基于转录组学筛选静原鸡肌内脂肪沉积的候选基因

doi:10.7668/hbnxb.20194927

摘要/Abstract

摘要：

肌内脂肪(IMF)是重要的肉质性状之一,也是影响鸡肉质风味的重要指标。为了深入研究静原鸡不同发育阶段IMF形成的作用机理,以42(B1)、126(B2)、180(B3)日龄静原鸡(每阶段各10只)的胸肌为研究对象,利用转录组测序筛选调控静原鸡IMF沉积的候选基因。采用实时荧光定量PCR(RT-qPCR)验证测序数据的可靠性。结果显示,在B1vsB2比较组中鉴定出1 008个差异表达基因。在B1vsB3比较组中鉴定出1 396个差异表达基因。在B2vsB3比较组中鉴定出696个差异表达基因。3个比较组的GO功能富集条目主要涉及RNA聚合酶Ⅱ对转录的正向调节、细胞黏附性、细胞膜和原生质膜,相同的蛋白质结合等。同时,PPAR信号通路、FOXO信号通路、胰岛素信号通路、MAPK信号通路、脂肪酸降解、脂肪酸代谢等在KEGG通路中显著富集。这些通路中显著富集的FABP1、FOXO1、FAS和CPT1A等基因可能是调控静原鸡IMF沉积的关键候选基因。

关键词: 静原鸡, 肌内脂肪, 转录组, 肉品质

Abstract:

Intramuscular fat(IMF)is one of the important meat quality traits and an important indicator of chicken meat flavour.In order to investigate the mechanism of IMF formation at different developmental stages of Jingyuan chicken,the present study was carried out to screen candidate genes regulating IMF deposition in Jingyuan chicken by transcriptome sequencing using the pectoral muscles of 42(B1),126(B2),and 180(B3)day-old Jingyuan chicken(10 chickens at each stage).Real-time Fluorescence Quantitative PCR(RT-qPCR)was used to verify the reliability of the sequencing data.The results showed that 1 008 differentially expressed genes were identified in the B1vsB2 comparison group.1 396 differentially expressed genes were identified in the B1vsB3 comparison group. 696 differentially expressed genes were identified in the B2vsB3 comparison group.The GO function-enriched entries in the three comparison groups were mainly related to the positive regulation of transcription by RNA polymerase Ⅱ,cell adhesion,cell membrane and protoplasmic membrane,and binding of the same proteins.Meanwhile,PPAR signaling pathway,FOXO signaling pathway,insulin signaling pathway,MAPK signaling pathway,fatty acid degradation,and fatty acid metabolism were significantly enriched in KEGG pathway.Genes such as FABP1,FOXO1,FAS,and CPT1A,which were significantly enriched in these pathways,may be the key candidate genes to regulate IMF deposition in Jingyuan chicken.

Key words: Jingyuan chicken, Intramuscular fat, Transcriptome, Meat quality

中图分类号:

S831.2

胡佳欢, 赵薇, 禹保军, 赵金燕, 付希, 张统, 陈亚飞, 顾亚玲, 张娟. 基于转录组学筛选静原鸡肌内脂肪沉积的候选基因[J]. 华北农学报, 2025, 40(S1): 329-339. doi: 10.7668/hbnxb.20194927.

HU Jiahuan, ZHAO Wei, YU Baojun, ZHAO Jinyan, FU Xi, ZHANG Tong, CHEN Yafei, GU Yaling, ZHANG Juan. Transcriptomics-based Screening of Candidate Genes for Intramuscular Fat Deposition in Jingyuan Chickens[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(S1): 329-339. doi: 10.7668/hbnxb.20194927.

http://www.hbnxb.net/CN/Y2025/V40/IS1/329

图/表 12

表1 基因引物序列

Tab.1 Primer sequences of broilers

基因 Gene	基因登录号 Gene entry number	引物序列(5'-3') Primer sequences(5'-3')	退火温度/℃ Annealing temperature	产物长度/bp Product length
FAS	NM_001199487.2	F:TTGTTCGTCATCACCGTCTA	58	264
		R:TGCTTTGCTCAGTTTTGTCA
GLP2R	NM_001163248.1	F:ATGGAAGTGCAGGAAACGT	58	296
		R:GGCACGCAAGATGAAAGAT
ABCG1	XM_025145525.3	F:CGGTCCATCATCTGCACAAT	58	163
		R:GTACTGCCCTTACCAGTCGG
MSTN	NM_001001461.2	F:GAGACGATTATCACAATGCCTAC	58	294
		R:TTGCAGCACTGTCTTCACATC
PNPLA2	NM_001113291.2	F:AGCAGGGCTATCGGGATG	58	284
		R:TTGGGAGGCAAGTGTTCG
ACACB	XM_040648481.2	F:CTTCTCCCACCCAAATAGC	58	193
		R:CACTTCATCAAAGTTCCTCGT

表2 42,126,180日龄静原母鸡IMF含量

Tab.2 IMF content of 42,126 and 180 days old Jingyuan chicken

个体号 Individual number	42日龄 42 days old	126日龄 126 days old	180日龄 180 days old
1	0.32	0.81	1.08
2	0.33	0.83	1.08
3	0.33	0.89	1.11
4	0.35	0.95	1.21
5	0.35	1.01	1.26
6	0.40	1.01	1.34
7	0.43	1.05	1.54
8	0.45	1.12	1.65
9	0.46	1.25	1.73
10	0.48	1.28	1.87
平均值±标准差	0.39±0.08	1.14±0.25	1.64±0.64
mean±s

表3 测序数据质量评估

Tab.3 Quality evaluation of sequencing data

样本名称 Sample name	原始读数 Raw reads	有效读数 Clean reads	有效读数/% Vaild reads	Q20/%	Q30/%	GC含量/% GC content
B1_1	44 641 996	43 603 376	97.67	99.95	96.95	49.50
B1_2	42 203 114	41 257 270	97.76	99.95	97.04	49.50
B1_3	44 595 972	43 532 480	97.62	99.95	96.98	49.00
B1_4	45 753 490	44 642 976	97.57	99.96	97.08	49.50
B1_5	49 088 696	47 828 862	97.43	99.95	96.92	49.00
B1_6	34 765 710	32 666 642	93.96	99.93	97.23	48.50
B1_7	34 482 536	32 364 054	93.86	99.94	97.22	48.50
B1_8	51 129 984	48 000 686	93.88	99.93	97.24	48.50
B1_9	36 737 614	34 327 564	93.44	99.93	97.20	48.50
B1_10	48 130 186	44 860 422	93.21	99.93	97.19	48.50
B2_1	36 313 250	35 568 816	97.95	99.96	97.40	47.50
B2_2	39 982 564	38 974 364	97.48	99.96	97.24	49.00
B2_3	40 405 820	39 409 366	97.53	99.96	97.42	48.50
B2_4	39 202 292	38 260 934	97.60	99.95	97.16	48.50
B2_5	39 477 124	38 527 366	97.59	99.96	97.24	49.50
B2_6	39 308 358	36 822 402	93.68	99.92	97.25	48.50
B2_7	54 702 700	51 272 360	93.73	99.92	97.13	48.50
B2_8	38 046 870	35 779 792	94.04	99.93	97.21	48.50
B2_9	37 486 270	35 152 722	93.77	99.93	97.17	48.50
B2_10	42 487 358	40 047 664	94.26	99.93	97.25	48.50
B3_1	39 399 168	38 287 554	97.18	99.96	97.29	49.50
B3_2	37 621 922	36 675 748	97.49	99.96	97.51	49.50
B3_3	47 136 518	46 038 676	97.67	99.95	96.80	49.50
B3_4	44 487 340	43 411 616	97.58	99.95	96.90	48.50
B3_5	45 429 302	44 357 812	97.64	99.96	96.96	49.00
B3_6	51 633 934	48 451 864	93.84	99.94	97.27	48.50
B3_7	52 871 666	49 369 848	93.38	99.93	97.07	48.50
B3_8	48 356 002	45 427 508	93.94	99.93	97.10	48.50
B3_9	39 153 982	36 827 204	94.06	99.93	96.99	48.00
B3_10	43 223 728	40 618 378	93.97	99.93	96.97	48.50

图1 差异表达基因火山图和聚类分析 A、D.B1vsB2;B、E.B1vsB3;C、F.B2vsB3。红色.上调差异表达基因;蓝色.下调差异表达基因;灰色.差异不显著基因。

Fig.1 Volcano plot and cluster analysis of differentially expressed genes A,D.B1vsB2;B,E.B1vsB3;C,F.B2vsB3.Red.Up-regulated differentially expressed genes;Blue.Depressed differentially expressed genes;Grey.Genes with insignificant differences.

图2 B1vsB2比较组差异表达基因富集的 GO功能条目 1.RNA聚合酶Ⅱ对转录的正向调节;2.G蛋白偶联受体信号通路;3.DNA复制对转录正向调节;4.信号转导;5.RNA 聚合酶对转录的调节;6.DNA复制对转录的调节;7.免疫应答;8.RNA聚合酶N对转录的负调控;9.蛋白磷酸化;10.基因表达的正向调节;11.炎症反应;12.细胞黏附;13.离子输运;14.DNA复制对转录负向调节;15.趋化因子介导的信号通路;16.ERK1 和 ERK2 级联的正向调节;17.凋亡过程的正向调节;18.细胞群体增殖的正向调节;19.凋亡过程的负调控;20.细胞外基质组织;21.细胞群体增殖的负调控;22.骨骼肌肉组织发育;23.趋药性;24.蛋白激酶 B 信号转导的正向调节;25.T细胞分化;26.细胞质;27.原生质膜;28.细胞膜;29.细胞核;30.膜整体成分;31.细胞液;32.核质;33.细胞外空间;34.细胞外区域;35.线粒体;36.质膜不可分割的组成部分;37.细胞骨架;38.内质网;39.高尔基体;40.细胞膜内有界器官;41.相同的蛋白质结合;42.RNA聚合酶Ⅱ顺式调节区序列特异性;43.DNA 结合转录因子活性;44.金属离子结合;45.DNA 结合;46.序列特异性双链 DNA 结合;47.DNA 结合转录激活因子活性,RNA 聚合体;48.ATP 结合;49.序列特异性 DNA 结合;50.G蛋白偶联受体活性。

Fig.2 GO functional entries enriched for differentially expressed genes in the B1vsB2 comparison group 1.Positive regulation of transcription by RNA polymerase Ⅱ;2.G protein-coupled receptor signalling pathway;3.Positive regulation of transcription,DNA-templated;4.Signal transduction;5.Regulation of transcription by RNA polymerase Ⅱ;6.Regulation of transcription,DNA-templated;7.Immune response;8.Negative regulation of transcription by RNA polymerase Ⅱ;9.Protein phosphorylation;10.Positive regulation of gene expression;11.Inflammatory response;12.Cell adhesion;13.Ion transport;14.Negative regulation of transcription,DNA-templated;15.Chemokine-mediated signaling pathways;16.Positive regulation of ERK1 and ERK2 cascades;17.Positive regulation of apoptotic processes;18.Positive regulation of cell population proliferation;19.Negative regulation of apoptotic processes;20.Extracellular matrix organization;21.Negative regulation of cell population proliferation;22.Skeletal muscle tissue development;23.Chemotaxis;24.Positive regulation of protein kinase B signaling;25.T cell differentiation;26.Cytoplasm;27.Plasma membrane;28.Membrane;29.Nucleus;30.Integral component of membrane;31.Cytosol;32.Nucleoplasm;33.Extracellular space;34.Extracellular region;35.Mitochondrion;36.Integral component of the plasma membrane;37.Cytoskeleton;38.Endoplasmic reticulum;39.Golgi apparatus;40.Intracellular membrane-bounded organelle;41.Identical protein binding;42.RNA polymerase Ⅱ cis-regulatory regulartory region sequence-specific;43.DNA binding transcription factor activity;44.Metal ion binding;45.DNA binding;46.Sequence-specific double-stranded DNA binding;47.DNA binding transcription activator activity,RNA polymer;48.ATP binding;49.Sequence-specific DNA binding;50.G protein-coupled receptor activity.

图3 B1vsB3比较组差异表达基因富集的 GO功能条目 1.RNA 聚合酶Ⅱ对转录的正向调节;2.RNA 聚合酶Ⅱ对转录的调控;3.信号转导;4.细胞外基质组织;5.细胞黏附;6.细胞群体增殖的负调控;7.G 蛋白偶联受体信号通路;8.基因表达的正向调节;9.蛋白质分析;10.细胞乳头增殖的正向调节;11.DNA复制对转录正向调节;12.DNA复制对转录的调节;13.免疫反应;14.RNA聚合酶Ⅱ对转录的负向调节;15.炎症反应;16.蛋白质磷酸化;17.胶原纤维组织;18.细胞对氨基酸刺激的应答;19.细胞-细胞黏附;20.DNA复制对转录负向调节;21.细胞迁移的负向调节;22.细胞迁移的正向调节;23.细胞分化;24.血管生成;25.细胞迁移;26.细胞膜;27.原生质膜;28.细胞质;29.细胞外空间;30.膜的组成成分;31.细胞核;32.细胞外区域;33.细胞质;34.核质;35.细胞质外基质;36.质膜的组成部分;37.细胞表面;38.基底膜;39.含褶皱的细胞外基质;40.质膜的外侧;41.相同的蛋白质结合;42.金属离子结合;43.RNA 聚合酶 Ⅱ 顺式调节区序列特异性;44.DNA 结合;45.DNA 结合转录因子活性;46.蛋白质同源二聚化活性;47.水解酶活性;48.ATP 结合;49.锌离子结合;50.DNA 结合转录因子活性,RNA 聚合酶。

Fig.3 GO functional entries enriched for differentially expressed genes in the B1vsB3 comparison group 1.Positive regulation of transcription by RNA polymerase Ⅱ;2.Regulation of transcription by RNA polymerase Ⅱ;3.Signal transduction;4.Extracellular matrix organisation;5.Cell adhesion;6.Negative regulation of cell population proliferation;7.G-protein-coupled receptor signaling pathway;8.Positive regulation of gene expression;9.Proteolysis;10.Positive regulation of cell population proliferation;11.Positive regulation of transcription,DNA-templated;12.Regulation of transcription,DNA-templated;13.Immune response;14.Negative regulation of transcription by RNA polymerase Ⅱ;15.Inflammatory response;16.Protein phosphorylation;17.Collagen fibre organisation;18.Cellular response to amino acid stimulus;19.Cell-cell adhesion;20.Negative regulation of transcription,DNA-templated;21.Negative regulation of cell migration;22.Positive regulation of cell migration;23.Cell differentiation;24.Angiogenesis;25.Cell migration;26.Membrane;27.Plasma membrane;28.Cytoplasm;29.Extracellular space;30.Integral component of membrane;31.Nucleus;32.Extracellular region;33.Cytosol;34.Nucleoplasm;35.Extracellular matrix;36.Integral component of plasma membrane;37.Cell surface;38.Basement membrane;39.Collagen-containing extracellular matrix;40.External side of plasma membrane;41.Identical protein binding;42.Metal ion binding;43.RNA polymerase Ⅱ cis-regulatory region sequence specific;44.DNA binding;45.DNA-binding transcription factor activity;46.Protein homodimerization activity;47.Hydrolase activity;48.ATP binding;49.Zinc ion binding;50.DNA-binding transcription factor activity,RNA polymerase.

图4 B2vsB3比较组差异表达基因富集的 GO功能条目 1.细胞黏附性;2.RNA 聚合酶Ⅱ对转录的负调控;3.胶原纤维组织;4.细胞外基质组织;5.DNA复制对转录正向调节;6.RNA 聚合酶 Ⅱ对转录的正向调节;7.RNA聚合酶Ⅱ对转录的调节;8.转录的 DNA 模板化调控;9.信号转导;10.细胞迁移;11.基因表达的正向调节;12.肺发育;13.心脏发育;14.细胞乳头增殖的负调控;15.细胞凋亡;16.细胞与基质黏附的正向调节;17.细胞对氨基酸刺激的反应;18.血管发育;19.细胞凋亡过程的正向调节;20.细胞迁移的正向调节;21.脂质代谢过程;22.G 蛋白偶联受体信号通路;23.对低氧状态的反应;24.骨矿化;25.膜电位调节;26.细胞外空间;27.细胞膜;28.细胞质;29.原生质膜;30.细胞核;31.细胞外区域;32.膜的组成部分;33.细胞外基质;34.核质;35.细胞质;36.含胶原蛋白的细胞外基质;37.细胞表面;38.基底膜;39.质膜的组成部分;40.细胞内膜-有界细胞器;41.相同的蛋白质结合;42.金属离子结合;43.钙离子结合;44.胶原结合;45.肝素结合;46.RNA聚合酶Ⅱ顺式区域序列特异性;47.DNA结合;48.锌离子结合;49.蛋白质同源二聚化活性;50.细胞外基质结构成分。

Fig.4 GO functional entries enriched for differentially expressed genes in the B2vsB3 comparison group 1.Cell adhesion;2.Negative regulation of transcription by RNA polymerase Ⅱ;3.Collagen fibril organization;4.Extracellular matrix organization;5.Positive regulation of transcription,DNA-templated;6.Positive regulation of transcription by RNA polymerase Ⅱ;7.Regulation of transcription by RNA polymerase Ⅱ;8.Regulation of transcription,DNA-templated;9.Signal transduction;10.Cell migration;11.Positive regulation of gene expression;12.Lung development;13.Heart development;14.Negative regulation of cell population proliferation;15.Proteolysis;16.Positive regulation of cell-substrate adhesion;17.Cell response to amino acid stimulus;18.Blood vessel development;19.Positive regulation of apoptotic processes;20.Positive regulation of cell migration;21.Lipid metabolic processes;22.G protein-coupled receptor signaling pathway;23.Response to hypoxia;24.Bone mineralization;25.Regulation of membrane potential;26.Extracellular space;27.Membrane;28.Cytoplasm;29.Plasma membrane;30.Nucleus;31.Extracellular region;32.Integral component of the membrane;33.Extracellular matrix;34.Nucleoplasm;35.Cytosol;36.Collagen-containing extracellular matrix;37.Cell surface;38.Basement membrane;39.Integral component of plasma membrane;40.Intracellular membranes-bounded organelle;41.Identical protein binding;42.Metal ion binding;43.Calcium ion binding;44.Collagen binding;45.Heparin binding;46.RNA polymerase Ⅱ cis-regulatory region sequence-specific;47.DNA binding;48.Zinc ion binding;49.Protein homodimerization activity;50.Extracellular matrix structural constituent.

图5 B1vsB2比较组差异表达基因富集的 KEGG功能条目 1.细胞因子-细胞因子受体相互作用;2.氮代谢;3.甘氨酸、丝氨酸和苏氨酸代谢;4.精氨酸生物合成;5.神经活性配体-受体相互作用;6.赖氨酸、天门冬氨酸和谷氨酸代谢;7.氨基酸的生物合成;8.ECM-受体相互作用;9.血管平滑肌收缩;10.病灶黏附;11.半胱氨酸和蛋氨酸代谢;12.TGF-beta 信号通路;13.心肌细胞中的肾上腺素能信号传导;14.吞噬细胞;15.粘蛋白O型糖的生物合成;16.醚脂代谢;17.血管内皮生长因子信号通路;18.ErbB信号通路;19.细胞黏附分子;20.钙离子信号通路。

Fig.5 KEGG functional entries enriched for differentially expressed genes in the B1vsB2 comparison group 1.Cytokine-cytokine receptor interactions;2.Nitrogen metabolism;3.Glycine,serine and threonine metabolism;4.Arginine biosynthesis;5.Neuroactive ligand-receptor interaction;6.Alaine,aspartateand glutamate metabolism;7.Biosynthesis of amino acid;8.ECM-receptor interactions;9.Vascular smooth muscle contraction;10.Focal adhesion;11.Cysteine and methionine metabolism;12.TGF-beta signaling pathway;13.Adrenergic signaling in cardiomyocytes;14.Phagosome;15.Mucin type O-glycan biosynthesis biosynthesis;16.Ether lipid metabolism;17.VEGF signaling pathway;18.ErbB signaling pathway;19.Cell adhesion mplecules;20.Calcium signaling pathway.

图6 B1vsB3比较组差异表达基因富集的 KEGG功能条目 1.ECM-受体相互作用;2.病灶黏附;3.细胞因子-细胞因子受体相互作用;4.细胞黏附分子;5.花生四烯酸代谢;6.精氨酸生物合成;7.刺猬信号通路;8.钙信号通路;9.α-亚麻酸代谢;10.亚油酸代谢;11.半胱氨酸和蛋氨酸代谢;12.吞噬细胞;13.神经活性配体-受体相互作用;14.药物代谢-细胞色素P450;15.甘氨酸、丝氨酸和苏氨酸代谢;16.黑色素生成;17.血管平滑肌收缩;18.肠道产生 IgA 的免疫网络;19.精氨酸和脯氨酸的代谢;20.肌动蛋白细胞骨架的调节。

Fig.6 KEGG functional entries enriched for differentially expressed genes in the B1vsB3 comparison group 1.ECM-receptor interaction;2.Focal adhesion;3.Cytokine-cytokine receptor interaction;4.Cell adhesion molecules;5.Arachidonic acid metabolism;6.Arginine biosynthesis;7.Hedgehog signaling pathway;8.Calcium signaling pathway;9.Alpha-linolenic acid metabolism;10.Linoleic acid metabolism;11.Cysteine and methionine metabolism;12.Phagosome;13.Neuroactive ligand-receptor interaction;14.Drug metabolism-cytochrome P450;15.Glycine,serine and threonine metabolism;16.Melanogenesis;17.Vascular smooth muscle contraction;18.Intestinal immune network for IgA production;19.Arginine and proline metabolism;20.Regulation of actin cytoskeleton.

图7 B2vsB3比较组差异表达基因富集的 KEGG功能条目 1.ECM-受体相互作用;2.病灶黏附;3.脂肪细胞因子信号通路;4.花生四烯酸代谢;5.氮代谢;6.脂肪酸生物合成;7.牛磺酸和低牛磺酸代谢;8.亚油酸代谢;9.刺猬信号通路;10.胰岛素信号通路;11.FOXO信号通路;12.嘌呤代谢;13.药物代谢-其他酶;14.RIG-l样受体信号通路;15.Apelin信号通路;16.心肌细胞中的肾上腺素能信号通路;17.mTOR信号通路;18.自噬-动物;19.吞噬体;20.钙离子信号通路。

Fig.7 KEGG functional entries enriched for differentially expressed genes in the B2vsB3 comparison group 1.ECM-receptor interaction;2.Focal adhesion;3.Adipocytokine signaling pathway;4.Arachidonic acid metabolism;5.Nitrogen metabolism;6.Fatty acid biosynthesis;7.Taurine and hypotaurinr metabolism;8.Linoleic acid metabolism;9.Hedgehog signalling pathway;10.Insulin signalling pathway;11.FOXO signalling pathway;12.Purine metabolism;13.Drug metabolism-other enzymes;14.RIG-l-like receptor signaling pathway;15.Apelin signaling pathway;16.Adrenergic signalling in cardiomyocytes;17.mTOR signaling pathway;18.Autophagy-animal;19.Phagosome;20.Calcium signaling pathway.

表4 与肌内脂肪沉积有关的KEGG通路与差异表达mRNA富集结果

Tab.4 KEGG pathway associated with intramuscular fat deposition with differentially expressed mRNA enrichment results

比较组 Comparison group	KEGG富集通路 KEGG enrichment pathway	基因名称 Gene name	差异倍数 Log₂ (Fold Change)	比较组 Comparison group	KEGG富集通路 KEGG enrichment pathway	基因名称 Gene name	差异倍数 Log₂ (Fold Change)
B1vsB2	MAPK 信号通路	TGFB3	1.95	B2vsB3	FOXO 信号通路	IRS4	1.03
		DUSP1	1.26			TGFB3	-2.39
	FOXO 信号通路	TGFB3	1.95			FOXO1	1.05
		FOXO1	-1.40		脂肪细胞因子信号通路	IRS4	1.03
	PPAR 信号通路	CPT1A	-1.55			CPT1A	1.76
		FABP1	1.19			ACACB	1.68
	胰岛素信号通路	FOXO1	-1.40		胰岛素信号通路	IRS4	1.03
	脂肪酸降解	CPT1A	-1.55			FOXO1	1.05
	脂肪酸代谢	CPT1A	-1.55			ACACB	1.68
	脂肪细胞因子信号通路	CPT1A	-1.55		脂肪酸降解	CPT1A	1.76
B1vsB3	代谢途径	PNPLA2	1.82		PPAR 信号通路	CPT1A	1.76
	甘油酯代谢	PNPLA2	1.82		脂肪酸代谢	ACACB	1.68
	脂肪酸的生物合成	ACACB	1.54			CPT1A	1.76
	胰岛素信号通路	ACACB	1.54		脂肪酸的生物合成	ACACB	1.68
	丙酮酸代谢	ACACB	1.54		代谢途径	PNPLA2	1.10
	脂肪细胞因子信号通路	ACACB	1.54			ACACB	1.68
	MAPK 信号通路	FAS	-1.27		丙酮酸代谢	ACACB	1.68
	ABC 转运体	ABCG1	-2.96			CPT1A	1.76
B2vsB3	MAPK 信号通路	TGFB3	-2.39		甘油酯代谢	PNPLA2	1.10

图8 差异表达基因RT-qPCR比较分析不同小写字母表示不同比较组之间差异显著(P<0.05)。

Fig.8 Comparative RT-qPCR analysis of differentially expressed genes Different lowercase letters indicate significant differences between different comparisons(P<0.05).

参考文献 38

[1]	Jin C L, Zeng H R, Gao C Q, Yan H C, Tan H Z, Wang X Q. Dietary supplementation with pioglitazone hydrochloride and chromium methionine manipulates lipid metabolism with related genes to improve the intramuscular fat and fatty acid profile of yellow-feathered chickens[J]. Journal of the Science of Food and Agriculture, 2020, 100(3):1311-1319.doi:10.1002/jsfa.10146. URL
[2]	赵薇, 曹国伟, 王卫振, 邓占钊, 张娟, 顾亚玲, 虎红红, 黄增文. 静原鸡胸肌与腿肌肌苷酸特异性沉积相关基因的生物信息学分析[J]. 中国农业大学学报, 2022, 27(5):227-239.doi:10.11841/j.issn.1007-4333.2022.05.21.
	Zhao W, Cao G W, Wang W Z, Deng Z Z, Zhang J, Gu Y L, Hu H H, Huang Z W. Bioinformatics analysis of genes related to inosine monophosphate specific deposition in chest muscle and leg muscle of Jingyuan chicken[J]. Journal of China Agricultural University, 2022, 27(5):227-239.
[3]	吕亚宁, 贺琛昕, 兰旅涛. 猪肌内脂肪与肉品质的关系及其影响因素的研究进展[J]. 中国畜牧兽医, 2020, 47(2):554-563.doi:10.16431/j.cnki.1671-7236.2020.02.027.
	Lyu Y N, He C X, Lan L T. Research advances on the relationship between intramuscular fat and meat quality and influence factor of intramuscular fat in pigs[J]. China Animal Husbandry & Veterinary Medicine, 2020, 47(2):554-563.
[4]	章琳俐, 李丽, 朱志明, 缪中纬, 辛清武, 郑嫩珠. 基于RNA-seq鉴定连城白鸭肉质风味相关候选基因[J]. 农业生物技术学报, 2021, 29(4):711-722.doi:10.3969/j.issn.1674-7968.2021.04.009.
	Zhang L L, Li L, Zhu Z M, Miao Z W, Xin Q W, Zheng N Z. Identification of candidate genes related to meat flavor in Liancheng white duck(Anas platyrhynchos)based on RNA-seq[J]. Journal of Agricultural Biotechnology, 2021, 29(4):711-722.
[5]	San J S, Du Y T, Wu G F, Xu R F, Yang J C, Hu J M. Transcriptome analysis identifies signaling pathways related to meat quality in broiler chickens the extracellular matrix(ECM)receptor interaction signaling pathway[J]. Poultry Science, 2021, 100(6):101135.doi:10.1016/j.psj.2021.101135.
[6]	Xiao C, Sun T T, Yang Z L, Xu W W, Wang J, Zeng L H, Deng J X, Yang X R. Transcriptome landscapes of differentially expressed genes related to fat deposits in Nandan-Yao chicken[J]. Functional & Integrative Genomics, 2021, 21(1):113-124.doi:10.1007/s10142-020-00764-7.
[7]	Liu L, Liu X J, Cui H X, Liu R R, Zhao G P, Wen J. Transcriptional insights into key genes and pathways controlling muscle lipid metabolism in broiler chickens[J]. BMC Genomics, 2019, 20(1):863.doi:10.1186/s12864-019-6221-0. pmid: 31729950
[8]	Yu B J, Cai Z Y, Liu J M, Zhao W, Fu X, Gu Y L, Zhang J. Transcriptome and co-expression network analysis reveals the molecular mechanism of inosine monophosphate-specific deposition in chicken muscle[J]. Frontiers in Physiology, 2023, 14:1199311.doi:10.3389/fphys.2023.1199311.
[9]	Wang S S, Zhang Y, Yuan X Y, Pan R, Yao W C, Zhong L, Song Q Q, Zheng S H, Wang Z X, Xu Q, Chang G B, Chen G H. Identification of differentially expressed microRNAs during preadipocyte differentiation in Chinese crested duck[J]. Gene, 2018, 661:126-132.doi:10.1016/j.gene.2018.03.085. pmid: 29604463
[10]	Gan L, Yan J, Liu Z J, Feng M, Sun C. Adiponectin prevents reduction of lipid-induced mitochondrial biogenesis via AMPK/ACC2 pathway in chicken adipocyte[J]. Journal of Cellular Biochemistry, 2015, 116(6):1090-1100.doi:10.1002/jcb.25064. pmid: 25536013
[11]	Haunerland N H, Spener F. Fatty acid-binding proteins insights from genetic manipulations[J]. Progress in Lipid Research, 2004, 43(4):328-349.doi:10.1016/j.plipres.2004.05.001. pmid: 15234551
[12]	农伟伦, 毕英杰, 李宏旭, 张丽. 甘肃黑猪H-FABP基因克隆、SNPs筛查及生物信息学分析[J]. 浙江农业学报, 2018, 30(3):378-385.doi:10.3969/j.issn.1004-1524.2018.03.04.
	Nong W L, Bi Y J, Li H X, Zhang L. H-FABP gene cloning,SNPs screening and bioinformatics analysis in Gansu black pig[J]. Acta Agriculturae Zhejiangensis, 2018, 30(3):378-385.
[13]	杨蓉. 贵州地方猪胶原蛋白特性及其对肉质的影响研究[D]. 贵阳: 贵州大学, 2020.doi:10.27047/d.cnki.ggudu.2020.001138.
	Yang R. Study on the characteristics of collagen in Guizhou local pigs and its influence on meat quality[D]. Guiyang: Guizhou University, 2020.
[14]	Thompson J, Winter N, Terwey D, Bratt J, Banaszak L. The crystal structure of the liver fatty acid-binding protein a complex with two bound oleates[J]. Journal of Biological Chemistry, 1997, 272(11):7140-7150.doi:10.1074/jbc.272.11.7140. pmid: 9054409
[15]	Mao H, Xu X, Liu H, Cao H, Dong X, Xu N, Zou X, Yin Z. The temporal-spatial patterns,polymorphisms and association analysis with meat quality traits of FABP1 gene in domestic pigeons(Columba livia)[J]. British Poultry Science, 2020, 61(3):232-241.doi:10.1080/00071668.2020.1724880. pmid: 32063032
[16]	杨绍康, 常振宇, 严飞飞, 商鹏, 强巴央宗. 藏猪和大约克猪FABP1基因多态性及差异表达分析[J]. 高原农业, 2023, 7(4):400-408.doi:10.19707/j.cnki.jpa.2023.04.008.
	Yang S K, Chang Z Y, Yan F F, Shang P, Qiang Y Z. Analysis of FABP1 gene polymorphism and differential expression between Tibetan and Yorkshire pigs[J]. Journal of Plateau Agriculture, 2023, 7(4):400-408.
[17]	熊讯, 阮涌, 许厚强. 基于转录组分析干扰FABP1基因对猪肌内脂肪沉积的影响[J]. 南方农业学报, 2023, 54(3):724-734.doi:10.3969/j.issn.2095-1191.2023.03.008.
	Xiong X, Ruan Y, Xu H Q. Transcriptome-based analysis of the effects of interference with FABP1 gene on intramuscular fat deposition in pigs[J]. Journal of Southern Agriculture, 2023, 54(3):724-734.
[18]	Wang Y J, Tang K Q, Zhang W, Guo W L, Wang Y N, Zan L S, Yang W C. Fatty acid-binding protein 1 increases steer fat deposition by facilitating the synthesis and secretion of triacylglycerol in liver[J]. PLoS One, 2019, 14(4):e0214144.doi:10.1371/journal.pone.0214144.
[19]	Zhang Q, Shi H, Liu W, Wang Y, Wang Q, Li H. Differential expression of L-FABP and L-BABP between fat and lean chickens[J]. Genetics and Molecular Research, 2013, 12(4):4192-4206.doi:10.4238/2013.october.7.5. pmid: 24114214
[20]	Adiguzel D, Celik-Ozenci C. FoxO1 is a cell-specific core transcription factor for endometrial remodeling and homeostasis during menstrual cycle and early pregnancy[J]. Human Reproduction Update, 2021, 27(3):570-583.doi:10.1093/humupd/dmaa060. pmid: 33434267
[21]	赵航, 张运佳, 树林一, 宋光耀. FOXO1在肝脏糖脂代谢中的作用[J]. 基础医学与临床, 2019, 39(1):115-119.doi:10.16352/j.issn.1001-6325.2019.01.039.
	Zhao H, Zhang Y J, Shu L Y, Song G Y. Effect of FOXO1 on glucose and lipid metabolism in liver[J]. Basic & Clinical Medicine, 2019, 39(1):115-119.
[22]	Kurakazu I, Akasaki Y, Hayashida M, Tsushima H, Goto N, Sueishi T, Toya M, Kuwahara M, Okazaki K, Duffy T, Lotz M K, Nakashima Y. FOXO1 transcription factor regulates chondrogenic differentiation through transforming growth factor β1 signaling[J]. Journal of Biological Chemistry, 2019, 294(46):17555-17569.doi:10.1074/jbc.RA119.009409. pmid: 31601652
[23]	熊祥平. Sirt1通过PI3K-Akt-mTOR信号途径对鹅肝细胞脂质沉积的影响研究[D]. 雅安: 四川农业大学, 2016.
	Xiong X P. Effect of Sirt1 on lipid deposition in goose liver cells through PI3K-Akt-mTOR signaling pathway[D]. Yaan: Sichuan Agricultural University, 2016.
[24]	温小庆. 高NEFA经FOXO1途径诱导奶牛子宫内膜上皮细胞凋亡的调节机制[D]. 大庆: 黑龙江八一农垦大学, 2023.doi:10.27122/d.cnki.ghlnu.2023.000048.
	Wen X Q. Regulatory mechanism of apoptosis induced by high NEFA via FOXO1 pathway in endometrial epithelial cells of dairy cows[D]. Daqing: Heilongjiang Bayi Agricultural Reclamation University, 2023.
[25]	李家旋, 宋倩倩, 徐鸣鸿, 周婷婷, 严丹, 田慧月, 常国斌, 陈国宏, 王志秀. FoxO1基因对鸭脂肪沉积的作用及机制初探[J]. 中国家禽, 2022, 44(7):114-118.doi:10.16372/j.issn.1004-6364.2022.07.018.
	Li J X, Song Q Q, Xu M H, Zhou T T, Yan D, Tian H Y, Chang G B, Chen G H, Wang Z X. Preliminary study on effect and mechanism of FoxO1 gene on fat deposition in duck[J]. China Poultry, 2022, 44(7):114-118.
[26]	Dai J Y, Liang K, Zhao S, Jia W T, Liu Y, Wu H K, Lv J, Cao C, Chen T, Zhuang S T, Hou X M, Zhou S J, Zhang X N, Chen X W, Huang Y Y, Xiao R P, Wang Y L, Luo T P, Xiao J Y, Wang C. Chemoproteomics reveals baicalin activates hepatic CPT1 to ameliorate diet-induced obesity and hepatic steatosis[J]. Proceedings of the National Academy of Sciences of the United States of America, 2018, 115(26):E5896-E5905.doi:10.1073/pnas.1801745115.
[27]	Morash A J, McClelland G B. Regulation of carnitine palmitoyltransferase(CPT)I during fasting in rainbow trout(Oncorhynchus mykiss)promotes increased mitochondrial fatty acid oxidation[J]. Physiological and Biochemical Zoology, 2011, 84(6):625-633.doi:10.1086/662552. URL
[28]	Tan S K, Welford S M. Lipid in renal carcinoma:queen bee to target?[J]. Trends in Cancer, 2020, 6(6):448-450.doi:10.1016/j.trecan.2020.02.017. URL
[29]	Thapa D, Wu K Y, Stoner M W, Xie B X, Zhang M L, Manning J R, Lu Z P, Li J H, Chen Y, Gucek M, Playford M P, Mehta N N, Harmon D, O'Doherty R M, Jurczak M J, Sack M N, Scott I. The protein acetylase GCN5L1 modulates hepatic fatty acid oxidation activity via acetylation of the mitochondrial β-oxidation enzyme HADHA[J]. Journal of Biological Chemistry, 2018, 293(46):17676-17684.doi:10.1074/jbc.AC118.005462. URL
[30]	梁计峻, 林亚秋, 俞雨阳, 王永, 朱江江. 山羊CPT1A基因的克隆表达及肌内脂肪含量的相关性分析[J]. 华北农学报, 2019, 34(5):231-238.doi:10.7668/hbnxb.201751529.
	Liang J J, Lin Y Q, Yu Y Y, Wang Y, Zhu J J. Cloning and expression of goat CPT1A gene and its correlation with intramuscular fat content[J]. Acta Agriculturae Boreali-Sinica, 2019, 34(5):231-238.
[31]	叶凤江. CPT1A、MTP基因干扰对三种糖诱导的鹅肝细胞脂质沉积的影响[D]. 雅安: 四川农业大学, 2018.
	Ye F J. Effects of CPT1A and MTP gene interference on lipid deposition in goose liver cells induced by three sugars[D]. Yaan: Sichuan Agricultural University, 2018.
[32]	屠云洁, 巨晓军, 单艳菊, 姬改革, 刘一帆, 章明, 邹剑敏, 束婧婷. 花山麻鸡CPT1A基因时空表达规律研究[J]. 中国家禽, 2021, 43(6):20-24.doi:10.16372/j.issn.1004-6364.2021.06.004.
	Tu Y J, Ju X J, Shan Y J, Ji G G, Liu Y F, Zhang M, Zou J M, Shu J T. Temporal and spatial expression of CPT1A gene in Huashan partridge chicken[J]. China Poultry, 2021, 43(6):20-24.
[33]	王燕新, 廖圆圆, 阿依木古丽, 徐红伟, 杨具田, 蔡勇. 绵羊脂肪代谢与调控机理研究进展[J]. 黑龙江畜牧兽医, 2020(17):33-36.doi:10.13881/j.cnki.hljxmsy.2019.10.0118.
	Wang Y X, Liao Y Y, A Y M G L, Xu H W, Yang J T, Cai Y. Recent progress on the mechanism of fat metabolism in sheep[J]. Heilongjiang Animal Science and Veterinary Medicine, 2020(17):33-36.
[34]	吴雨, 苑洪霞, 陈祥. PPARγ与FAS基因在白洗猪及苏白杂交F₁代不同组织中的表达分析[J]. 中国畜牧杂志, 2020, 56(3):66-69.doi:10.19556/j.0258-7033.20190225-05.
	Wu Y, Yuan H X, Chen X. Expression analysis of PPARγ and FAS genes in different tissues of white washed pigs and the F₁ generation of Subai crosses[J]. Chinese Journal of Animal Husbandry, 2020, 56(3):66-69.
[35]	Zhao S M, Ren L J, Chen L, Zhang X, Cheng M L, Li W Z, Zhang Y Y, Gao S Z. Differential expression of lipid metabolism related genes in porcine muscle tissue leading to different intramuscular fat deposition[J]. Lipids, 2009, 44(11):1029-1037.doi:10.1007/s11745-009-3356-9. pmid: 19847466
[36]	Lashkari S, Moller J W, Jensen S K, Hellgren L I, Sørensen M T, Theil P K, Sejrsen K. Changes in long-chain fatty acid composition of milk fat globule membrane and expression of mammary lipogenic genes in dairy cows fed sunflower seeds and rumen-protected choline[J]. Journal of Animal Physiology and Animal Nutrition, 2020, 104(6):1606-1619.doi:10.1111/jpn.13386. URL
[37]	Lyu Y T, Zhang S H, Guan W T, Chen F, Zhang Y Z, Chen J, Liu Y. Metabolic transition of milk triacylglycerol synthesis in response to varying levels of palmitate in porcine mammary epithelial cells[J]. Genes & Nutrition, 2018, 13(1):18.doi:10.1186/s12263-018-0606-6.
[38]	曲桂娟, 董晓庆, 孟轲音, 杨连玉, 丁尚红, 秦贵信. 不同杂交组合肉牛背最长肌FAS mRNA表达及其对肌内脂肪沉积的影响[J]. 中国兽医学报, 2016, 36(7):1183-1185,1211.doi:10.16303/j.cnki.1005-4545.2016.07.20.
	Qu G J, Dong X Q, Meng K Y, Yang L Y, Ding S H, Qin G X. FAS mRNA expression of different hybridized combination beef cattle and its effect on intramuscular fat deposition[J]. Chinese Journal of Veterinary Science, 2016, 36(7):1183-1185,1211.

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