转录组数据揭示滩羊肌内脂肪沉积相关lncRNAs及其相关基因与通路

doi:10.7668/hbnxb.20195563

摘要/Abstract

摘要： 肌内脂肪沉积作为影响肉品质(如嫩度、风味和多汁性)的关键因素,其调控机制与长链非编码 RNA(lncRNA)对脂肪生成和脂质代谢的调节密切相关。基于前期研究发现,滩羊肌内脂肪含量显著高于杜泊羊和小尾寒羊,以这3个绵羊品种的背最长肌为研究对象,通过转录组测序筛选差异表达 mRNAs(DE mRNAs)和 lncRNAs(DE lncRNAs)及其相关通路,旨在揭示滩羊高肌内脂肪沉积的分子机制。结果表明:在小尾寒羊和滩羊组筛选出836个DE mRNAs,832个DE lncRNAs,杜泊羊和滩羊组筛选出578个DE mRNAs,700个DE lncRNAs。两组分别取交集共得到226 DE mRNAs,308个DE lncRNAs,通过对308个DE lncRNAs进行靶基因预测,共预测到1 475个靶基因,其中预测到co_expression(共表达)靶基因687个,co_localization(共定位)靶基因有788个;通过对DE mRNAs进行GO和KEGG富集发现,其中与脂肪沉积相关的通路有肌动蛋白细胞骨架调节、催乳素信号通路等,其中FGF14、FGF9等基因被报道参与动物的脂肪代谢过程。通过对预测到的共表达靶基因和共定位靶基因进行GO和KEGG富集分析,发现它们富集到与脂肪代谢的相关通路,如cAMP信号通路、催乳素信号通路、胰岛素信号通路等;通过对通路和功能分析进一步筛选出和脂肪代谢相关的CREBBP、ATP1B3、PPP1R3C、PIK3R1等基因,这些基因在绵羊肌内脂肪沉积方面可能发挥重要调控作用。随机选择的6个DE mRNAs和6个DE lncRNAs的qRT-PCR结果与转录组测序结果趋势一致。

关键词: 滩羊, 肌内脂肪沉积, 转录组, lncRNA

Abstract:

Intramuscular fat deposition,as a key factor affecting meat quality(e.g.tenderness,flavor,and juiciness),is closely related to the regulation of lipogenesis and lipid metabolism by long non-coding RNA(lncRNA).Based on the previous finding that the intramuscular fat content of Tan sheep was significantly higher than that of Dorper sheep and Small-tailed Han sheep,the dorsal longest muscle of these three sheep breeds was used as the target,and differentially expressed mRNAs(DE mRNAs)and lncRNAs(DE lncRNAs)and their related pathways were screened by transcriptome sequencing,with the aim of revealing molecular mechanism of high intramuscular fat deposition in Tan sheep.The results showed:836 DE mRNAs and 832 DE lncRNAs were identified in the Small-tailed Han sheep vs.Tan sheep group;578 DE mRNAs and 700 DE lncRNAs were screened in the Dorper sheep vs.Tan sheep group.The intersection of the two groups fielded a total of 226 DE mRNAs and 308 DE lncRNAs.Target gene prediction for the 308 DE lncRNAs identified 1 475 potential targets,including 687 co_expression(co-expressed)target genes and 788 co_localization(co-localized)target genes.GO and KEGG enrichment analysis of DE mRNAs revealed fat deposition-related pathways such as regulation of actin cytoskeleton and prolactin signaling pathway,with genes like FGF14 and FGF9 were reported to participate in animal fat metabolism.Enrichment analysis of co-expressed and co-localized target genes identified fat metabolism-related pathways,including the cAMP signaling pathway,prolactin signaling pathway,and insulin signaling pathway.Further functional analysis screened key genes associated with fat metabolism(CREBBP,ATP1B3,PPP1R3C, and PIK3R1),which may play crucial regulatory roles in ovine intramuscular fat deposition.The qRT-PCR results of 6 randomly selected DE mRNAs and 6 randomly selected DE lncRNAs were consistent with transcriptome sequencing data.

Key words: Tan sheep, Intramuscular fat deposition, Transcriptome, lncRNA

中图分类号:

S826

王丁香, 李承隆, 郭晨浩, 冯登侦, 康晓龙. 转录组数据揭示滩羊肌内脂肪沉积相关lncRNAs及其相关基因与通路[J]. 华北农学报, 2025, 40(6): 202-212. doi: 10.7668/hbnxb.20195563.

WANG Dingxiang, LI Chenglong, GUO Chenhao, FENG Dengzhen, KANG Xiaolong. Transcriptomic Data Reveal Sheep Intramuscular Fat Deposition-Associated LncRNAs and Their Associated Genes and Pathways[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(6): 202-212. doi: 10.7668/hbnxb.20195563.

http://www.hbnxb.net/CN/Y2025/V40/I6/202

图/表 6

表1 mRNA和lncRNA引物信息

Tab.1 Primer information of mRNA and lncRNA

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	产物长度/bp Product length	退火温度/℃ Tm
PRKAA2	F:AGCAGAGGTCTGGTTCCTCA	158	60
	R:AGGTGAGACAGAGGACGACA
FGF9	F:AGTGGACTCTACCTCGGCAT	158	60
	R:ACGTAGAACCTCCGTCCAGT
FGF2	F:ACTTTAAGGACCCCAAGCGG	170	60
	R:ACGGTTTGCACACACTCCTT
TNC	F:ATGGGGAGATACGGGGACAA	115	60
	R:TTCGGAAGTTGCTGGGTCTC
CRH	F:CTTCCTCCGCCTGGGTAAC	103	60
	R:CTTGTCCGGAGAAAGGCGAC
CASR	F:CTTTGCTGTGCTGGGCATTT	125	60
	R:AGAGCAGGGAGAAGAGGAGG
GAPDH	F:TCGGAGTGAACGGATTCGG	162	60
	R:TGATGACGAGCTTCCCGTTC
TCONS_00033873	F:GTGCCGCCTCTGTGAACTTG	143	60
	R:GGTGACTGCTCTTCTGCTTTCTG
TCONS_00128473	F:TCAGACACAACTTAGCGACTACAC	83	60
	R:AAGCCTCCACCACCTCCAAC
TCONS_00112422	F:GGCTACCTGATGCGAAGAACTG	100	60
	R:CATCCAACCATCTCATCCTCTGTC
TCONS_00153755	F:TCTGGCTTTGTTTGAAGGGTTGTG	75	60
	R:CACCACCACTCCGAGGAATCC
TCONS_00193487	F:CCTGAGTGACTGAACTGAACTGAAG	135	60
	R:GGAAGGACAGATGATGAAGCAGAC
TCONS_00150301	F:TTGTGATTCGCTCTCCTCTTGTTG	109	60
	R:ATTGCTGCTTCTGCCGTTACTG

表2 3个品种绵羊肌内脂肪含量

Tab.2 Intramuscular fat content in three sheep breeds

品种 Breeds	肌内脂肪/% Intramuscular fat content
杜泊羊D	3.41±0.41b
滩羊 T	5.42±2.35a
小尾寒羊XH	2.09±0.82b

图1 DE mRNAs 和 DE lncRNAs的韦恩图和火山图 A、D.TvsH&TvsD 2组的DE mRNAs、DE lncRNAs的韦恩图;B、C分别为TvsD组和TvsH组 DE mRNAs的火山图;图E、F分别为TvsD组和TvsH组DE lncRNAs的火山图。

Fig.1 Venn and volcano plots of DE mRNAs and DE lncRNAs A,D show the Venn diagram of DE mRNAs and DE lncRNAs of the two groups TvsH & TvsD;B and C.Volcano plots of DE mRNAs from the TvsD and TvsH groups;E and F.Volcano plots of DE lncRNAs from the TvsD and TvsH groups.

图2 差异表达mRNAs的 GO和KEGG富集分析 A.DE mRNAs GO富集分析:1.染色质沉默,2.蛋白质去乙酰化,3.蛋白质脱酰基化,4.基因表达的负调控表观遗传学,5.大分子脱酰基化,6.膜融合,7.膜组织,8.烟酰胺代谢过程,9.烟酰胺生物合成过程,10.真核翻译起始因子2复合物,11.病毒膜,12.纤毛基部,13.生长因子活性,14.受体结合,15.分子转导剂活性,16.蛋白质结合,17.细胞外配体门控离子通道活性,18.组蛋白去乙酰化酶活性,19.NAD依赖的组蛋白去乙酰化酶活性,20.去乙酰化酶活性,21.蛋白质去乙酰化酶活性,22.NAD依赖的蛋白质去乙酰化酶活性,23.水解酶活性,作用于碳氮键(而不是肽),在线性酰胺中,24.生长因子受体结合,25.细胞因子受体结合,26.磷脂酰肌醇磷酸酯激酶活性,27.受体活性,28.烟酰胺合成酶活性,29.配体门控离子通道活性,30.配体门控通道活性;B.DE mRNAs KEGG富集分析:1.刺猬信号通路,2.吗啡成瘾,3.昼夜节律,4.昼夜节律调控,5.逆行内分泌素信号传导,6.尼古丁成瘾,7.细菌侵袭上皮细胞,8.GABA能突触,9.神经活性配体-受体相互作用,10.长时程抑制,11.黑色素生成,12.粘蛋白型O-聚糖生物合成,13.黑色素瘤,14.谷氨酸能突触,15.卵母细胞减数分裂,16.非洲锥虫病,17.肌动蛋白细胞骨架调节,18.HTLV-I感染,19.转化生长因子-β信号通路,20.催乳素信号通路。

Fig.2 GO and KEGG enrichment analysis of DE mRNAs A.GO enrichment analysis of DE mRNAs:1.Chromatin silencing,2.Protein deacetylation,3.Protein deacylation,4.Negative regulation of gene expression,epigenetic,5.Macromolecule deacylation,6.Membrane fusion,7.Membrane organization,8.Nicotianamine metabolic process,9.Nicotianamine biosynthetic process,10.Eukaryotic translation initiation factor 2 complex,11.Virion membrane,12.Ciliary base,13.Growth factor activity,14.Receptor binding,15.Molecular transducer activity,16.Protein binding,17.Extracellular ligand-gated ion channel activity,18.Histone deacetylase activity,19.NAD-dependent histone deacetylase activity,20.Deacetylase activity,21.Protein deacetylase activity,22.NAD-dependent protein deacetylase activity,23.Hydrolase activity,acting on carbon-nitrogen(but not peptide)bonds,in linear amides,24.Growth factor receptor binding,25.Cytokine receptor binding,26.Phosphatidylinositol phosphate kinase activity,27.Receptor activity,28.Nicotianamine synthase activity,29.Ligand-gated ion channel activity,30.Ligand-gated channel activity;B.KEGG enrichment analysis of DE mRNAs:1.Hedgehog signaling pathway,2.Morphine addiction,3.Circadian rhythm,4.Circadian entrainment,5.Retrograde endocannabinoid signaling,6.Nicotine addiction,7.Bacterial invasion of epithelial cells,8.GABA ergic synapse,9.Neuroactive ligand-receptor interaction,10.Long-term depression,11.Melanogenesis,12.Mucin type O-Glycan biosynthesis,13.Melanoma,14.Glutamatergic synapse,15.Oocyte meiosis,16.African trypanosomiasis,17.Regulation of actin cytoskeleton,18.HTLV-I infection,19.TGF-beta signaling pathway,20.Prolactin signaling pathway.

图3 差异表达lncRNAs的GO和KEGG富集分析 A.DE lncRNAs的共表达靶基因GO富集分析:1.胺生物合成过程,2.细胞生物胺生物合成过程,3.多胺代谢过程,4.多胺生物合成过程,5.细胞生物胺代谢过程,6.细胞胺代谢过程,7.铵离子代谢过程,8.精胺生物合成过程,9.精胺代谢过程,10.免疫系统过程,11.骨骼系统发育,12.软骨细胞分化,13.软骨细胞增殖,14.软骨发育,15.结缔组织发育,16.再生,17.组织再生,18.发育生长,19.免疫反应,20.胺代谢过程,21.rRNA加工,22.rRNA代谢过程,23.对真菌的反应,24.藻胆体,25.质膜蛋白复合物,26.三价铁结合,27.细胞因子活性,28.G蛋白偶联受体结合,29.趋化因子活性,30.趋化因子受体结合;B.DE lncRNAs的共表达靶基因KEGG富集分析:1.矿物质吸收,2.真核生物核糖体生物发生,3.甲型流感,4.沙门氏菌感染,5.军团病,6.阿米巴病,7.类风湿关节炎,8.胆汁分泌,9.NOD样受体信号通路,10.核糖体,11.单纯疱疹感染,12.心肌收缩,13.朊病毒病,14.酪氨酸代谢,15.cAMP信号通路,16.药物代谢-其他酶,17.半胱氨酸和蛋氨酸代谢,18.色氨酸代谢,19.心肌细胞肾上腺素能信号传导,20.醚脂代谢; C.DE lncRNAs的共定位靶基因GO富集分析:1.一价无机阳离子运输,2.细胞含氮化合物生物合成过程,3.钾离子运输,4.芳香族化合物生物合成过程,5.杂环生物合成过程,6.核转录mRNA分解代谢过程,无义介导的衰变,7.有机环状化合物生物合成过程,8.无机阳离子跨膜运输,9.细胞含氮化合物代谢过程,10.无机离子跨膜运输,11.大分子生物合成过程,12.含核碱基化合物生物合成过程,13.DNA转录模板,14.核酸转录模板,15.DNA转录调控模板,16.核酸转录调控模板,17.细胞大分子生物合成过程,18.基因表达,19.RNA生物合成过程,20.阳离子跨膜运输,21.质子运输双区ATP酶复合物,催化结构域,22.细胞内,23.细胞,24.细胞部分,25.磷脂酶抑制剂活性,26.脂肪酶抑制剂活性,27.内向整流钾通道活性,28.核酸结合,29.钾离子跨膜转运体活性,30.电压门控钾通道活性;D.DE lncRNAs的共定位靶基因KEGG富集分析:1.抗原加工和呈递,2.RIG-I样受体信号通路,3.破骨细胞分化,4.血小板活化,5.味觉转导,6.细胞黏附分子(CAMs),7.核糖体,8.子宫内膜癌,9.结核病,10.胰岛素信号通路,11.乙醛酸和二羧酸代谢,12.单纯疱疹感染,13.甲型流感,14.GnRH信号通路,15.胃酸分泌,16.甲状腺癌,17.神经营养因子信号通路,18.昼夜节律,19.催乳素信号通路,20.丙型肝炎。

Fig.3 GO and KEGG enrichment analysis of DE lncRNAs A.GO enrichment analysis of co-expressed target genes for DE lncRNAs:1.Amine biosynthetic process,2.Cellular biogenic amine biosynthetic process,3.Polyamine metabolic process,4.Polyamine biosynthetic process,5.Cellular biogenic amine metabolic process,6.Cellular amine metabolic process,7.Ammonium ion metabolic process,8.Spermine biosynthetic process,9.Spermine metabolic process,10.Immune system process,11.Skeletal system development,12.Chondrocyte differentiation,13.Chondrocyte proliferation,14.Cartilage development,15.Connective tissue development,16.Regeneration,17.Tissue regeneration,18.Developmental growth,19.Immune response,20.Amine metabolic process,21.rRNA processing,22.rRNA metabolic process,23.Response to fungus,24.Phycobilisome,25.Plasma membrane protein complex,26.Ferric iron binding,27.Cytokine activity,28.G-protein coupled receptor binding,29.Chemokine activity,30.Chemokine receptor binding;B.KEGG enrichment analysis of co-expressed target genes for DE lncRNAs:1.Mineral absorption,2.Ribosome biogenesis in eukaryotes,3.Influenza A,4.Salmonella infection,5.Legionellosis,6.Amoebiasis,7.Rheumatoid arthritis,8.Bile secretion,9.NOD-like receptor signaling pathway,10.Ribosome,11.Herpes simplex infection,12.Cardiac muscle contraction,13.Prion diseases,14.Tyrosine metabolism,15.cAMP signaling pathway,16.Drug metabolism-other enzymes,17.Cysteine and methionine metabolism,18.Tryptophan metabolism,19.Adrenergic signaling in cardiomyocytes 20.Ether lipid metabolism;C.GO enrichment analysis of co-located target genes for DE lncRNAs:1.Monovalent inorganic cation transport,2.Cellular nitrogen compound biosynthetic process,3.Potassium ion transport,4.Aromatic compound biosynthetic process,5.Heterocycle biosynthetic process,6.Nuclear-transcribed mRNA catabolic process,nonsense-mediated decay,7.Organic cyclic compound biosynthetic process,8.Inorganic cation transmembrane transport,9.Cellular nitrogen compound metabolic process,10.Inorganic ion transmembrane transport,11.Macromolecule biosynthetic process,12.Nucleobase-containing compound biosynthetic process,13.Transcription,DNA-templated,14.Nucleic acid-templated transcription,15.Regulation of transcription,DNA-templated,16.Regulation of nucleic acid-templated transcription,17.Cellular macromolecule biosynthetic process,18.Gene expression,19.RNA biosynthetic process,20.Cation transmembrane transport,21.Proton-transporting two-sector ATPase complex,catalytic domain,22.Intracellular,23.Cell,24.Cell part,25.Phospholipase inhibitor activity,26.Lipase inhibitor activity,27.Inward rectifier potassium channel activity,28.Nucleic acid binding,29.Potassium ion transmembrane transporter activity,30.Voltage-gated potassium channel activity;D.KEGG enrichment analysis of co-located target genes for DE lncRNAs:1.Antigen processing and presentation,2.RIG-I-like receptor signaling pathway,3.Osteoclast differentiation,4.Platelet activation,5.Taste transduction,6.Cell adhesion molecules(CAMs),7.Ribosome,8.Endometrial cancer,9.Tuberculosis,10.Insulin signaling pathway,11.Glyoxylate and dicarboxylate metabolism,12.Herpes simplex infection,13.Influenza A,14.GnRH signaling pathway,15.Gastric acid secretion,16.Thyroid cancer,17.Neurotrophin signaling pathway,18.Circadian rhythm,19.Prolactin signaling pathway,20.Hepatitis C.

图4 DE mRNAs和DE lncRNAs的qRT-PCR验证

Fig.4 qRT-PCR validation of DE mRNAs and DE lncRNAs

参考文献 43

[1]	Gao X G, Wang Z Y, Miao J, Xie L, Dai Y, Li X M, Chen Y, Luo H L, Dai R T. Influence of different production strategies on the stability of color,oxygen consumption and metmyoglobin reducing activity of meat from Ningxia Tan sheep[J]. Meat Science, 2014, 96(2):769-774.doi:10.1016/j.meatsci.2013.09.026. URL
[2]	Zhang C, Zhang H, Liu M, Zhao X G, Luo H L. Effect of breed on the volatile compound precursors and odor profile attributes of lamb meat[J]. Foods, 2020, 9(9):1178.doi:10.3390/foods9091178. URL
[3]	EEr H, Ma L N, Xie X L, Ma J F, Ma X M, Yue C J, Ma Q, Liang X J, Ding W, Li Y K. Genetic polymorphism association analysis of SNPs on the species conservation genes of Tan sheep and Hu sheep[J]. Tropical Animal Health and Production, 2020, 52(3):915-926.doi:10.1007/s11250-019-02063-1. pmid: 32026291
[4]	Cloete S W P, Snyman M A, Herselman M J. Productive performance of dorper sheep[J]. Small Ruminant Research, 2000, 36(2):119-135.doi:10.1016/S0921-4488(99)00156-X. pmid: 10760448
[5]	Ma Y, Zheng Z, Zhang L G, Dou Y J, Bai U, Liu X, Zhang X R, Su X H, Zhang L. Effects study of early weaning time on the growth and development of dairy sheep lamb[J]. International Journal of Food Science and Agriculture, 2021, 5(3):411-420.doi:10.26855/ijfsa.2021.09.011. URL
[6]	徐红伟. 三种尾脂型绵羊脂肪沉积性状表型测定及关联基因筛选研究[D]. 兰州:甘肃农业大学, 2019. doi:10.27025/d.cnki.ggsnu.2019.000017.
	Xu H W. Research on phenotypic determination of fat deposition traits and screening of associated genes in three kinds of tail fat sheep[D]. Lanzhou: Gansu Agricultural University,2019.
[7]	Jathar S, Kumar V, Srivastava J, Tripathi V. Technological developments in lncRNA biology[J]. Advances in Experimental Medicine and Biology, 2017,1008:283-323.doi:10.1007/978-981-10-5203-3_10.
[8]	孙昊然, 李新海. 基于RNA-seq技术鉴定绵羊毛囊发育相关lncRNAs[J]. 华北农学报, 2024, 39(1):211-218.doi:10.7668/hbnxb.20194552.
	Sun H R, Li X H. Identification of Sheep hair follicles development-related lncRNAs based on RNA-seq technology[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(1):211-218. doi: 10.7668/hbnxb.20194552
[9]	Xiao C, Wei T, Liu L X, Liu J Q, Wang C X, Yuan Z Y, Ma H H, Jin H G, Zhang L C, Cao Y. Whole-transcriptome analysis of preadipocyte and adipocyte and construction of regulatory networks to investigate lipid metabolism in sheep[J]. Frontiers in Genetics, 2021,12:662143.doi:10.3389/fgene.2021.662143.
[10]	Wang Z G, Luo Z C, Dai Z, Zhong Y T, Liu X G, Zuo C Q. Long non-coding RNA lnc-OAD is required for adipocyte differentiation in 3T3-L1 preadipocytes[J]. Biochemical and Biophysical Research Communications, 2019, 511(4):753-758.doi:10.1016/j.bbrc.2019.02.133. pmid: 30833079
[11]	Yuan Z H, Ge L, Sun J Y, Zhang W B, Wang S H, Cao X K, Sun W. Integrative analysis of Iso-Seq and RNA-seq data reveals transcriptome complexity and differentially expressed transcripts in sheep tail fat[J]. PeerJ, 2021,9:e12454.doi:10.7717/peerj.12454.
[12]	Ma L, Zhang M, Jin Y Y, Erdenee S, Hu L Y, Chen H, Cai Y, Lan X Y. Comparative transcriptome profiling of mRNA and lncRNA related to tail adipose tissues of sheep[J]. Frontiers in Genetics, 2018,9:365.doi:10.3389/fgene.2018.00365.
[13]	Han F H, Li J, Zhao R R, Liu L R, Li L L, Li Q, He J N, Liu N. Identification and co-expression analysis of long noncoding RNAs and mRNAs involved in the deposition of intramuscular fat in Aohan fine-wool sheep[J]. BMC Genomics, 2021, 22(1):98.doi:10.1186/s12864-021-07385-9. pmid: 33526009
[14]	Liu T Y, Feng H, Yousuf S, Xie L L, Miao X Y. Differential regulation of mRNAs and lncRNAs related to lipid metabolism in Duolang and Small Tail Han sheep[J]. Scientific Reports, 2022,12:11157.doi:10.1038/s41598-022-15318-z.
[15]	刘远, 李文杨, 吴贤锋, 黄勤楼, 高承芳, 陈鑫珠, 张晓佩. 福清山羊与努比亚黑山羊背最长肌比较转录组分析[J]. 中国农业科学, 2019, 52(14):2525-2537.doi:10.3864/j.issn.0578-1752.2019.14.011.
	Liu Y, Li W Y, Wu X F, Huang Q L, Gao C F, Chen X Z, Zhang X P. Transcriptome analysis of differentially gene expression associated with longissimus Doris tissue in Fuqing goat and Nubian black goat[J]. Scientia Agricultura Sinica, 2019, 52(14):2525-2537.
[16]	王燕燕, 杨军祥, 徐建峰, 成伟伟, 何茂昌, 王珂, 高登伟, 谢文章. 陶湖杂交 F₁代与萨湖杂交 F₁代羔羊背最长肌转录组分析[J]. 饲料研究, 2022, 45(11):56-60.doi:10.13557/j.cnki.issn1002-2813.2022.11.013.
	Wang Y Y, Yang J X, Xu J F, Cheng W W, He M C, Wang K, Gao D W, Xie W Z. Transcriptome analysis of longissimus dorsi muscle of Taohu F₁ and Sahu F₁ lambs[J]. Feed Research, 2022, 45(11):56-60.
[17]	Arora R, Naveen Kumar S, Sudarshan S, Fairoze M N, Kaur M, Sharma A, Girdhar Y, Sreesujatha R M, Devatkal S K, Ahlawat S, Vijh R K, Manjunatha S S. Transcriptome profiling of longissimus thoracis muscles identifies highly connected differentially expressed genes in meat type sheep of India[J]. PLoS One, 2019, 14(6):e0217461.doi:10.1371/journal.pone.0217461. URL
[18]	Noce A. Analyzing the genetic variation of dairy Sarda and Spanish meat ovine breeds[D]. Italy: University of Sassari, 2016.
[19]	Shamsi F, Xue R D, Huang T L, et al. FGF6 and FGF9 regulate UCP1 expression independent of brown adipogenesis[J]. Nature Communications, 2020,11:1421.doi:10.1038/s41467-020-15055-9.
[20]	黄凯. FGF9对山羊肌内脂肪细胞分化的调控作用[D].成都:西南民族大学, 2020.doi:10.27417/d.cnki.gxnmc.2020.000210.
	Huang K. Regulation of FGF9 on the differentiation of goat intramuscular adipocytes[D]. Chengdu: Southwest University for Nationalities,2020.
[21]	Onzima R B, Upadhyay M R, Doekes H P, Brito L F, Bosse M, Kanis E, Groenen M A M, Crooijmans R P M A. Genome-wide characterization of selection signatures and runs of homozygosity in Ugandan goat breeds[J]. Frontiers in Genetics, 2018,9:318.doi:10.3389/fgene.2018.00318.
[22]	包荣坤. 番茄红素通过调控脂质代谢拮抗DEHP致肝脏毒性的研究[D]. 哈尔滨: 东北农业大学, 2021.doi:10.27010/d.cnki.gdbnu.2021.000168.
	Bao R K. Lycopene antagonizes DEHP induced hepatotoxicity by regulating lipid metabolism[D].Harbin: Northeast Agricultural University,2021.
[23]	Yang Y Y, Han L, Yu Q L, Gao Y F, Song R D. Study of the AMP-activated protein kinase role in energy metabolism changes during the postmortem aging of yak longissimus dorsal[J]. Animals, 2020, 10(3):427.doi:10.3390/ani10030427. URL
[24]	Kong D L, Cui J J, Fu J C. DbcAMP regulates adipogenesis in sheep inguinal preadipocytes[J]. Lipids in Health and Disease, 2017, 16(1):93.doi:10.1186/s12944-017-0478-6. pmid: 28526050
[25]	Ravnskjaer K, Madiraju A, Montminy M. Role of the cAMP pathway in glucose and lipid metabolism[M]// Metabolic Control.Cham: Spinger International Publishing,2015:29-49.doi:10.1007/164_2015_32.
[26]	Moon Y, Tong T, Kang W, Park T. Filbertone ameliorates adiposity in mice fed a high-fat diet via activation of cAMP signaling[J]. Nutrients, 2019, 11(8):1749.doi:10.3390/nu11081749. URL
[27]	梁鹏, 张稳, 冯登侦, 强浩, 荣轩, 孟科. 基于转录组学筛选绵羊肉质性状相关候选基因[J]. 华北农学报, 2022, 37(4):220-231.doi:10.7668/hbnxb.20192944.
	Liang P, Zhang W, Feng D Z, Qiang H, Rong X, Meng K. Screening candidate genes related to meat quality traits in sheep based on transcriptome[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(4):220-231. doi: 10.7668/hbnxb.20192944
[28]	Lee H L, Qadir A S, Park H J, Chung E, Lee Y S, Woo K M, Ryoo H M, Kim H J, Baek J H. cAMP/protein kinase A signaling inhibits Dlx5 expression via activation of CREB and subsequent C/EBPβ induction in 3T3-L1 preadipocytes[J]. International Journal of Molecular Sciences, 2018, 19(10):3161.doi:10.3390/ijms19103161. URL
[29]	梁浩, 李敏, 董翔宇, 李辉, 杜志强. 肉鸡腹脂双向选择系中RNA编辑位点的鉴定研究[J]. 畜牧兽医学报, 2017, 48(9):1611-1623.doi:10.11843/j.issn.0366-6964.2017.09.006.
	Liang H, Li M, Dong X Y, Li H, Du Z Q. Identification of RNA editing sites in chicken lines divergently selected for abdominal fat content[J]. Acta Veterinaria et Zoo technica Sinica, 2017, 48(9):1611-1623.
[30]	靳易. 谷胱甘肽-S-转移酶M2调控细胞内脂滴含量的机制研究[D]. 武汉:华中农业大学, 2021.doi:10.27158/d.cnki.ghznu.2021.000148.
	Jin Y. The study on the mechanism of glutathione-S-transferase M2 regulating the content of lipid droplets content in cells[D]. Wuhan: Huazhong Agricultural University,2021.
[31]	胡凯召. 奶山羊注射5-HTP的全转录组测序分析及ATF3基因在乳腺上皮细胞中的功能验证[D]. 杨凌: 西北农林科技大学, 2020.doi:10.27409/d.cnki.gxbnu.2020.001652.
	Hu K Z. Whole transcriptome sequencing analysis of dairy goats injected with 5-HTP and function verification of ATF3 gene in goat mammary epithelial cells[D]. Yangling: Northwest A&F University, 2020.
[32]	宋昀静, 田彦梅, 孟科, 尤科梅, 冯登侦. 不同性别滩羊背最长肌中肌肉发育相关LncRNA的筛选及分析[J]. 华北农学报, 2024, 39(1):219-227.doi:10.7668/hbnxb.20194341.
	Song Y J, Tian Y M, Meng K, You K M, Feng D Z. Screening and analysis of LncRNA related to muscle development in longissimus dorsi muscle of different sex Tan sheep[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(1):219-227. doi: 10.7668/hbnxb.20194341
[33]	Avruch J. Insulin signal transduction through protein kinase cascades[J]. Molecular and Cellular Biochemistry, 1998, 182(1/2):31-48.doi:10.1023/A:100682310945.
[34]	Brozinick J T Jr, Birnbaum M J. Insulin,but not contraction,activates Akt/PKB in isolated rat skeletal muscle[J]. The Journal of Biological Chemistry, 1998, 273(24):14679-14682.doi:10.1074/jbc.273.24.14679. URL
[35]	Czech M P. Fat targets for insulin signaling[J]. Molecular Cell, 2002, 9(4):695-696.doi:10.1016/S1097-2765(02)00509-9. pmid: 11983159
[36]	Lee J H, Woo K J, Kim M A, Hong J, Kim J, Kim S H, Han K I, Iwasa M, Kim T J. Heat-killed Enterococcus faecalis prevents adipogenesis and high fat diet-induced obesity by inhibition of lipid accumulation through inhibiting C/EBP-α and PPAR-γ in the insulin signaling pathway[J]. Nutrients, 2022, 14(6):1308.doi:10.3390/nu14061308. URL
[37]	Yan X, Huang Y, Zhao J X, Long N M, Uthlaut A B, Zhu M J, Ford S P, Nathanielsz P W, Du M. Maternal obesity-impaired insulin signaling in sheep and induced lipid accumulation and fibrosis in skeletal muscle of offspring[J]. Biology of Reproduction, 2011, 85(1):172-178.doi:10.1095/biolreprod.110.089649. pmid: 21349823
[38]	崔潇, 李成萍, 张海燕, 刘雪燕, 周国利. PPP1R3C基因在3T3-L1前脂肪细胞分化过程中的功能研究[J]. 聊城大学学报(自然科学版), 2021, 34(4):95-102.doi:10.19728/j.issn1672-6634.2021.04.012.
	Cui X, Li C P, Zhang H Y, Liu X Y, Zhou G L. Study on function of PPP1R3C gene during differentiation of 3T3-L1 preadipocytes[J]. Journal of Liaocheng University(Natural Science Edition), 2021, 34(4):95-102.
[39]	Zhang Y X, Gu J, Wang L, Zhao Z L, Pan Y, Chen Y. Ablation of PPP1R3G reduces glycogen deposition and mitigates high-fat diet induced obesity[J]. Molecular and Cellular Endocrinology, 2017, 439:133-140.doi:10.1016/j.mce.2016.10.036. pmid: 27815211
[40]	Yang C Y, Wang Z X, Song Q Q, Dong B Q, Bi Y L, Bai H, Jiang Y, Chang G B, Chen G H. Transcriptome sequencing to identify important genes and lncRNAs regulating abdominal fat deposition in ducks[J]. Animals, 2022, 12(10):1256.doi:10.3390/ani12101256. URL
[41]	Ma M T, Cai B L, Kong S F, Zhou Z, Zhang J, Zhang X Q, Nie Q H. PPARGC1A is a moderator of skeletal muscle development regulated by miR-193b-3p[J]. International Journal of Molecular Sciences, 2022, 23(17):9575.doi:10.3390/ijms23179575. URL
[42]	Ramayo-Caldas Y, Fortes M R S, Hudson N J, et al. A marker-derived gene network reveals the regulatory role of PPARGC1A,HNF4G,and FOXP3 in intramuscular fat deposition of beef cattle[J]. Journal of Animal Science, 2014, 92(7):2832-2845.doi:10.2527/jas.2013-7484. pmid: 24778332
[43]	Moreira G C M, Godoy T F, Boschiero C, et al. Variant discovery in a QTL region on chromosome 3 associated with fatness in chickens[J]. Animal Genetics, 2015, 46(2):141-147.doi:10.1111/age.12263. pmid: 25643900

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