疆褐牛的日增重和总增重均无显著差异,而散栏饲养的中国西门塔尔牛日增重显著高于栓系饲养。

1.3 同品种不同时期中国黄牛胴体性状对比

王秀娟等[25]对比了2023年平凉红牛(24~36月龄)和2023年之前的测定结果,2023年强度育肥后平凉红牛的体重相较于2019年(22~29月龄)增加6.96%,胴体重、净肉重、净肉重相较于2010年(24月龄)分别增加长12.61%、19.16%、3.82%,肉骨比相较于2012年(22~27月龄)增加27.96%。农胜虎等[26]对比了2018年及2006年文山牛的胴体性状,结果显示文山牛胴体重(306.85 kg vs.136.3 kg)、屠宰率(60.23% vs.44.9%)和净肉率(49.78% vs.36.8%)均比2006年有所增加。王华等[16]对2024年和2006年思南牛的胴体性状进行对比,结果表明2024年思南牛平均胴体重182.81 kg,屠宰率55.98%,净肉率47.25%,肉骨比5.77,与2006年相比均有所增加,2006年思南牛胴体重、屠宰率、净肉率和肉骨比分别为153.1 kg、53.6%、44.2%和5.4。以上结果表明,无论是培育品种还是地方品种,中国黄牛的胴体性状均有较大提高,不仅表明选育有显著成效,而且表明随着养殖规模扩大和育肥技术提高,地方品种产肉潜力也在不断提高。

2 影响中国黄牛育肥性能的分子生物学机制

华西牛是中国农业科学院北京畜牧兽医研究所通过40多年不断改良的肉牛新品种[7],是以西门塔尔牛为主,夏洛莱牛、三河牛和蒙古牛杂交培育而成[27],并于2021年经农业农村部审定通过成为我国首个自主培育的肉牛新品种。Liu等[28]对202头华西牛5个发育阶段(出生、6月龄、12月龄、18月龄和24月龄)的体重、眼肌面积和背膘厚度进行了全基因组关联研究(genome-wide association studies,GWAS),同时还分析了10头青年牛(出生后320~403 d)和10头成年牛(出生后900~930 d)背最长肌的公开转录组测序(RNA-Seq)数据,鉴定出关键基因单酰基甘油脂肪酶(monoacylglycerol lipase,MGLL)、鞘磷脂合酶1(sphingomyelin synthase 1,SGMS1)、分选连接蛋白29(sorting nexin 29,SNX29)和A激酶锚定蛋白6(A-kinase anchoring protein 6,AKAP6)与脂质代谢、脂肪生成和肌肉生长相关。其中,MGLL基因编码α/β水解酶超家族(α/β hydrolase superfamily)的丝氨酸水解酶,催化单酰基甘油酯转化为游离脂肪酸和甘油。已有研究表明MGLL基因与天祝白牦牛背最长肌肌内脂肪沉积相关,经KEGG通路分析MGLL基因富集在脂肪细胞中脂肪分解的调节通路和叉头框蛋白O(FoxO)信号通路[29];AKAP6基因是成肌细胞分化、肌管形成和肌肉再生的重要调节因子,它通过肌细胞增强因子2A(myocyte enhancer factor 2A,MEF2A)促进肌细胞生成素表达,而肌细胞生成素通过直接结合AKAP6启动子上调AKAP6表达[30]。
平凉红牛是以甘肃省平凉市当地黄牛为母本,与秦川牛、南德温牛、利木赞牛经过多年杂交选育改良而成,2008年成功注册为全国首例活牛类证明商标[31],平凉红牛具有适应性强、耐粗饲、大理石纹明显的特点,能提供优质高档牛肉[32]。Wang等[33]对10头平凉红牛阉牛的半膜肌、背最长肌、肱三头肌和腰大肌的RNA-Seq数据进行了分析,筛选出21个候选基因参与了肌肉质量和个体发育,其中真核翻译延长因子1α(eukaryotic translation elongation factor 1 alpha,EEF1A)对蛋白质合成至关重要,它存在2种不同的亚型(EEF1A1和EEF1A2),EEF1A2是调节与肌生成相关的基因,有研究显示与乌珠穆沁羊(低肌肉高脂肪)相比,EEF1A2基因在特塞尔羊(高肌肉低脂肪)中过表达[34];苯丙酰-tRNA合成酶亚基β(phenylalanyl-tRNA synthetase subunit beta,FARSB)在RNA结合和蛋白质生物合成中发挥积极作用,是1个与坎钦牛断奶体重相关且有潜力的候选基因[35];线粒体在肌肉质量的调节过程中起着重要作用,线粒体的质量控制系统可识别并解决线粒体功能障碍,这对骨骼肌质量维持至关重要[36],在这个调控系统中,PTEN诱导的激酶1(PTEN-induced kinase 1,PINK1)是1种丝氨酸/苏氨酸激酶,在线粒体损伤情况下被激活,电压依赖性阴离子通道1(voltage dependent anion channel 1,VDAC1)通过线粒体钙单转运通道促进线粒体对钙的摄取,负责调节线粒体自噬和细胞凋亡[37];线粒体外膜转位酶70(translocase of outer mitochondrial membrane 70,TOMM70)和PINK1相互作用作为初始受体参与线粒体前提蛋白质的输入,此外TOMM70还通过调控钙离子(Ca2+)的转运效率影响线粒体能量代谢和氧化应激水平,从而对线粒体的质量控制发挥重要作用[38]。
鲁西牛是我国5大黄牛地方良种之一,具有肉质鲜美、肌内脂肪均匀沉积、大理石纹明显的优点[39]。Hu等[40]对山东核心保种场的16头鲁西牛进行了全基因组测序(whole-genome sequencing,WGS),并将174个已发表的牛基因组数据进行联合分析,鉴定出与鲁西牛肌肉生长分化、生长性状相关的候选基因:胞质分裂作用因子3(dedicator of cytokinesis 3,DOCK3)、Ral GTP酶激活蛋白催化亚基α1(Ral GTPase activating protein catalytic subunit alpha 1,RALGAPA1)、生长激素受体(growth hormone receptor,GHR)、DAZ相关蛋白1(DAZ-associated protein 1,DAZAP1)、介质复合物亚基13(mediator complex subunit 13,MED13)、PLAG1锌指蛋白(PLAG1 zinc finger,PLAG1)和肝配蛋白A2(ephrin A2,EFNA2);其中,DOCK3基因调节正常的骨骼肌再生和葡萄糖代谢,敲除DOCK3基因后会导致小鼠肌肉结构受损,运动活动降低,肌纤维再生受损和代谢功能障碍[41];DAZAP1基因是RNA结合蛋白,对小鼠正常生长、发育和生育能力有重要作用[42];PLAG1基因编码锌指蛋白,已有研究证明该基因与内洛尔牛的出生体重相关[43]。
郏县红牛属于肉役兼用型牛,是我国著名的地方品种之一,于2006年被列入国家品种资源保护名录,其肉质细嫩多汁,大理石纹明显,具有生产雪花牛肉的潜力[44]。Xia等[45]对核心育种场的30头郏县红牛的全基因组测序数据以及已发表的131头牛的基因组数据进行联合分析,研究发现有2个与肌肉发育相关的基因:磷酸酶和肌动蛋白调节因子1(phosphatase and actin regulator 1,PHACTR1)是磷酸酶和肌动蛋白调节家族的成员,参与调节肌动蛋白细胞骨架的重组,是纤维肌发育不良的第一个遗传易感性位点[46];EYA转录辅激活因子和磷酸酶(EYA-transcriptional coactivator and phosphatase,EYA)基因家族具有蛋白磷酸酶功能,其酶活性对调控编码生长控制和信号分子的基因、调节前体细胞增殖至关重要[47],它与肌肉的正常发育有关,EYA3基因敲除小鼠表现出运动减弱[48]。
昭通牛因主产于云南省昭通市而得名,属役肉兼用型黄牛[49],是昭通市饲养的古老畜种之一,不仅具有优良的肉质,而且还具有耐热耐湿性。Lv等[50]以20头昭通牛为研究对象对全基因组重测序数据进行分析,研究发现与骨发育和代谢相关的基因包括碳酸酐酶10(carbonic anhydrase 10,CA10)、γ-氨基丁酸A型受体亚基γ3(gamma-aminobutyric acid type a receptor subunit gamma 3,GABRG3)、胶质介素(gliomedin,GLDN)、棕榈油酰蛋白羧酸酯酶(notum, palmitoleoyl-protein carboxylesterase,NOTUM)和淋巴增强子结合因子1(lymphoid enhancer binding factor 1,LEF1);其中,CA10基因属于碳酸酐酶家族的锌金属酶,参与骨骼矿物质的溶解和骨吸收[51],LEF1基因是Wnt/Lrp5/β-连环蛋白信号级联的转录效应器,调节成骨细胞分化、骨密度和骨骼强度[52]。与肌肉发育相关的基因包括富含亮氨酸重复序列和Ig结构域蛋白2(leucine rich repeat and Ig domain containing 2,LINGO2)、肌球蛋白VB(myosin VB,MYO5B);其中,MYO5B基因作为1种V类肌球蛋白,在内吞体回收进入质膜过程中起着关键作用[53],MYO5B基因还对于肌肉和脂肪细胞中葡萄糖转运蛋白4(glucose transporter type 4,GLUT4)的转位至关重要[54]。
文山牛属肉用型黄牛地方品种,是中国现存单体存栏量最大、遗传基因最为完整且品种独特的地方黄牛品种[55]。任洪辉[56]对1 060头文山牛的17个体尺性状进行全基因组关联分析,筛选到24个候选基因与骨稳态和体节发育等生物学过程相关,其中HIVEP锌指3(HIVEP zinc finger 3,HIVEP3)也称为Schnurri-3(Shn3),它是一种锌指蛋白可,通过控制RUNX家族转录因子2(RUNX family transcription factor 2,RUNX2)的蛋白水平来调节骨量,而RUNX2是成骨细胞分化的主要转录调节因子[57];乙酰辅酶A合成酶(acetoacetyl-CoA synthetase,AACS)是一种酮体利用酶,能在胞质溶胶中将乙酰乙酸转化成乙酰辅酶A,提供乙酰基单元作为脂肪生成的前体,在脂肪细胞分化中起关键作用[58];卷曲螺旋结构域蛋白92(coiled-coil domain containing 92,CCDC92)和动力蛋白轴丝重链10(dynein axonemal heavy chain 10,DNAH10)参与脂肪细胞功能和分化[59]。
临沧高峰牛是云南省高峰牛群之一,是较为原始的牛品种,具有优越的耐热性和抗逆性,Sun等[60]利用全基因组重测序对22头临沧驼峰牛的基因组多样性进行研究,结果显示与骨骼肌发育相关的基因有转录因子12(transcription factor 12,TCF12)、SUMO特异性肽酶2(SUMO specific peptidase 2,SENP2)、驱动蛋白家族成员1C(kinesin family member 1C,KIF1C)和前纤维蛋白1(profilin 1,PFN1);其中,TCF12是肌肉干细胞染色质重塑的关键调控因子,其既能调控肌肉细胞的特化过程,又参与骨骼肌的发育与再生[61],是骨骼肌中的一个强效促分化因子[62];SENP2是一种蛋白质编码基因,可将新合成的小泛素样修饰剂1(small ubiquitin like modifier 1,SUMO1)加工成可结合形式,是骨骼肌脂肪酸代谢的重要调节因子,在调节肌肉生长抑制因子表达和肌肉生成中起重要作用[63];KIF1C在肌肉生成和成年肌肉中起着维持膜循环的作用[64];PFN1是控制肌动蛋白纤维结构的关键参与者,它参与迁移等许多细胞活动,是骨骼发育的关键因子[65]。
Mei等[66]对闽南牛(n=13)和蒙古牛(n=13)利用全基因组测序筛选出与肉产量相关的潜在选择基因,在闽南牛中这些基因包括溶质载体家族38成员3(solute carrier family 38 member 3,SLC38A3)、外泌体复合物组分3(exocyst complex component 3,EXOC3)、信号转导及转录激活因子5B(signal transducer and activator of transcription 5B,STAT5B)、核膜蛋白(emerin,EMD)和肾上腺素能受体β2(adrenoceptor beta 2,ADRB2)等;其中,SLC38A3、EXOC3、STAT5B、EMD也被作为日本本地牛种Kuchinoshima-Ushi全基因组重测序中肉类性状的候选基因[67];ADRB2编码的β-肾上腺素能受体在调节代谢作用,特别是脂肪分解、胰岛素抗性和能量平衡中发挥重要作用,其多态性对秦川牛选择的胴体性状有影响[68]。在蒙古牛中与肉产量相关的潜在选择基因包括Hes相关家族BHLH转录因子含YRPW模体类似物(Hes related family BHLH transcription factor with YRPW motif like,HEYL)、溶质载体家族29成员1(solute carrier family 29 member 1,SLC29A1)、釉原蛋白相互作用蛋白11(tuftelin interacting protein 11,TFIP11)、天冬酰胺合成酶(asparagine synthetase,ASNS)、核糖体蛋白L15(ribosomal protein L15,RPL15)、视黄醇脱氢酶5(retinol dehydrogenase 5,RDH5)、肌醇1,4,5-三磷酸受体3型(inositol 1,4,5-trisphosphate receptor type 3,ITPR3)、载脂蛋白F(apolipoprotein F,APOF)和核受体共激活因子1(nuclear receptor coactivator 1,NCOA1)等;其中,HEYL、SLC29A1、TFIP11、ASNS、RPL15、RDH5、ITPR3同样也作为日本本地牛种Kuchinoshima-Ushi肉类性状的候选基因[67];APOF在脂质代谢中具有关键功能,NCOA1被认为是牛骨骼和肌肉发育、繁殖、胚胎和器官发育的候选基因[69]。
Liu等[70]对8个中国品种201头牛[凉山牛(n=22)、南丹牛(n=23)、平武牛(n=23)、文山牛(n=21)、昭通牛(n=23)、海南牛(n=26)、雷琼(牛n=30)、陆丰牛(n=33)]使用Illumina牛高密度单核苷酸多态性(high density-single nucleotide polymorphism,HD-SNP)阵列对所有个体进行基因分型,并利用整合单倍型分数(integrated haplotype score,iHS)方法检测到钙结合素2(calsyntenin 2,CLSTN2)、二氢嘧啶脱氢酶(dihydropyrimidine dehydrogenase,DPYD)和硫酸软骨素合酶3(chondroitin sulfate synthase 3,CHSY3)与陆丰牛肉质和脂肪酸有关,CLSTN2参与突触组装和突触传递的正调节,但也有研究表明CLSTN2在肥胖小鼠的网膜脂肪组织和皮下脂肪组织中均未显著上调[71];DPYD编码的蛋白质是一种嘧啶分解代谢酶,该基因的表达量与韩牛大理石纹脂肪含量呈正相关[72],CHSY3编码一种与高尔基体相关的跨膜蛋白,该基因与肌肉蒸煮损失相关[73]。

3 小结与展望

中国黄牛育肥性能受到品种、地域、营养水平以及饲喂方式等的影响;虽然提高饲粮营养水平不能显著改善中国黄牛的胴体性状(如屠宰率、净肉率等),但选育和养殖技术的进步能显著提升中国黄牛的产肉潜力;分子生物学机制研究发现多个关键基因(如EEF1A2、MGLL等)与肌肉生长、脂肪沉积等相关。综上所述,品种选育、精准饲喂及遗传改良可提升中国黄牛育肥性能,未来研究应聚焦于基因调控机制的深入解析,结合基因组选择技术优化育种策略,同时推动遗传资源保护以及中国黄牛产业高质量发展。

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