Article(id=1307007598729912781, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, articleNumber=null, orderNo=null, doi=10.11843/j.issn.0366-6964.2026.01.018, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1750089600000, receivedDateStr=2025-06-17, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1789544494564, onlineDateStr=2026-09-16, pubDate=1769097600000, pubDateStr=2026-01-23, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789544494564, onlineIssueDateStr=2026-09-16, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789544494564, creator=13701087609, updateTime=1789544494564, updator=13701087609, issue=Issue{id=1307007396681900041, tenantId=1146029695717560320, journalId=1306274286668972124, year='2026', volume='57', issue='1', pageStart='1', pageEnd='568', issueExtLink='null', onlineDate='null', pubDate='1769097600000', pubDateStr='2026-01-23', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789544446392, creator='13701087609', updateTime=1789609890784, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1307281890638057732, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1307281890738721029, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=206, endPage=220, ext={EN=ArticleExt(id=1307007598939627982, articleId=1307007598729912781, tenantId=1146029695717560320, journalId=1306274286668972124, language=EN, title=Comparative Analysis of Histomorphology and Transcriptome in Pectoral Muscle Tissues between Shitou and Wuzong Geese, columnId=1307007413870158054, journalTitle=Acta Veterinaria et Zootechnica Sinica, columnName=ANIMAL GENETICS AND BREEDING, runingTitle=null, highlight=null, articleAbstract=

This study aimed to compare the morphological characteristics of pectoral muscle myofibers and transcriptomic profiles between Shitou geese and Wuzong geese at 40- and 90-day-old stages, and to identify candidate genes underlying divergent breast muscle growth across breeds and ages. A total of 3 male Shitou geese and 3 male Wuzong geese from the same batch were randomly selected at 40 and 90 days of age. Pectoral muscle tissues were collected for morphological analysis of muscle fibers using H&E staining. Transcriptome sequencing was performed to identify differentially expressed genes (DEGs) related to muscle growth, and key candidate genes were further validated by reverse transcription quantitative PCR (RT‑qPCR). H&E staining results revealed that at 40 days of age, the density of pectoral muscle fibers in Shitou geese was significantly higher than that in Wuzong geese (P<0.001), while the diameter of pectoral muscle fibers was significantly smaller (P<0.001). At 90 days of age, however, the density of pectoral muscle fibers in Shitou geese was significantly lower than that in Wuzong geese (P<0.05), and no significant difference was observed in pectoral muscle fiber diameter between the two breeds (P>0.05). Transcriptome sequencing of pectoral muscle identified 5 719 DEGs in total, including 3 907 DEGs between different ages within the same breed and 1 812 DEGs between the two breeds at the same age. KEGG enrichment analysis revealed that pathways including ECM-receptor interaction, cell adhesion molecules, and focal adhesion critically regulated pectoral muscle growth across ages within breeds. Cytokine-cytokine receptor interaction and cell adhesion molecule pathways were key regulators of growth differences between breeds at the same age. Eight candidate genes (COL4A2, ITGB3, CAV1, CCND2, MYLK2, PDGFRA, ACTN2, and TNNC2) closely associated with pectoral muscle development were identified. RT-qPCR validation confirmed high correlation (r>0.90) with transcriptome data, supporting the reliability of sequencing results. In summary, this study revealed significant variations in myofiber morphology across developmental stages and between Shitou and Wuzong goose breeds. Key pathways including focal adhesion, ECM-receptor interaction, and cell adhesion molecules were found to mediate pectoral muscle myofibre hypertrophy through regulating the expression of candidate genes (e.g., COL4A2, ITGB3, CAV1, etc.). These findings establish a theoretical foundation for molecular marker-assisted breeding in meat geese, accelerating genetic improvement programs and new breed development.

, authors=Zeting XIE, Xiaoyun HOU, Yinkai ZHAN, Suiyuan LIN, Purui ZHU, Shiqi GUO, Shuang QIN, Yangbing LUO, Xiujin LI, Yunmao HUANG, Zhongping WU, Xumeng ZHANG, authorsList=Zeting XIE, Xiaoyun HOU, Yinkai ZHAN, Suiyuan LIN, Purui ZHU, Shiqi GUO, Shuang QIN, Yangbing LUO, Xiujin LI, Yunmao HUANG, Zhongping WU, Xumeng ZHANG, authorCompany=null, correspAuthors=Zhongping WU, Xumeng ZHANG, authorNote=null, correspAuthorsNote=
WU Zhongping, E-mail:
ZHANG Xumeng, E-mail:
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旨在对比狮头鹅和乌鬃鹅在40和90日龄时,其胸肌肌纤维形态和转录组的差异,挖掘导致不同日龄、不同鹅种胸肌生长差异的候选基因。本研究随机选取同批次健康的40、90日龄的狮头鹅和乌鬃鹅公鹅各3只,采集胸肌组织,利用H&E染色分析肌纤维形态差异,结合转录组测序筛选出与肌肉生长相关的差异表达基因(DEGs),并采用实时荧光定量PCR(RT-qPCR)验证关键候选基因。H&E染色结果发现,40日龄时,狮头鹅的胸肌肌纤维密度极显著高于乌鬃鹅(P<0.001),胸肌肌纤维直径极显著低于乌鬃鹅(P<0.001),而90日龄时,狮头鹅的胸肌肌纤维密度显著低于乌鬃鹅(P<0.05),胸肌纤维直径无显著差异(P>0.05)。胸肌转录组测序共筛选出DEGs 5 719个,不同日龄同一鹅种间的DEGs 3 907个,同一日龄不同鹅种间的DEGs 1 812个。对胸肌DEGs进行KEGG通路富集分析发现,ECM-受体相互作用、细胞黏附分子和黏着斑等6个通路在不同日龄同一鹅种间胸肌生长中发挥关键调控作用,细胞因子-细胞因子受体相互作用和细胞黏附分子通路在同一日龄不同鹅种间胸肌生长中发挥关键调控作用,并从中筛选出8个与胸肌生长发育密切相关的候选基因,分别是COL4A2、ITGB3、CAV1、CCND2、MYLK2、PDGFRA、ACTN2和TNNC2。通过对上述基因进行RT-qPCR验证,发现其结果与转录组测序结果相关系数均在0.90以上,说明测序结果可靠。综上,本研究发现,不同日龄及品种间的肌纤维形态差异显著,黏着斑通路、ECM-受体相互作用和细胞黏附分子等通路可通过调控COL4A2、ITGB3和CAV1等8个候选基因表达介导两品种胸肌肌纤维的肥大过程。本研究将为肉鹅分子标记辅助育种提供数据基础,进而推动肉鹅遗传改良和新品种培育。

, authors=谢泽婷, 侯晓韵, 詹胤锴, 林随源, 朱溥睿, 郭诗琦, 秦双, 罗阳冰, 李秀金, 黄运茂, 伍仲平, 张续勐, authorsList=谢泽婷, 侯晓韵, 詹胤锴, 林随源, 朱溥睿, 郭诗琦, 秦双, 罗阳冰, 李秀金, 黄运茂, 伍仲平, 张续勐, authorCompany=null, correspAuthors=伍仲平, 张续勐, authorNote=

谢泽婷,本科生,主要从事动物遗传育种研究,E-mail:

谢泽婷与侯晓韵为同等贡献作者

侯晓韵,本科生,主要从事动物遗传育种研究,E-mail:

, correspAuthorsNote=
伍仲平,主要从事动物遗传育种研究,E-mail:
张续勐,主要从事动物遗传育种研究,E-mail:
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A. The images from left to right and from top to bottom show HE-stained sections of 40 d Wuzong goose, 40 d Shitou goose, 90 d Wuzong goose, and 90 d Shitou goose, respectively (scale bar=20 μm); B. The statistics of myofiber density; C. The statistics of myofiber diameter. *. P<0.05; ***. P<0.001

, figureFileSmall=y5PZGA25D9+wEhh9vvb9MQ==, figureFileBig=zxDT7SOC7HoP9WprFP6/Yg==, tableContent=null), ArticleFig(id=1307286467873628294, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=图1, caption=40、90日龄乌鬃鹅(WZE)和狮头鹅(STE)的胸肌H&E染色切片及肌纤维密度和直径统计

A.从左到右从上到下依次为40日龄乌鬃鹅、40日龄狮头鹅、90 日龄乌鬃鹅、90日龄狮头鹅的H&E染色切片图,比例尺为20 μm;B.肌纤维密度统计图;C.肌纤维直径统计图。*.P<0.05;***.P<0.001

, figureFileSmall=y5PZGA25D9+wEhh9vvb9MQ==, figureFileBig=zxDT7SOC7HoP9WprFP6/Yg==, tableContent=null), ArticleFig(id=1307286467965902983, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Fig.2, caption=Spearman’s correlation coefficient analysis of pectoral muscle transcriptome sequencing data from Wuzong and Shitou geese at 40 d and 90 d, figureFileSmall=OUwx4qoxA77Ii45W31SgfA==, figureFileBig=onX28Y99iVjo3H5xocHv6A==, tableContent=null), ArticleFig(id=1307286468163035272, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=图 2, caption=40、90日龄狮头鹅与乌鬃鹅胸肌转录组测序数据Spearman相关系数分析

***. P<0.001

, figureFileSmall=OUwx4qoxA77Ii45W31SgfA==, figureFileBig=onX28Y99iVjo3H5xocHv6A==, tableContent=null), ArticleFig(id=1307286468230144137, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Fig.3, caption=Cluster analysis of pectoral muscle transcriptome sequencing data in Shitou goose and Wuzong goose at 40 d and 90 d, figureFileSmall=ZNz2NsZkbjo8vL+/EKfLcw==, figureFileBig=9QSyZoYdxD0bX4JjQsFajQ==, tableContent=null), ArticleFig(id=1307286468297253002, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=图3, caption=40、90日龄狮头鹅与乌鬃鹅胸肌转录组测序数据聚类分析, figureFileSmall=ZNz2NsZkbjo8vL+/EKfLcw==, figureFileBig=9QSyZoYdxD0bX4JjQsFajQ==, tableContent=null), ArticleFig(id=1307286468360167563, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Fig.4, caption=Scatter plot of KEGG pathway enrichment of DEGs in the pectoral muscles of Wuzong and Shitou geese at 40 d and 90 d

The vertical axis represents pathway names, and the horizontal axis represents the enrichment factors. The color of the circles indicate the q-value of pathway enrichment, the smaller the q-value, the redder the color. The size of the circles represent the number of genes enriched in the pathway, the larger the circle, the greater the number of enriched genes

, figureFileSmall=SXz0JJ/SBvuP/tDRVfTzgw==, figureFileBig=u4xoanhgzZikFlUNmUQJEA==, tableContent=null), ArticleFig(id=1307286468423082124, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=图4, caption=40、90日龄狮头鹅与乌鬃鹅胸肌DEGs的KEGG通路富集散点图

纵坐标表示通路名称,横坐标为富集因子。圆圈颜色代表通路富集的q 值,q 值越小颜色越红,圆圈的大小表示该通路中富集到的基因数目,点越大富集到的基因数目越多

, figureFileSmall=SXz0JJ/SBvuP/tDRVfTzgw==, figureFileBig=u4xoanhgzZikFlUNmUQJEA==, tableContent=null), ArticleFig(id=1307286470079832205, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Fig.5, caption=Correlation analysis of transcriptome sequencing data and real-time fluorescence quantitative PCR data, figureFileSmall=5bVsp5uP1O4Vx85BYGHBAA==, figureFileBig=Breqp6oYfzXj+hBcsBcHIQ==, tableContent=null), ArticleFig(id=1307286470197272718, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=图5, caption=转录组测序数据与实时荧光定量PCR数据相关性分析, figureFileSmall=5bVsp5uP1O4Vx85BYGHBAA==, figureFileBig=Breqp6oYfzXj+hBcsBcHIQ==, tableContent=null), ArticleFig(id=1307286470289547407, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Table 1, caption=

Primer sequences for the target genes and internal control genes used in the experiment

, figureFileSmall=null, figureFileBig=null, tableContent=

基因名称

Gene name

正向(5′→3′)

Forward

反向(3′→5′)

Reverse

GenBank序列号

Accession number

β-actinATGTCGCCCTGGATTTCGCACAGGACTCCATACCCAAGAAXM_066977989.1
COL4A2GGTTTGGTTTGTCAGGATCTGGAAACACTGGCATCCTCCACTXM_048080798.2
ITGB3AGGGATGAATCTGGCAATCTACACAAACTCAGCTTCACCCTGCXM_066994002.1
CAV1GATCCTCCTGCCTCCGTTATGTACAGAAAGCCCTCCGAGTXM_066991389.1
CCND2ACAAGCACAGATGTGGACTGAAGGATTGCTTTGCTGTTGCXM_066977338.1
MYLK2TCATCAAAGAGAAAGGCGGCATTTCTTGAGCAGCACCTGGXM_066978075.1
PDGFRATCAAATGAGGCAGTTGTCGGGGAACAATAGTCGGCAGAGGXM_013172769.3
ACTN2AAGCCTGCCTCATCAGTCTACAAACTCAGCTTCACCCTGCXM_066994002.1
TNNC2CTTCCGCATCTTCGACAAGAACGTCTTCTATGTCCTCCTCGGTXM_048050648.2
), ArticleFig(id=1307286470369239184, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=表1, caption=

试验相关的基因以及内参基因的引物序列

, figureFileSmall=null, figureFileBig=null, tableContent=

基因名称

Gene name

正向(5′→3′)

Forward

反向(3′→5′)

Reverse

GenBank序列号

Accession number

β-actinATGTCGCCCTGGATTTCGCACAGGACTCCATACCCAAGAAXM_066977989.1
COL4A2GGTTTGGTTTGTCAGGATCTGGAAACACTGGCATCCTCCACTXM_048080798.2
ITGB3AGGGATGAATCTGGCAATCTACACAAACTCAGCTTCACCCTGCXM_066994002.1
CAV1GATCCTCCTGCCTCCGTTATGTACAGAAAGCCCTCCGAGTXM_066991389.1
CCND2ACAAGCACAGATGTGGACTGAAGGATTGCTTTGCTGTTGCXM_066977338.1
MYLK2TCATCAAAGAGAAAGGCGGCATTTCTTGAGCAGCACCTGGXM_066978075.1
PDGFRATCAAATGAGGCAGTTGTCGGGGAACAATAGTCGGCAGAGGXM_013172769.3
ACTN2AAGCCTGCCTCATCAGTCTACAAACTCAGCTTCACCCTGCXM_066994002.1
TNNC2CTTCCGCATCTTCGACAAGAACGTCTTCTATGTCCTCCTCGGTXM_048050648.2
), ArticleFig(id=1307286470444736657, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Table 2, caption=

Uniform quality analysis of pectoral muscle transcriptome sequencing data from Wuzong and Shitou geese at 40 d and 90 d

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

质控数据

Quality control data

GC含量/%

GC content

≥Q30/%

比对上的片段数

Aligned Reads

S40X-125 993 68652.8992.6539 532 520(76.04%)
S40X-229 258 92751.9993.0144 956 487(76.83%)
S40X-326 641 01251.6492.4741 752 034(78.36%)
S90X-128 035 90352.8893.1343 059 724(76.79%)
S90X-228 889 93552.6292.9744 994 751(77.87%)
S90X-326 895 65351.3792.9542 246 027(78.54%)
W40X-138 520 71952.6492.9959 236 364(76.89%)
W40X-229 841 66951.2392.4847 271 191(79.20%)
W40X-321 850 22452.1092.8334 262 682(78.40%)
W90X-131 078 12851.0192.7449 449 086(79.56%)
W90X-242 480 65051.2292.3266 544 240(78.32%)
W90X-333 421 26951.3492.5053 187 288(79.57%)
), ArticleFig(id=1307286470549594258, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=表 2, caption=

40、90日龄狮头鹅与乌鬃鹅胸肌转录组测序数据的质量评估

, figureFileSmall=null, figureFileBig=null, tableContent=

样品

Sample

质控数据

Quality control data

GC含量/%

GC content

≥Q30/%

比对上的片段数

Aligned Reads

S40X-125 993 68652.8992.6539 532 520(76.04%)
S40X-229 258 92751.9993.0144 956 487(76.83%)
S40X-326 641 01251.6492.4741 752 034(78.36%)
S90X-128 035 90352.8893.1343 059 724(76.79%)
S90X-228 889 93552.6292.9744 994 751(77.87%)
S90X-326 895 65351.3792.9542 246 027(78.54%)
W40X-138 520 71952.6492.9959 236 364(76.89%)
W40X-229 841 66951.2392.4847 271 191(79.20%)
W40X-321 850 22452.1092.8334 262 682(78.40%)
W90X-131 078 12851.0192.7449 449 086(79.56%)
W90X-242 480 65051.2292.3266 544 240(78.32%)
W90X-333 421 26951.3492.5053 187 288(79.57%)
), ArticleFig(id=1307286470650257555, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Table 3, caption=

Statistics on the number of differentially expressed genes in the pectoral muscles of Wuzong and Shitou geese at 40 d and 90 d

, figureFileSmall=null, figureFileBig=null, tableContent=

差异表达基因集

DEGs set

差异表达基因数目

Number of DEGs

上调基因数目

Number of up-regulated genes

下调基因数目

Number of down-regulated genes

不同日龄同一鹅种 Same goose breed with different ages
S40X vs S90X1 9128361 076
W40X vs W90X1 9959231 072
同一日龄不同鹅种 Different goose breeds with the same age
S40X vs W40X319187132
S90X vs W90X1 493858635
), ArticleFig(id=1307286470725755028, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=表3, caption=

40、90日龄狮头鹅与乌鬃鹅胸肌差异表达基因数目统计表

, figureFileSmall=null, figureFileBig=null, tableContent=

差异表达基因集

DEGs set

差异表达基因数目

Number of DEGs

上调基因数目

Number of up-regulated genes

下调基因数目

Number of down-regulated genes

不同日龄同一鹅种 Same goose breed with different ages
S40X vs S90X1 9128361 076
W40X vs W90X1 9959231 072
同一日龄不同鹅种 Different goose breeds with the same age
S40X vs W40X319187132
S90X vs W90X1 493858635
), ArticleFig(id=1307286470805446805, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Table 4, caption=

GO significantly enriched with DEGs in terms related to muscle formation

, figureFileSmall=null, figureFileBig=null, tableContent=

分组

Group

生物过程

Biological process

q值

q-value

基因名称

Gene name

S40X vsS90X胶原纤维组织0.02DDR2、MKX、LOC106044084、LOX、P3H1、COL2A1、ANXA2、COL5A2、LUM
S40X vsW40X钙离子稳态0.05LOC106031155、LOC106031415、CSRP3
骨骼肌卫星细胞增殖的调节0.05ANGPT1
肌丝滑动0.05TNNT2
Rho蛋白信号转导的调节0.05Anser_cygnoides_newGene_5510、PLEKHG5、ARHGEF9
肌肉组织发育的正向调控0.05PPARGC1A
血管发生的正向调控0.05ASB4
肌肉器官发育0.05MYF5、ETV1
对肌肉活动的响应0.05PPARGC1A
细胞黏附的调节0.05S1PR1、LAMA5
细胞呼吸的正向调控0.05PPARGC1A
ATP生物合成过程的正向调控0.05PPARGC1A
W40X vs W90X炎症反应0.01Anser_cygnoides_newGene_9361、IL18、LOC106029986、LOC106041040、NCF1、BMPR1B、LOC106041921、LOC106041918、NOS2、PTAFR、C3、ITGB6、KIT
W90X vsS90X炎症反应0.01Anser_cygnoides_newGene_9361、IL18、LOC106029986、LOC106041040、NCF1、LOC106041921、LOC106041918、NOS2、PTAFR、C3、KIT
), ArticleFig(id=1307286470901915798, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=表 4, caption=

GO显著富集的与成肌相关条目中的DEGs

, figureFileSmall=null, figureFileBig=null, tableContent=

分组

Group

生物过程

Biological process

q值

q-value

基因名称

Gene name

S40X vsS90X胶原纤维组织0.02DDR2、MKX、LOC106044084、LOX、P3H1、COL2A1、ANXA2、COL5A2、LUM
S40X vsW40X钙离子稳态0.05LOC106031155、LOC106031415、CSRP3
骨骼肌卫星细胞增殖的调节0.05ANGPT1
肌丝滑动0.05TNNT2
Rho蛋白信号转导的调节0.05Anser_cygnoides_newGene_5510、PLEKHG5、ARHGEF9
肌肉组织发育的正向调控0.05PPARGC1A
血管发生的正向调控0.05ASB4
肌肉器官发育0.05MYF5、ETV1
对肌肉活动的响应0.05PPARGC1A
细胞黏附的调节0.05S1PR1、LAMA5
细胞呼吸的正向调控0.05PPARGC1A
ATP生物合成过程的正向调控0.05PPARGC1A
W40X vs W90X炎症反应0.01Anser_cygnoides_newGene_9361、IL18、LOC106029986、LOC106041040、NCF1、BMPR1B、LOC106041921、LOC106041918、NOS2、PTAFR、C3、ITGB6、KIT
W90X vsS90X炎症反应0.01Anser_cygnoides_newGene_9361、IL18、LOC106029986、LOC106041040、NCF1、LOC106041921、LOC106041918、NOS2、PTAFR、C3、KIT
), ArticleFig(id=1307286470977413271, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=EN, label=Table 5, caption=

KEGG significantly enriched with DEGs in the myogenesis-related pathways

, figureFileSmall=null, figureFileBig=null, tableContent=

分组

Group

通路名称

Pathway

q值

q-value

基因名称

Gene name

A组 A group(S40X vs S90X)ECM-受体相互作用<0.01COL4A5、COL4A6、COL1A2、SDC4、CD44、COL4A2、THBS2、SDC1、VWF、ITGA9、ITGB3、COL6A6、COMP、THBS3、COL2A1、LOC106049842、TNXB、ITGB8、HMMR、CHAD、ITGB4、CD36、ITGA1、ITGA2、COL4A3、SV2B
A组 A group(S40X vsS90X)细胞黏附分子<0.01NCAM1、CADM1、ESAM、JAM3、SDC4、VCAN、NEGR1、LRRC4C、CD276、NEO1、SDC1、SDC3、ITGA9、CNTN1、NRCAM、OCLN、CLDN19、CLDN10、LOC106029335、ITGB8、CLDN5、CDH5、PTPRF、PECAM1、PTPRM、CD99、CD58、JAM2、CDH2、COLEC10
A组 A group(S40X vsS90X)黏着斑<0.01LOC106038554、COL4A5、COL4A6、COL1A2、FLT1、CAV2、CAV1、PIK3R1、COL4A2、THBS2、VWF、ITGA9、ITGB3、CCND2、COL6A6、MYLK2、COMP、THBS3、MYLK4、SHC2、VEGFA、COL2A1、LOC106049842、TNXB、ITGB8、PDGFRA、KDR、MYL2、CRKL、PDGFC、FLT4、PIK3R3、SHC4、CHAD、ITGB4、ITGA1、ITGA2、COL4A3、CAV3、FIGF、FLNB、PARVG
C组 C group(W40X vsW90X)ECM-受体相互作用<0.01COL4A5、COL4A6、COL1A2、LAMA5、SPP1、LAMB1、LAMB4、ITGA11、CD44、LAMC3、AGRN、THBS2、VWF、ITGA9、GP1BA、COL6A6、THBS3、LOC106049058、COL2A1、COL6A2、LOC106049839、LOC106049842、TNXB、ITGB5、ITGB6、GP1BB、LAMA2、LAMA4、COL6A3、CHAD、TNC、LOC106037602、COLEC10
C组 C group(W40X vsW90X)细胞因子-细胞因子受体相互作用<0.01Anser_cygnoides_newGene_1760、IL2RA、IL18、TNFRSF1B、TNFRSF8、TNFRSF6B、BMPR1B、LOC106041570、IL18R1、IL1R2、LOC106041918、LOC106041919、CXCL12、IFNAR1、IL10RB、TNFRSF18、TNFSF13B、AMH、LOC106045913、IL4R、CD40、IL1RAP、CSF1、CNTFR、LOC106048595、LOC106048600、NGFR、VEGFA、CCR7、IL1B、PDGFRA、KIT、IL17RA、TGFB2、BMPR1A、CXCL14、CX3CL1、PDGFA、LOC106035752、LOC106035885、IL2RB、IL7R、TNFSF15、IL13RA1、IL2RG、TNFRSF21、LOC106038377、LOC106038356、LOC106038357
C组 C group(W40X vsW90X)黏着斑<0.01COL4A5、COL4A6、COL1A2、PIK3CB、LAMA5、SPP1、VAV3、PIK3CG、LAMB1、LAMB4、ITGA11、LAMC3、VAV2、PIK3CD、CCND1、THBS2、LOC106044678、VWF、ITGA9、CCND3、LOC106045852、COL6A6、PIK3R5、ZYX、ACTN2、THBS3、RASGRF1、LOC106049058、SHC2、VEGFA、COL2A1、COL6A2、LOC106049839、LOC106049842、TNXB、ITGB5、ITGB6、PDGFRA、MYL2、LAMA2、LAMA4、PDGFA、BIRC2、COL6A3、PRKCA、CHAD、IGF1、RAC2、PAK7、CAV3、TNC、LOC106037602
C组 C group(W40X vsW90X)细胞黏附分子<0.01Anser_cygnoides_newGene_6728、CD2、CD80、VCAM1、NCAM1、CADM1、NLGN4X、JAM3、CD34、LOC106041540、LOC106041602、NTNG1、PTPRC、NRXN1、NEGR1、LRRC4C、PDCD1、ITGA9、LOC106046390、CD40、CNTN1、CD4、CD86、LOC106049813、LOC106029726、CD28、ITGB2、CD8A、PTPRF、COLEC10
C组 C group(W40X vsW90X)钙信号通路<0.01Anser_cygnoides_newGene_6800 、Anser_cygnoides_newGene_6806 、CAMK2D、ADCY7、HTR7、HRH1、PHKG1、P2RX1、CAMK4、ADRB2、NOS2、LOC106044837、TBXA2R、ADORA2B、CACNA1S、PTAFR、AVPR1A、PLCB2、TNNC2、PLN、PTK2B、LOC106049872、PDE1A、TACR2、PDGFRA、SLC8A3、CACNA1H、ERBB4、PRKCA、SPHK1、AGTR1、ADCY2、HTR2C、GRPR、PHKA2
C组 C group(W40X vs W90X)肌动蛋白细胞骨架的调控<0.01FGFR2、ARHGEF4、PIK3CB、ELN、VAV3、PIK3CG、IQGAP2、ITGA11、VAV2、EZR、PIK3CD、FGFR1、LOC106044678、ITGA9、LOC106045852、FGF6、PIK3R5、ACTN2、LOC106049058、FGFR4、LOC106049839、LOC106029514、ITGB5、ITGB6、ABI2、ITGB2、PDGFRA、MYL2、ENAH、CFL2、ARPC1B、FGF1、PDGFA、GNA13、RAC2、PAK7、SCIN
D组 D group(S90X vsW90X)细胞因子-细胞因子受体相互作用<0.01Anser_cygnoides_newGene_1760、TNFRSF11B、IL2RA、IL18、TNFRSF1B、TNFRSF19、LOC106041570、IL18R1、IL1R2、LOC106041918、LOC106041919、IFNAR1、IL10RB、TNFRSF18、TNFSF13B、AMH、LOC106045913、IL1RAP、CSF1、CNTFR、CCR7、IL1B、PDGFRA、KIT、TGFB2、CXCL14、FLT4、PDGFA、LOC106035752、LOC106035885、IL7R、TNFSF15、IL2RG、LOC106038356、LOC106038357
D组 D group(S90X vsW90X)细胞黏附分子0.01CD80、VCAM1、ESAM、CD34、LOC106041540、LOC106041602、PTPRC、VCAN、PDCD1、LOC106046592、CNTN1、CD4、OCLN、CD86、LOC106049813、LOC106029504、CD28、ITGB2、CDH5、JAM2、CDH2、ALCAM
), ArticleFig(id=1307286471057105048, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307007598729912781, language=CN, label=表5, caption=

KEGG显著富集的与成肌相关通路中的DEGs

, figureFileSmall=null, figureFileBig=null, tableContent=

分组

Group

通路名称

Pathway

q值

q-value

基因名称

Gene name

A组 A group(S40X vs S90X)ECM-受体相互作用<0.01COL4A5、COL4A6、COL1A2、SDC4、CD44、COL4A2、THBS2、SDC1、VWF、ITGA9、ITGB3、COL6A6、COMP、THBS3、COL2A1、LOC106049842、TNXB、ITGB8、HMMR、CHAD、ITGB4、CD36、ITGA1、ITGA2、COL4A3、SV2B
A组 A group(S40X vsS90X)细胞黏附分子<0.01NCAM1、CADM1、ESAM、JAM3、SDC4、VCAN、NEGR1、LRRC4C、CD276、NEO1、SDC1、SDC3、ITGA9、CNTN1、NRCAM、OCLN、CLDN19、CLDN10、LOC106029335、ITGB8、CLDN5、CDH5、PTPRF、PECAM1、PTPRM、CD99、CD58、JAM2、CDH2、COLEC10
A组 A group(S40X vsS90X)黏着斑<0.01LOC106038554、COL4A5、COL4A6、COL1A2、FLT1、CAV2、CAV1、PIK3R1、COL4A2、THBS2、VWF、ITGA9、ITGB3、CCND2、COL6A6、MYLK2、COMP、THBS3、MYLK4、SHC2、VEGFA、COL2A1、LOC106049842、TNXB、ITGB8、PDGFRA、KDR、MYL2、CRKL、PDGFC、FLT4、PIK3R3、SHC4、CHAD、ITGB4、ITGA1、ITGA2、COL4A3、CAV3、FIGF、FLNB、PARVG
C组 C group(W40X vsW90X)ECM-受体相互作用<0.01COL4A5、COL4A6、COL1A2、LAMA5、SPP1、LAMB1、LAMB4、ITGA11、CD44、LAMC3、AGRN、THBS2、VWF、ITGA9、GP1BA、COL6A6、THBS3、LOC106049058、COL2A1、COL6A2、LOC106049839、LOC106049842、TNXB、ITGB5、ITGB6、GP1BB、LAMA2、LAMA4、COL6A3、CHAD、TNC、LOC106037602、COLEC10
C组 C group(W40X vsW90X)细胞因子-细胞因子受体相互作用<0.01Anser_cygnoides_newGene_1760、IL2RA、IL18、TNFRSF1B、TNFRSF8、TNFRSF6B、BMPR1B、LOC106041570、IL18R1、IL1R2、LOC106041918、LOC106041919、CXCL12、IFNAR1、IL10RB、TNFRSF18、TNFSF13B、AMH、LOC106045913、IL4R、CD40、IL1RAP、CSF1、CNTFR、LOC106048595、LOC106048600、NGFR、VEGFA、CCR7、IL1B、PDGFRA、KIT、IL17RA、TGFB2、BMPR1A、CXCL14、CX3CL1、PDGFA、LOC106035752、LOC106035885、IL2RB、IL7R、TNFSF15、IL13RA1、IL2RG、TNFRSF21、LOC106038377、LOC106038356、LOC106038357
C组 C group(W40X vsW90X)黏着斑<0.01COL4A5、COL4A6、COL1A2、PIK3CB、LAMA5、SPP1、VAV3、PIK3CG、LAMB1、LAMB4、ITGA11、LAMC3、VAV2、PIK3CD、CCND1、THBS2、LOC106044678、VWF、ITGA9、CCND3、LOC106045852、COL6A6、PIK3R5、ZYX、ACTN2、THBS3、RASGRF1、LOC106049058、SHC2、VEGFA、COL2A1、COL6A2、LOC106049839、LOC106049842、TNXB、ITGB5、ITGB6、PDGFRA、MYL2、LAMA2、LAMA4、PDGFA、BIRC2、COL6A3、PRKCA、CHAD、IGF1、RAC2、PAK7、CAV3、TNC、LOC106037602
C组 C group(W40X vsW90X)细胞黏附分子<0.01Anser_cygnoides_newGene_6728、CD2、CD80、VCAM1、NCAM1、CADM1、NLGN4X、JAM3、CD34、LOC106041540、LOC106041602、NTNG1、PTPRC、NRXN1、NEGR1、LRRC4C、PDCD1、ITGA9、LOC106046390、CD40、CNTN1、CD4、CD86、LOC106049813、LOC106029726、CD28、ITGB2、CD8A、PTPRF、COLEC10
C组 C group(W40X vsW90X)钙信号通路<0.01Anser_cygnoides_newGene_6800 、Anser_cygnoides_newGene_6806 、CAMK2D、ADCY7、HTR7、HRH1、PHKG1、P2RX1、CAMK4、ADRB2、NOS2、LOC106044837、TBXA2R、ADORA2B、CACNA1S、PTAFR、AVPR1A、PLCB2、TNNC2、PLN、PTK2B、LOC106049872、PDE1A、TACR2、PDGFRA、SLC8A3、CACNA1H、ERBB4、PRKCA、SPHK1、AGTR1、ADCY2、HTR2C、GRPR、PHKA2
C组 C group(W40X vs W90X)肌动蛋白细胞骨架的调控<0.01FGFR2、ARHGEF4、PIK3CB、ELN、VAV3、PIK3CG、IQGAP2、ITGA11、VAV2、EZR、PIK3CD、FGFR1、LOC106044678、ITGA9、LOC106045852、FGF6、PIK3R5、ACTN2、LOC106049058、FGFR4、LOC106049839、LOC106029514、ITGB5、ITGB6、ABI2、ITGB2、PDGFRA、MYL2、ENAH、CFL2、ARPC1B、FGF1、PDGFA、GNA13、RAC2、PAK7、SCIN
D组 D group(S90X vsW90X)细胞因子-细胞因子受体相互作用<0.01Anser_cygnoides_newGene_1760、TNFRSF11B、IL2RA、IL18、TNFRSF1B、TNFRSF19、LOC106041570、IL18R1、IL1R2、LOC106041918、LOC106041919、IFNAR1、IL10RB、TNFRSF18、TNFSF13B、AMH、LOC106045913、IL1RAP、CSF1、CNTFR、CCR7、IL1B、PDGFRA、KIT、TGFB2、CXCL14、FLT4、PDGFA、LOC106035752、LOC106035885、IL7R、TNFSF15、IL2RG、LOC106038356、LOC106038357
D组 D group(S90X vsW90X)细胞黏附分子0.01CD80、VCAM1、ESAM、CD34、LOC106041540、LOC106041602、PTPRC、VCAN、PDCD1、LOC106046592、CNTN1、CD4、OCLN、CD86、LOC106049813、LOC106029504、CD28、ITGB2、CDH5、JAM2、CDH2、ALCAM
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狮头鹅和乌鬃鹅胸肌组织形态学及转录组比较分析
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谢泽婷 , 侯晓韵 , 詹胤锴 , 林随源 , 朱溥睿 , 郭诗琦 , 秦双 , 罗阳冰 , 李秀金 , 黄运茂 , 伍仲平 , 张续勐
畜牧兽医学报 | 遗传育种 2026,57(1): 206-220
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畜牧兽医学报 |遗传育种 2026 , 57 (1) : 206 -220
狮头鹅和乌鬃鹅胸肌组织形态学及转录组比较分析
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谢泽婷与侯晓韵为同等贡献作者

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谢泽婷与侯晓韵为同等贡献作者

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侯晓韵,本科生,主要从事动物遗传育种研究,E-mail:

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侯晓韵,本科生,主要从事动物遗传育种研究,E-mail:

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谢泽婷, 侯晓韵 , 詹胤锴, 林随源, 朱溥睿, 郭诗琦, 秦双, 罗阳冰, 李秀金, 黄运茂, 伍仲平 , 张续勐
作者信息
  • 仲恺农业工程学院动物科技学院,广州 510225
通讯作者:
伍仲平,主要从事动物遗传育种研究,E-mail:
张续勐,主要从事动物遗传育种研究,E-mail:
作者简介:

谢泽婷,本科生,主要从事动物遗传育种研究,E-mail:

谢泽婷与侯晓韵为同等贡献作者

侯晓韵,本科生,主要从事动物遗传育种研究,E-mail:

Comparative Analysis of Histomorphology and Transcriptome in Pectoral Muscle Tissues between Shitou and Wuzong Geese
Zeting XIE, Xiaoyun HOU , Yinkai ZHAN, Suiyuan LIN, Purui ZHU, Shiqi GUO, Shuang QIN, Yangbing LUO, Xiujin LI, Yunmao HUANG, Zhongping WU , Xumeng ZHANG
Affiliations
  • College of Animal Sciences and Technology,Zhongkai University of Agriculture and Engineering,Guangzhou 510225,China
  • Corresponding Author:
    WU Zhongping, E-mail:
    ZHANG Xumeng, E-mail:
出版时间: 2026-01-23 doi: 10.11843/j.issn.0366-6964.2026.01.018
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旨在对比狮头鹅和乌鬃鹅在40和90日龄时,其胸肌肌纤维形态和转录组的差异,挖掘导致不同日龄、不同鹅种胸肌生长差异的候选基因。本研究随机选取同批次健康的40、90日龄的狮头鹅和乌鬃鹅公鹅各3只,采集胸肌组织,利用H&E染色分析肌纤维形态差异,结合转录组测序筛选出与肌肉生长相关的差异表达基因(DEGs),并采用实时荧光定量PCR(RT-qPCR)验证关键候选基因。H&E染色结果发现,40日龄时,狮头鹅的胸肌肌纤维密度极显著高于乌鬃鹅(P<0.001),胸肌肌纤维直径极显著低于乌鬃鹅(P<0.001),而90日龄时,狮头鹅的胸肌肌纤维密度显著低于乌鬃鹅(P<0.05),胸肌纤维直径无显著差异(P>0.05)。胸肌转录组测序共筛选出DEGs 5 719个,不同日龄同一鹅种间的DEGs 3 907个,同一日龄不同鹅种间的DEGs 1 812个。对胸肌DEGs进行KEGG通路富集分析发现,ECM-受体相互作用、细胞黏附分子和黏着斑等6个通路在不同日龄同一鹅种间胸肌生长中发挥关键调控作用,细胞因子-细胞因子受体相互作用和细胞黏附分子通路在同一日龄不同鹅种间胸肌生长中发挥关键调控作用,并从中筛选出8个与胸肌生长发育密切相关的候选基因,分别是COL4A2、ITGB3、CAV1、CCND2、MYLK2、PDGFRA、ACTN2和TNNC2。通过对上述基因进行RT-qPCR验证,发现其结果与转录组测序结果相关系数均在0.90以上,说明测序结果可靠。综上,本研究发现,不同日龄及品种间的肌纤维形态差异显著,黏着斑通路、ECM-受体相互作用和细胞黏附分子等通路可通过调控COL4A2、ITGB3和CAV1等8个候选基因表达介导两品种胸肌肌纤维的肥大过程。本研究将为肉鹅分子标记辅助育种提供数据基础,进而推动肉鹅遗传改良和新品种培育。

狮头鹅  /  乌鬃鹅  /  胸肌  /  肌肉生长  /  差异表达基因

This study aimed to compare the morphological characteristics of pectoral muscle myofibers and transcriptomic profiles between Shitou geese and Wuzong geese at 40- and 90-day-old stages, and to identify candidate genes underlying divergent breast muscle growth across breeds and ages. A total of 3 male Shitou geese and 3 male Wuzong geese from the same batch were randomly selected at 40 and 90 days of age. Pectoral muscle tissues were collected for morphological analysis of muscle fibers using H&E staining. Transcriptome sequencing was performed to identify differentially expressed genes (DEGs) related to muscle growth, and key candidate genes were further validated by reverse transcription quantitative PCR (RT‑qPCR). H&E staining results revealed that at 40 days of age, the density of pectoral muscle fibers in Shitou geese was significantly higher than that in Wuzong geese (P<0.001), while the diameter of pectoral muscle fibers was significantly smaller (P<0.001). At 90 days of age, however, the density of pectoral muscle fibers in Shitou geese was significantly lower than that in Wuzong geese (P<0.05), and no significant difference was observed in pectoral muscle fiber diameter between the two breeds (P>0.05). Transcriptome sequencing of pectoral muscle identified 5 719 DEGs in total, including 3 907 DEGs between different ages within the same breed and 1 812 DEGs between the two breeds at the same age. KEGG enrichment analysis revealed that pathways including ECM-receptor interaction, cell adhesion molecules, and focal adhesion critically regulated pectoral muscle growth across ages within breeds. Cytokine-cytokine receptor interaction and cell adhesion molecule pathways were key regulators of growth differences between breeds at the same age. Eight candidate genes (COL4A2, ITGB3, CAV1, CCND2, MYLK2, PDGFRA, ACTN2, and TNNC2) closely associated with pectoral muscle development were identified. RT-qPCR validation confirmed high correlation (r>0.90) with transcriptome data, supporting the reliability of sequencing results. In summary, this study revealed significant variations in myofiber morphology across developmental stages and between Shitou and Wuzong goose breeds. Key pathways including focal adhesion, ECM-receptor interaction, and cell adhesion molecules were found to mediate pectoral muscle myofibre hypertrophy through regulating the expression of candidate genes (e.g., COL4A2, ITGB3, CAV1, etc.). These findings establish a theoretical foundation for molecular marker-assisted breeding in meat geese, accelerating genetic improvement programs and new breed development.

Shitou goose  /  Wuzong goose  /  pectoral muscles  /  muscle growth  /  differentially expressed genes
谢泽婷, 侯晓韵, 詹胤锴, 林随源, 朱溥睿, 郭诗琦, 秦双, 罗阳冰, 李秀金, 黄运茂, 伍仲平, 张续勐. 狮头鹅和乌鬃鹅胸肌组织形态学及转录组比较分析. 畜牧兽医学报, 2026 , 57 (1) : 206 -220 . DOI: 10.11843/j.issn.0366-6964.2026.01.018
Zeting XIE, Xiaoyun HOU, Yinkai ZHAN, Suiyuan LIN, Purui ZHU, Shiqi GUO, Shuang QIN, Yangbing LUO, Xiujin LI, Yunmao HUANG, Zhongping WU, Xumeng ZHANG. Comparative Analysis of Histomorphology and Transcriptome in Pectoral Muscle Tissues between Shitou and Wuzong Geese[J]. Acta Veterinaria et Zootechnica Sinica, 2026 , 57 (1) : 206 -220 . DOI: 10.11843/j.issn.0366-6964.2026.01.018
鹅作为我国重要的经济家禽,其在2024年的出栏量达到5.69亿只,产值高达592亿元[1]。但其在生产上仍面临新品种选育程度不够、育种方式落后等问题[1]。而胸肌作为肉鹅主要产肉部位,其胚胎期时成肌细胞增殖的程度直接决定了出生后肌纤维数目,而出生后肌纤维生长和肌内脂肪沉积直接影响产肉效率与肉品质[2⁃3]。通过对比分析不同性别、日龄、体型及生长速度肉鹅胸肌肌肉组织形态差异,并借助转录组测序精准筛选出与肌肉生长发育相关的关键基因,可为肉鹅分子标记辅助育种提供靶基因,实现对肉鹅种资资源的高效利用[1,4⁃6]。
值得注意的是,在肉鹅实际养殖生产过程中可划分为育雏期(0~4周龄)、育肥期(5~8周龄)和育成期(9~26周龄)3个阶段[7]。其中,处于育肥期40日龄的肉鹅生长速度最快,日增重可达到100 g,而育成期90日龄时肉鹅的肌纤维生长趋于成熟,肌束间脂肪沉积,风味物质积累,是肉鹅最佳上市日龄[8⁃9]。肉鹅阶段生长特征与肌肉组织形态及分子调控机制密切相关,目前已有多项研究通过组织形态学和转录组学探究不同日龄、不同鹅种肌肉生长发育的分子机制。Yang等[4]通过对0、14、28、42、56和70日龄的雄性和雌性浙江白鹅的胸肌和腿肌进行性别因素比较分析,发现70日龄雄性浙江白鹅胸肌和腿肌肌纤维直径和面积显著高于雌性,且IGF1、GHRHR和NCAPG-LCORL基因在不同性别肉鹅肌肉生长发育中有着重要的调控作用。Zhao等[5]对2、4、6、8和10周龄的伊犁鹅腿肌进行组织形态和日龄因素的转录组比较分析,发现伊犁鹅腿肌肌纤维直径和横截面积随日龄增加而增大,而肌纤维密度随日龄增加而下降,且粘附斑块、ECM-受体相互作用和紧密连接等信号通路和ITGB3、ITGB6和ACTB等基因在伊犁鹅肌肉纤维生长发育中发挥着重要的调控作用。Hu等[6]对6、10和30周龄生长速度较快的朗德鹅和生长速度较慢的四川白鹅的胸肌进行品种内和品种间转录组比较分析,筛选出肌肉肥大相关基因CTB、IGF1、GHRHR和NCAPG-LCOR。狮头鹅和乌鬃鹅都是广东的地方品种,二者体型和肉质差异较大,是研究不同体型鹅种肌肉生长发育分子机制的良好试验模型[10⁃11]。Tang等[12]对5周龄的狮头鹅和乌鬃鹅腿肌进行转录组测序,发现CXCL12、SSTR4、FABP5等87个差异表达基因可能与腿肌生长有关。本研究团队前期对狮头鹅和乌鬃鹅胸肌的研究结果显示,二者发育模式存在较大差异,在胚胎15日龄时狮头鹅的胸肌肌纤维密度和直径显著高于乌鬃鹅,且MYL2、MYL3、TNNI2、TNNC2、CAV3、CACNA1S和NOS1基因共同调控狮头鹅和乌鬃鹅胚胎期胸肌的发育[13]。虽然目前已有多项关于鹅肌肉生长发育的研究,并筛选出多个候选基因,但目前关于不同体型鹅种出生后胸肌生长的具体调控机制尚不清晰。
为进一步探究导致乌鬃鹅与狮头鹅胸肌生长差异的分子机制,本试验以40、90日龄乌鬃鹅与狮头鹅为研究对象,通过组织形态学分析胸肌的肌纤维密度、直径,结合转录组测序技术和GO、KEGG富集分析,筛选出40、90日龄狮头鹅和乌鬃鹅胸肌中与肌肉生长相关的关键基因并分析其生物学功能。本研究将为肉鹅分子标记辅助育种提供靶基因,并为肉鹅的遗传改良和新品种培育提供借鉴。
本试验选取同批次健康的40、90日龄狮头鹅、乌鬃鹅公鹅各3只,由清远市金羽丰鹅业有限公司提供,所有试验鹅饲养管理条件相似。在禁食12 h后,采用颈静脉放血法处死,采集同侧胸肌组织,一部分立即投入液氮速冻,后转入-80 ℃保存,一部分置于4%的多聚甲醛溶液中固定,用于H&E染色。
取相同位置相同大小胸肌组织样本,置于4%多聚甲醛溶液中固定24~48 h,再经乙醇梯度脱水、透明、浸蜡和石蜡包埋处理后,使用切片机将包埋组织切成4 μm厚的切片,然后利用苏木素、伊红及相应活化剂进行染色。风干后滴20 μL中性树脂,并用镊子小心盖上盖玻片封片。将制备好的切片先经过CaseViewer(v2.2)扫描软件获取20.0倍放大图像后,正置于Axio Imager Z1显微镜(Carl Zeiss AG,Germany),以400倍放大倍数拍摄并采集每个区域的显微图像。每张切片随机选取3个非重叠视野,使用Adobe Photoshop(v2021)软件统计各个视野肌纤维数量,并随机选择6根肌纤维测量其直径,然后按照公式(1)、(2)计算肌纤维密度和肌纤维直径。
肌纤维密度(根·mm-2)= 肌纤维数量总和视野面积×3
肌纤维直径(mm)= 肌纤维直径总和肌纤维数量
RNA文库构建及Illumina HiSeqTM 2000高通量测序由北京百迈客生物科技有限公司(Biomarker Technologies)完成。按照Trizol试剂制造商提供的操作规程,采用Trizol法从冷冻胸肌组织样本中提取总RNA。用Agilent 2100生物分析仪(Agilent Technologies,USA)评估RNA的质量和完整性(RNA integrity number,RIN)。分别从40、90日龄的狮头鹅和乌鬃鹅各3份独立肌肉组织中筛选RIN大于6的RNA进行文库构建。利用Illumina HiSeqTM2000平台(Illumina,USA)生成大量的FASTQ格式的测序原始数据(raw read),使用Fastp软件去除N的比例大于10%的Read和质量值Q≤10的碱基数占整条Read的50%以上的Reads,同时计算Phred碱基质量值大于20的碱基数占总碱基数(Clean data)的比例(Q20)、GC含量和序列重复水平,得到质控数据(Clean read)。通过HISAT2(v2.2.1)和StringTie软件(v2.2.3),以中国家鹅(Anser cygnoides domesticus)基因组数据(版本为AnsCyg_PRJNA183603_v1.0)为参考基因组,对Reads进行组装、定量和文库饱和度分析。
通过StringTie(v2.2.3)软件的最大流量算法,对样品中的Mapped Reads数目和转录本长度进行归一化,采用每千碱基每百万片段数(fragments per kilobase of transcript per million fragments mapped,FPKM)作为衡量基因表达水平的指标,其计算公式见下方。使用Origin(v2024)软件计算斯皮尔曼相关系数(Spearman’s correlation coefficient),评估各样本间的生物学重复相关性后,使用R(v2024)软件进行热图聚类分析(Heatmap)和两两一组差异表达分析,并以|Fold Change|≥2且错误发现率(false discovery rate,FDR)<0.01为阈值筛选DEGs。接着利用GO数据库和KEGG数据库对DEGs进行GO功能分类和信号通路富集分析。最后对q<0.05的GO、KEGG通路中的DEGs进行注释,结合肌肉生长发育的相关文献,从中筛选出与肌肉生长有关的基因。
FPKM=cDNA FragmentsMapped FragmentsMillions*Transcript Length(kb)
公式中,cDNA Fragments表示比对到某一转录本上的片段数目,即双端Reads数目;Mapped Fragments(Millions)表示比对到转录本上的片段总数,以106为单位;Transcript Length(kb)表示转录本长度,以103个碱基为单位。
通过Trizol法用RNA提取测序分析的相同样本中的总RNA,按照ReverTra Ace qPCR RT Master Mix with gDNA Remover逆转录试剂盒(TOYOBO,JPN)说明书将RNA逆转录为cDNA。通过与NCBI GenBank数据库中的序列比对,使用Primer Premier 5.0软件设计目标基因及内参基因β-actin的引物(表1)。采用SYBR Select Master Mix(Thermo Fisher Scientific,USA),按说明书配制10 μL反应体系。在QuantStudio 7 Flex实时定量PCR仪(Thermo Fisher Scientific,USA)上运行反应程序,利用RT-qPCR方法检测基因表达,采用2-ΔΔCT法计算相对mRNA的表达量。
使用SPSS(v27.0)和GraphPad Prism(v7.0)软件进行统计分析及作图,数据表示为“平均值±标准差(SD)”。肌纤维直径、密度数据经过One-way ANOVA的方法对比平均值后,用Duncan多重比较检验组间差异,RT-qPCR结果与转录组测序结果的相关性则采用皮尔逊相关分析法计算相关系数。统计图中*表示不同组间差异显著(P<0.05),**表示不同组间差异极显著(P<0.01),***表示不同组间差异极其显著(P<0.001)。
通过H&E染色对40、90日龄乌鬃鹅和狮头鹅胸肌肌纤维进行形态学观察发现,胸肌肌纤维40日龄时横切面多呈圆形或椭圆形,少数为多边形;90日龄时,横切面以多边形为主,少数为圆形或椭圆形;细胞核经苏木素染色呈蓝紫色,呈扁平椭圆形,位于细胞边缘。计算肌纤维密度和直径后发现,40日龄时,狮头鹅胸肌肌纤维密度极显著高于乌鬃鹅(P<0.001),胸肌肌纤维直径极显著低于乌鬃鹅(P<0.001),而90日龄时,狮头鹅胸肌肌纤维密度显著低于乌鬃鹅(P<0.05),胸肌肌纤维直径无显著差异(P>0.05)(图1)。以上结果表明40、90日龄两鹅种胸肌生长差异明显。
对40、90日龄狮头鹅和乌鬃鹅的胸肌共12个样本进行转录组测序,共得到48.586 Gb Clean reads,各样品GC碱基含量为51.01%~52.89%,Q30碱基百分比≥92.32%(表2)。通过HISAT2和StringTie软件将过滤后各样品的clean Reads与中国家鹅(Anser cygnoides domesticus)参考基因组(版本:AnsCyg_PRJNA183603_v1.0)进行序列比对。结果显示,各样品的Mapped Reads与参考基因组的比对效率为76.04%~79.57%(表2)。上述结果表明,转录组测序数据质量较高,可用于后续试验进一步分析。
计算所有基因的FPKM值并利用Spearman相关系数评估样本之间的相关性,结果显示,各样本之间的相关系数≥0.87(图2),说明样本之间的相关性较强,表达模式存在较高相似性。基于FPKM值进行聚类分析,结果显示,40和90日龄的狮头鹅与乌鬃鹅胸肌样本分别聚集在一起(图3)。
通过使用R(v2024)软件DEseq2包对转录组测序数据进行差异表达分析,将|Fold Change|≥2且错误发现率(false discovery rate,FDR)<0.01为阈值筛选DEGs。共筛选出DEGs 5 719个,其中上调基因2 804个,下调基因2 915个,不同日龄同一鹅种间的DEGs 3 907个,其中上调基因1 759个,下调基因2 148个,同一日龄不同鹅种间的DEGs 1 812个,其中上调基因1 045个,下调基因767个(表3)。上述结果表明,生长阶段对基因表达的影响大于品种本身的遗传差异。
运用GO数据库进行对各组的DEGs富集分析结果发现,S40X vs S90X显著富集在胶原纤维组织(collagen fibril organization)条目,S40X vs W40X显著富集在细胞钙离子稳态(cellular calcium ion homeostasis)、骨骼肌卫星细胞增殖的调节(regulation of skeletal muscle satellite cell proliferation)、肌丝滑动(muscle filament sliding)等11条与肌肉生长发育相关的生物过程条目,W40X vs W90X和S90X vs W90X都显著富集到炎症反应(inflammatory response)条目中(表4)。
为进一步了解不同DEGs在40、90日龄狮头鹅与乌鬃鹅胸肌中的功能,利用KEGG数据库对各组的DEGs进行通路富集分析。选取每组显著性q值最小的前20个通路作散点图(图4)。从图中可以看出,A组(S40X vs S90X)、B组(S40X vs W40X)、C组(W40X vs W90X)和D组(S90X vs W90X)分别有10、0、9、6条通路显著富集(q<0.05)。其中筛选出6条与肌肉生长发育相关的信号通路,分别是细胞外基质-受体相互作用(ECM-receptor interaction)、细胞粘附分子(cell adhesion molecules (CAMs))、局灶性黏附(focal adhesion)、细胞因子-细胞因子受体相互作用(cytokine-cytokine receptor interaction)、钙信号通路(calcium signaling pathway)、肌动蛋白细胞骨架调控(regulation of actin cytoskeleton)。推测这些通路和通路中的DEGs对狮头鹅和乌鬃鹅的生长发育起着重要的调控作用(图4、表5)。
为进一步验证转录组测序数据的准确性,本试验利用RT-qPCR对8个与狮头鹅和乌鬃鹅胸肌生长相关的DEGs(COL4A2、ITGB3、CAV1、CCND2、MYLK2、PDGFRA、ACTN2和TNNC2)进行验证(图5)。结果表明,8个基因的mRNA表达量变化趋势都与转录组测序结果一致(r≥0.90),表明测序数据可靠。
胸肌作为禽类重要的产肉组织,由大量肌纤维组成,在肌生成过程中肌纤维的生长发育可分为胚胎期和出生后两个阶段[14⁃15]。胚胎期肌源性前体细胞迁移、增殖并分化为成肌细胞,一部分融合形成成熟肌管,最终发育为成熟的肌纤维;另一部分成肌细胞不发生融合,而在肌纤维表面形成保持增殖能力的卫星细胞,在出生后与肌纤维融合,增强肌纤维蛋白合成与生长能力,使肌纤维体积不断增大[16⁃17]。肌纤维形态结构的变化不仅决定了肌肉的生长发育,还决定了禽类的产肉量及肉品质[18]。值得注意的是,狮头鹅体型大、肉质紧实,乌鬃鹅体型小、肉质细嫩,是研究不同体型鹅种肌肉生长发育分子机制的良好试验模型[10, 11]。此前,本研究团队对胚胎期狮头鹅和乌鬃鹅肌肉的研究发现,转录起始位点(transcription start sites,TSS)周围生肌基因的早期去甲基化和MyoD1启动子的去甲基化与狮头鹅腿肌发育快于乌鬃鹅密切相关,且MYL2、MYL3、TNNI2、TNNC2、CAV3、CACNA1S和NOS1基因参与调控胸肌发育[13,19-20]。但目前关于狮头鹅和乌鬃鹅出生后胸肌肥大的分子机制研究还处于空白状态。因此,本研究旨在对出生后两鹅种胸肌肥大进行系统探究以填补这一空白。
出生后肌肉生长主要表现为肌纤维肥大以及肌纤维类型发生转化,通过H&E染色切片有助于了解狮头鹅和乌鬃鹅胸肌肌肉的生长模式[21]。本研究结果发现,两鹅种在40~90日龄期间胸肌生长明显,狮头鹅肌纤维直径增大的速度显著快于乌鬃鹅,且90日龄时狮头鹅胸肌肌纤维密度显著低于乌鬃鹅,但胸肌肌纤维直径差异不明显。这与王金辉等[22]的研究结果相符,即乌鬃鹅胸肌和腿肌肌肉纤维在1~40日龄生长最迅速,但40日龄后肌纤维生长潜力不如狮头鹅。Zhang等[13]通过将胚胎期狮头鹅和乌鬃鹅胸肌肌纤维形态进行对比,发现胚胎期15日龄时狮头鹅胸肌肌纤维密度和直径显著高于乌鬃鹅,而胚胎期23胚龄~1日龄两鹅种胸肌肌纤维直径差异不显著。王金辉等[20]研究发现,胚胎期15日龄时狮头鹅腿肌肌纤维直径和密度明显高于乌鬃鹅。肌肉纤维的变化与机体生长发育密切相关,Tang等[12]通过Logistic的生长曲线参数模型模拟1~84日龄狮头鹅和乌鬃鹅体型和体重变化,发现狮头鹅的生长高峰出现在14~63日龄,乌鬃鹅的生长高峰出现在7~49日龄。综上表明,狮头鹅和乌鬃鹅的肌肉生长发育具有明显的时序和品种差异,胚胎期15胚龄~1日龄和40~90日龄都是狮头鹅和乌鬃鹅胸肌肌肉生长的关键时期,且狮头鹅较乌鬃鹅的生长高峰晚并具有更强的肌肉生长潜力,但其生长速率可能在90日龄后开始放缓。
肌肉生长是一个复杂的过程,受多种因素的影响,基因及信号转导途径在其中发挥至关重要的作用[23]。40、90日龄狮头鹅和乌鬃鹅(即A组(S40X vs S90X)和C组(W40X vs W90X))在转录组测序共筛选出3 907个DEGs,GO功能和KEGG通路富集分析结果显示,DEGs主要富集在胶原纤维组织条目、炎症反应条目和ECM-受体相互作用、细胞黏附分子、黏着斑通路、细胞因子-细胞因子受体相互作用、钙信号和肌动蛋白细胞骨架通路中。这与Hu等[6]发现黏着斑通路是调控6、10、30周龄的朗德鹅(生长速度较快)和四川白鹅(生长速度较慢)胸肌生长发育的关键通路的研究结果基本相符。武敏等[18]也发现,ECM-受体相互作用和黏着斑通路在不同生长速度黄羽肉鸡胸肌肌肉生长发育中具有重要作用。这与Zhang等[13]研究在胚胎期的发现有所差异,即狮头鹅和乌鬃鹅胸肌胚胎15、23胚龄的DEGs主要富集于早期发育通路和代谢通路。已有研究发现,细胞黏附分子可以通过介导肌细胞与胶原纤维组织、胶原蛋白、蛋白多糖等细胞外基质的黏附作用,随肌肉的生长改变肌肉的形态[24⁃26]。也有研究发现,细胞因子-细胞因子受体相互作用能够动态调节细胞内Ca2+浓度来维持细胞内稳态,进而调控肌球蛋白收缩性,促进肌肉细胞增殖与分化[27⁃29]。综上表明,在胚胎期狮头鹅和乌鬃鹅胸肌发育的差异主要由调控肌纤维发生的核心通路驱动,出生后两品种间的肌肉生长差异则主要由与肌纤维成熟有关的通路调控。
通路中显著富集的COL4A2、ITGB3、CAV1、CCND2、MYLK2、PDGFRA、ACTN2和TNNC2基因在胸肌快速生长过程中发挥关键调控作用。其中COL1A2基因可通过编码I型胶原蛋白α2链调控I型胶原蛋白的合成与组装,改变细胞外基质组成和ECM-受体信号传导,影响肌肉生长[18]。这与赵迪等[30]对天农麻鸡胴体性状调控机制的研究发现相似。ITGB3基因所编码整合素β3亚基可与αv亚基共同组成αvβ3整合素,介导肌纤维之间的黏附和信号转导,促进肌肉肥大[31]。这与Zhao 等[5]发现ITGB3基因是与调控2、4周龄伊犁鹅肌肉生长相关的重要调控基因的研究结果一致。CAV1编码小窝蛋白1是一种重要的脂肪酸结合蛋白,能将游离脂肪酸从质膜转移到脂滴内,促进脂质沉积和肌细胞分化[3]。这与CAV1在钢鹅肌纤维发育和肌内脂肪沉积过程中发挥重要作用的结果相似[3]。CCND2编码的细胞周期蛋白D2是细胞周期调控的关键效应因子,可以通过与周期蛋白依赖激酶4和周期蛋白依赖激酶6的结合控制G1期向S期的转换,促进肌肉细胞增殖[32]。这与Hu等[6]在四川鹅和朗德鹅的研究结果不同:CCND2基因通过增加脂质沉积和减少白细胞浸润来促进肌肉内细胞的增殖,从而调节胸肌的肥大。推测与试验所用的鹅种不同有关。MYLK2编码的肌球蛋白轻链激酶2,可通过催化肌球蛋白调节轻链丝氨酸残端的磷酸化,增强肌球蛋白ATP酶活性,从而促进肌球蛋白与肌动蛋白的相互作用,维持肌纤维结构与功能[5]。Zhao等[5]研究发现,伊犁鹅腿肌中MYLK2基因同样具有维持肌纤维结构与功能的作用。PDGFRA所编码血小板衍生生长因子受体α与血小板衍生生长因子结合后,发生二聚化和磷酸化反应,激活下游特定的胞内分子,调控肌纤维生长、增殖和分化[33]。与Huang等[34]的研究结果一致:PDGFRA基因与狮头鹅肌肉生长发育密切相关。CACNA1S基因编码电压门控钙离子通道α亚基通过调控肌肉收缩、线粒体功能及骨骼肌能量代谢,参与动物肌肉生长调控[35⁃36]。这与Zhang等[13]发现CACNA1S基因在乌鬃鹅胚胎期腿肌发育中发挥关键作用的结果相符。张军等[35]研究也发现CACNA1S基因可能是影响鸡胸肌发育的关键基因。TNNC2基因编码的快速骨骼肌肌钙蛋白,主要调控骨骼肌快肌收缩,其能够抑制Ca2+浓度,与肌球蛋白共同调节钙离子对肌动蛋白ATP酶的活性,在细胞器转运和肌肉收缩等多个细胞进程中都起着关键作用[37]。Zhang等[13]发现TNNC2基因在乌鬃鹅胚胎期腿肌发育中挥发关键作用。综上,肌肉生长相关基因COL4A2、ITGB3、CAV1、MYLK2、PDGFRA、ACTN2和TNNC2在狮头鹅与伊犁鹅、四川白鹅、朗德鹅等中表现出功能保守性,其可能通过改变ECM的组成调控胞内信号传导、细胞外基质重塑、肌纤维结构及代谢稳态,促进肌纤维增殖与肥大。而CCND2基因的调控模式在不同鹅种间存在显著差异,这可能与不同品种间的进化分化有关。
将40、90日龄狮头鹅和乌鬃鹅(即B组(S40X vs W40X)和D组(S90X vs W90X))DEGs进行GO功能和KEGG通路富集分析,发现DEGs显著富集在细胞钙离子稳态、骨骼肌卫星细胞增殖调控等条目和细胞因子-细胞因受体相互作用和细胞黏附分子通路中。已有研究发现,细胞因子-细胞因受体相互作用通路能够通过调控细胞间重要的调节因子,介导细胞与细胞、细胞与细胞外间质之间的黏附及信号传导,参与肌肉的生长、分化[38]。其中PDGFRA基因可能通过调控血小板衍生生长因子受体α与血小板衍生生长因子间的相互作用,激活下游特定的胞内分子,引起不同信号通路的级联反应,诱导肌卫星细胞或间充质干细胞分化成脂肪细胞,促进肌内脂肪的沉积,使得狮头鹅和乌鬃鹅胸肌肉质出现分化[33]。
目前,我国肉鹅品种、品系选育和配套系杂交利用尚处初期,缺乏系统的育种配套选育方法和高效的分子育种技术,难以协同改良多个关键经济性状,使得鹅的体重生长与繁殖性能、生长速度与肉品质及抗逆性的矛盾日益突出[1,39]。而本研究借助转录组测序分析40、90日龄狮头鹅和乌鬃鹅各基因间的相互作用和调控,准确挖掘出COL4A2、ITGB3、CAV1等8个与肌肉生长相关联的基因,既可为分子育种提供靶标,提高鹅产肉性状选择的准确性,提高育种效率,从而满足市场对于良种肉鹅的需求,还可为平衡改良体重生长、生长速度、肉品质和繁殖性状之间的矛盾提供借鉴[40]。然而由于肌肉生长是一个动态过程,而本研究只解析了狮头鹅和乌鬃鹅40、90日龄胸肌中基因转录情况,难以捕捉发育全程的基因表达动态和完全反映翻译后蛋白质的功能和活性[41, 42]。因此,后面还需结合蛋白质组学、代谢组学和单细胞转录组测序进行多组学联合分析,并利用干扰、过表达技术,在体外细胞试验或动物模型中进行基因干扰或过表达试验,进一步验证候选基因的功能、调控机制及其与环境的相互作用[43]。
本研究发现,狮头鹅和乌鬃鹅的胸肌生长模式存在显著差异,且40、90日龄是胸肌肌纤维形态变化的关键时期,黏着斑通路、ECM-受体相互作用和细胞黏附分子等通路可通过调控COL4A2、ITGB3和CAV1等8个候选基因表达,介导两品种胸肌肌纤维的肥大过程,进而决定胸肌生长差异。本研究为明确狮头鹅和乌鬃鹅肌肉生长分子调控机制奠定了基础,并为肉鹅分子育种提供了靶基因,将有望加快肉鹅的遗传改良和新品种培育进程。
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2026年第57卷第1期
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doi: 10.11843/j.issn.0366-6964.2026.01.018
  • 接收时间:2025-06-17
  • 首发时间:2026-09-16
  • 出版时间:2026-01-23
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  • 收稿日期:2025-06-17
基金
广东省省级乡村振兴战略专项资金种业振兴项目(2023-XDY-00-001)
广东省省级乡村振兴战略专项资金种业振兴项目(2022-XPY-00-011)
大学生创新创业训练计划项目(202511347257)
大学生创新创业训练计划项目(202511347234)
2024年度广东省重点建设学科科研能力提升项目(2024ZDJS004)
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    仲恺农业工程学院动物科技学院,广州 510225

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伍仲平,主要从事动物遗传育种研究,E-mail:
张续勐,主要从事动物遗传育种研究,E-mail:
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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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