柯乐猪CRISP3基因SNP鉴定及其对繁殖性状的影响

doi:10.3969/j.issn.1004-1524.20230951

浙江农业学报 ›› 2024, Vol. 36 ›› Issue (6): 1270-1278.DOI: 10.3969/j.issn.1004-1524.20230951

柯乐猪CRISP3基因SNP鉴定及其对繁殖性状的影响

向进(), 王春源, 吴燕, 谭元成, 杨酸, 张依裕^*()

贵州大学动物科学学院,高原山地动物遗传育种与繁殖教育部重点实验室,贵州省动物遗传育种与繁殖重点实验室,香猪研究所,贵州贵阳 550025

收稿日期:2023-08-07 出版日期:2024-06-25 发布日期:2024-07-02
作者简介:向进(1995—),男,贵州德江人,硕士研究生,研究方向为动物遗传育种与繁殖。E-mail:1911005146@qq.com
通讯作者: *张依裕,E-mail:zyy8yyc@163.com
基金资助:
贵州省科技支撑计划(黔科合支撑〔2021〕一般147号);贵州省科技人才项目(黔科合平台人才〔2021〕5630号);贵州省生猪产业发展项目(黔财农〔2021〕157号);贵州省生猪产业发展项目(黔财农〔2022〕183号)

CRISP3 gene SNP identification and its impact on reproductive traits in Kela pigs

XIANG Jin(), WANG Chunyuan, WU Yan, TAN Yuancheng, YANG Suan, ZHANG Yiyu^*()

Key Laboratory of Genetic Breeding and Reproduction of Plateau and Mountain Animals, Ministry of Education, Guizhou Provincial Key Laboratory of Animal Genetic Breeding and Reproduction, Institute of Swine, College of Animal Science, Guizhou University, Guiyang 550025, China

Received:2023-08-07 Online:2024-06-25 Published:2024-07-02

摘要/Abstract

摘要：

为分析富半胱氨酸的分泌蛋白3(cysteine rich secretory protein 3,CRISP3)基因单核苷酸多态性(single nucleotide polymorphism, SNP)位点对柯乐猪繁殖性状的影响,采用Sanger测序法检测250头健康经产柯乐猪母猪CRISP3基因的SNP突变位点,利用SPSS 22.0软件分析SNP位点与柯乐猪产活仔猪数、初生窝重、初生均重、断奶窝重、断奶仔猪数和断奶均重6个指标的关联性。结果显示,在柯乐猪CRISP3基因第4内含子共发现3个SNP突变位点,即g.43796318T>C、g.43796424T>A和g.43796425G>T,均存在3种基因型。g.43796318T>C和g.43796424T>A位点的优势基因型为TT,优势等位基因为T;g.43796425G>T位点的优势基因型为GG,优势等位基因为G。χ²检验显示,g.43796318T>C位点基因型分布符合Hardy-Weinberg平衡状态(P>0.05);g.43796424T>A和g.43796425G>T位点基因型分布均极显著(P<0.01)偏离Hardy-Weinberg平衡状态。3个SNP位点的多态信息含量(PIC)为0.25~0.50,表现为中度多态。连锁不平衡分析显示,g.43796424T>A与g.43796425G>T之间存在强连锁不平衡。关联分析显示,g.43796318T>C位点TT和CC基因型个体的初生窝重和初生均重均显著(P<0.05)高于TC基因型;g.43796424T>A位点TA基因型个体的产活仔猪数、初生窝重和断奶仔猪数均显著(P<0.05)高于TT和AA基因型;g.43796425G>T位点GT基因型个体的产活仔猪数、初生窝重和断奶仔猪数均显著(P<0.05)高于GG和TT基因型。双倍型关联分析显示,双倍型H2H4(CCTAGT)个体的产活仔猪数、初生窝重、断奶仔猪数和断奶窝重显著(P<0.05)高于H1H2(TCTAGT)个体和H2H2(CCAATT)个体;双倍型H2H2(CCAATT)个体的初生均重显著(P<0.05)高于除H4H4(CCTTGG)个体外的其他个体,且其断奶均重显著(P<0.05)高于其他双倍型个体。CRISP3基因第4内含子上的3个突变位点与柯乐猪的6个繁殖性状均存在显著关联,可作为柯乐猪繁殖性状选择的候选分子标记。

关键词: 柯乐猪, CRISP3基因, 单核苷酸多态性, 繁殖性状

Abstract:

To analyze the impact of single nucleotide polymorphism (SNP) loci in the Cysteine Rich Secretory Protein 3 (CRISP3) gene on reproductive traits of Kele pigs, Sanger sequencing was used to detect SNP mutations in the CRISP3 gene of 250 healthy and prolific Kele sows. SPSS 22.0 software was utilized to analyze the association between SNP loci and six reproductive traits in Kele pigs, including total number of piglets born, primary litter weight, average birth weight, weaning litter weight, number of weaned piglets, and average weaning weight. The result showed that in the CRISP3 gene of Kele pigs, three SNP mutations were identified in the fourth intron (g.43796318T>C, g.43796424T>A, and g.43796425G>T), each having three genotypes. The dominant genotypes for g.43796318T>C and g.43796424T>A were TT, with the dominant alleles being T, while the dominant genotype for g.43796425G>T was GG, with the dominant allele being G. The χ² test revealed that the genotype distribution of g.43796318T>C was in Hardy-Weinberg equilibrium (P>0.05), whereas the genotype distributions of g.43796424T>A and g.43796425G>T significantly (P<0.01) deviated from Hardy-Weinberg equilibrium. All three SNP loci exhibited moderate polymorphism with polymorphism information content (PIC) ranging from 0.25 to 0.50. Linkage disequilibrium analysis showed strong linkage disequilibrium between g.43796424T>A and g.43796425G>T. The association analysis demonstrated that the TT and CC genotypes of g.43796318T>C were significantly (P<0.05) higher in primary litter weight and average birth weight than those of the TC genotype. The TA genotype of g.43796424T>A had significantly (P<0.05) higher number of piglets born alive, primary litter weight, and number of weaned piglets compared to the TT and AA genotypes. The GT genotype of g.43796425G>T showed significantly (P<0.05) higher number of piglets born alive, primary litter weight, and number of weaned piglets than those of the GG and TT genotypes. In the diplotype analysis, individuals with the H2H4 (CCTAGT) diplotype had significantly (P<0.05) higher number of piglets born alive, primary litter weight, number of weaned piglets, and weaning litter weight compared to individuals with the H1H2 (TCTAGT) diplotype and H2H2 (CCAATT) diplotype. The H2H2 (CCAATT) individuals had significantly (P<0.05) higher average birth weight than individuals of other diplotypes except for H4H4 (CCTTGG), and its average weaning weight was significantly (P<0.05) higher than that of other diplotypes. The three mutation sites in intron 4 of CRISP3 gene were significantly associated with 6 reproductive traits of Kole pigs, which could be used as candidate molecular markers for selection of reproductive traits in Kole pigs.

Key words: Kela pig, CRISP3 gene, single nucleotide polymorphism, reproductive trait

中图分类号:

S828

向进, 王春源, 吴燕, 谭元成, 杨酸, 张依裕. 柯乐猪CRISP3基因SNP鉴定及其对繁殖性状的影响[J]. 浙江农业学报, 2024, 36(6): 1270-1278.

XIANG Jin, WANG Chunyuan, WU Yan, TAN Yuancheng, YANG Suan, ZHANG Yiyu. CRISP3 gene SNP identification and its impact on reproductive traits in Kela pigs[J]. Acta Agriculturae Zhejiangensis, 2024, 36(6): 1270-1278.

图/表 7

参考文献 28

[1]	陈琼, 王济民. 我国肉类消费现状与未来发展趋势[J]. 中国食物与营养, 2013, 19(6): 43-47.
	CHEN Q, WANG J M. Current situation and future trends of meat consumption in China[J]. Food and Nutrition in China, 2013, 19(6): 43-47. (in Chinese with English abstract)
[2]	贾梦雨, 李尚, 刘刁, 等. 母猪繁殖性状分子遗传标记研究进展[J]. 黑龙江畜牧兽医, 2022(1): 38-43.
	JIA M Y, LI S, LIU D, et al. Advances in molecular genetic markers for reproductive traits in sows[J]. Heilongjiang Animal Science and Veterinary Medicine, 2022(1): 38-43. (in Chinese with English abstract)
[3]	颜华, 王立贤, 王立刚, 等. 六个候选基因多态性与母猪繁殖性状关联分析[J]. 农业生物技术学报, 2009, 17(2): 249-255.
	YAN H, WANG L X, WANG L G, et al. Genetic polymorphism of six genetic loci and their correlation with reproductive traits[J]. Journal of Agricultural Biotechnology, 2009, 17(2): 249-255. (in Chinese with English abstract)
[4]	卢世豪, 崔浩, 闫耀晖, 等. VRTN基因型对猪生长和繁殖性能的影响[J]. 石河子大学学报(自然科学版), 2022, 40(4): 460-465.
	LU S H, CUI H, YAN Y H, et al. Association of VRTN gene polymorphisms with growth and reproductive trait in pig[J]. Journal of Shihezi University(Natural Science), 2022, 40(4): 460-465. (in Chinese with English abstract)
[5]	李若岚, 代宇星, 温作晨, 等. FSHR、LHβ基因多态性和法系长白猪繁殖性状的联合关联分析[J]. 中国畜牧杂志, 2021, 57(S1): 92-97.
	LI R L, DAI Y X, WEN Z C, et al. Joint association analysis of FSHR, LHβ gene polymorphism and reproductive traits of Landrace in legal system[J]. Chinese Journal of Animal Science, 2021, 57(S1): 92-97. (in Chinese)
[6]	周礼渝. 多群体全基因组关联性分析鉴别母猪繁殖性状的遗传位点[D]. 南昌: 江西农业大学, 2016.
	ZHOU L Y. Genome-wide association studies for female reproductive traits on three populations[D]. Nanchang: Jiangxi Agricultural University, 2016. (in Chinese with English abstract)
[7]	张宇, 周佳伟, 吴俊静, 等. 大白猪繁殖性状全基因组关联分析[J]. 中国畜牧杂志, 2022, 58(8): 94-99.
	ZHANG Y, ZHOU J W, WU J J, et al. Reproductive traits of Large White pig genome-wide association study[J]. Chinese Journal of Animal Science, 2022, 58(8): 94-99. (in Chinese)
[8]	DOTY A, BUHI W C, BENSON S, et al. Equine CRISP3 modulates interaction between spermatozoa and polymorphonuclear Neutrophils1[J]. Biology of Reproduction, 2011, 85(1): 157-164.
[9]	GONZALEZ S N, SULZYK V, WEIGEL MUÑOZ M, et al. Cysteine-rich secretory proteins (CRISP) are key players in mammalian fertilization and fertility[J]. Frontiers in Cell and Developmental Biology, 2021, 9: 800351.
[10]	KRÄTZSCHMAR J, HAENDLER B, EBERSPAECHER U, et al. The human cysteine-rich secretory protein (CRISP) family[J]. European Journal of Biochemistry, 1996, 236(3): 827-836.
[11]	UDBY L, COWLAND J B, JOHNSEN A H, et al. An ELISA for SGP28/CRISP-3, a cysteine-rich secretory protein in human neutrophils, plasma, and exocrine secretions[J]. Journal of Immunological Methods, 2002, 263(1/2): 43-55.
[12]	GIBBS G M, ROELANTS K, O’BRYAN M K. The CAP superfamily: cysteine-rich secretory proteins, antigen 5, and pathogenesis-related 1 proteins: roles in reproduction, cancer, and immune defense[J]. Endocrine Reviews, 2008, 29(7): 865-897.
[13]	BJARTELL A, JOHANSSON R, BJÖRK T, et al. Immunohistochemical detection of cysteine-rich secretory protein 3 in tissue and in serum from men with cancer or benign enlargement of the prostate gland[J]. The Prostate, 2006, 66(6): 591-603.
[14]	LIAO Q, KLEEFF J, XIAO Y, et al. Preferential expression of cystein-rich secretory protein-3 (CRISP-3) in chronic pancreatitis[J]. Histology and Histopathology, 2003, 18(2): 425-433.
[15]	EVANS J, D’SYLVA R, VOLPERT M, et al. Endometrial CRISP3 is regulated throughout the mouse estrous and human menstrual cycle and facilitates adhesion and proliferation of endometrial epithelial cells1[J]. Biology of Reproduction, 2015, 92(4): 99.
[16]	崔世泉, 李剑虹, 崔卫国, 等. 母猪哺乳初期的母性行为与催乳素受体基因多态性关系的初探[J]. 遗传, 2007, 29(1): 47-51.
	CUI S Q, LI J H, CUI W G, et al. Preliminary study on the relationship between sow maternal behaviour during early lactation and polymorphism of PRLR gene[J]. Hereditas, 2007, 29(1): 47-51. (in Chinese with English abstract)
[17]	杨酸, 郭小江, 杨红文, 等. 柯乐猪PRLR基因多态性与繁殖性状的关联性[J]. 浙江农业学报, 2023, 35(3): 556-564.
	YANG S, GUO X J, YANG H W, et al. Correlation of PRLR gene polymorphisms and reproductive traits in Kele pig[J]. Acta Agriculturae Zhejiangensis, 2023, 35(3): 556-564. (in Chinese with English abstract)
[18]	赵春萍, 张雄, 张静, 等. 不同屠宰体重对柯乐猪胴体性状、肉品质及风味的影响[J]. 黑龙江畜牧兽医, 2022(6): 36-41.
	ZHAO C P, ZHANG X, ZHANG J, et al. Effect of different slaughter weight on meat quality and flavor of Kele pigs[J]. Heilongjiang Animal Science and Veterinary Medicine, 2022(6): 36-41. (in Chinese with English abstract)
[19]	REDDY T, GIBBS G M, MERRINER D J, et al. Cysteine-rich secretory proteins are not exclusively expressed in the male reproductive tract[J]. Developmental Dynamics, 2008, 237(11): 3313-3323.
[20]	UPADHAYA S D, YUN H M, HUANG S Q, et al. Efficacy of dietary supplementation of fatty acid compound on performance and production in finishing pigs[J]. Tropical Animal Health and Production, 2017, 49(6): 1281-1288.
[21]	DA ROS V, BUSSO D, COHEN D J, et al. Molecular mechanisms involved in gamete interaction: evidence for the participation of cysteine-rich secretory proteins (CRISP) in sperm-egg fusion[J]. Society of Reproduction and Fertility Supplement, 2007, 65: 353-356.
[22]	ACHACHE H, REVEL A. Endometrial receptivity markers, the journey to successful embryo implantation[J]. Human Reproduction Update, 2006, 12(6): 731-746.
[23]	DUNCAN W C, SHAW J L V, BURGESS S, et al. Ectopic pregnancy as a model to identify endometrial genes and signaling pathways important in decidualization and regulated by local trophoblast[J]. PLoS One, 2011, 6(8): e23595.
[24]	SERROTE C M L, REINIGER L R S, SILVA K B, et al. Determining the polymorphism information content of a molecular marker[J]. Gene, 2020, 726: 144175.
[25]	WIGGINTON J E, CUTLER D J, ABECASIS G R. A note on exact tests of Hardy-Weinberg equilibrium[J]. The American Journal of Human Genetics, 2005, 76(5): 887-893.
[26]	SHAUL O. How introns enhance gene expression[J]. The International Journal of Biochemistry & Cell Biology, 2017, 91: 145-155.
[27]	NOTT A, MEISLIN S H, MOORE M J. A quantitative analysis of intron effects on mammalian gene expression[J]. RNA, 2003, 9(5): 607-617.
[28]	LE HIR H, NOTT A, MOORE M J. How introns influence and enhance eukaryotic gene expression[J]. Trends In Biochemical Sciences, 2003, 28(4): 215-220.

编辑推荐

Metrics

阅读次数

全文

433

HTML			PDF

最新录用	在线预览	正式出版	最新录用	在线预览	正式出版
0	0	8	0	0	425

来源	本网站	其他网站

次数	331	102
比例	76%	24%

摘要

144

最新录用	在线预览	正式出版

0	0	144

	来源	本网站

	次数	144
	比例	100%

引物名称 Primer name	引物序列(5'→3') Primer sequence (5'→3')	片段长度 Product length/bp	位置 Loci	退火温度 Annealing temperature/℃
CRISP3-F1 CRISP3-R1	TGATGGAACACGATGGAGGAT ATGGCAAACTCCGAAAAGGG	732	外显子3 Exon-3	60
CRISP3-F2	TTAAGGATCTGGTCTTGCCG	664	外显子5 Exon-5	60
CRISP3-R2	AGGCTAGTGCTCAAACATGA
CRISP3-F3	CCCATTGACATTTTGTGCCT	639	外显子6 Exon-6	60
CRISP3-R3	ATTGCTGTGAGCTGTAGTGT
CRISP3-F4	TCAGAGAACTTTTGGGTCCCT	524	外显子8 Exon-8	60
CRISP3-R4	CGGGTATCAAAAGCCAAGGT

引物名称 Primer name	引物序列(5'→3') Primer sequence (5'→3')	片段长度 Product length/bp	位置 Loci	退火温度 Annealing temperature/℃
CRISP3-F1 CRISP3-R1	TGATGGAACACGATGGAGGAT ATGGCAAACTCCGAAAAGGG	732	外显子3 Exon-3	60
CRISP3-F2	TTAAGGATCTGGTCTTGCCG	664	外显子5 Exon-5	60
CRISP3-R2	AGGCTAGTGCTCAAACATGA
CRISP3-F3	CCCATTGACATTTTGTGCCT	639	外显子6 Exon-6	60
CRISP3-R3	ATTGCTGTGAGCTGTAGTGT
CRISP3-F4	TCAGAGAACTTTTGGGTCCCT	524	外显子8 Exon-8	60
CRISP3-R4	CGGGTATCAAAAGCCAAGGT

品种 Breed	SNP位点 SNP loci	基因型频率 Genotype frequency			等位基因频率 Allele frequency		He	Ne	PIC	χ²-HWE
柯乐猪 KeLe pig	g.43796318T>C	TT (143) 0.572	TC (76) 0.304	CC (31) 0.124	T 0.724	C 0.276	0.400	1.666	0.320	14.320
	g.43796424T>A	TT(127) 0.508	TA (104) 0.416	AA (19) 0.076	T 0.716	A 0.284	0.407	1.685	0.324	0.131
	g.43796425G>T	GG(127) 0.508	GT (104) 0.416	TT (19) 0.076	G 0.716	C 0.284	0.407	1.685	0.324	0.131
大白猪 Large White pig	g.43796318T>C	TT (7) 0.35	TC (9) 0.45	CC(4) 0.2	T 0.575	C 0.425	0.489	1.956	0.369	0.126
	g.43796424T>A	TT(6) 0.3	TA (9) 0.45	AA (5) 0.25	T 0.525	A 0.475	0.499	1.995	0.374	0.191
	g.43796425G>T	GG(5) 0.25	GT (11) 0.55	TT (4) 0.2	G 0.525	C 0.475	0.499	1.995	0.374	0.211

品种 Breed	SNP位点 SNP loci	基因型频率 Genotype frequency			等位基因频率 Allele frequency		He	Ne	PIC	χ²-HWE
柯乐猪 KeLe pig	g.43796318T>C	TT (143) 0.572	TC (76) 0.304	CC (31) 0.124	T 0.724	C 0.276	0.400	1.666	0.320	14.320
	g.43796424T>A	TT(127) 0.508	TA (104) 0.416	AA (19) 0.076	T 0.716	A 0.284	0.407	1.685	0.324	0.131
	g.43796425G>T	GG(127) 0.508	GT (104) 0.416	TT (19) 0.076	G 0.716	C 0.284	0.407	1.685	0.324	0.131
大白猪 Large White pig	g.43796318T>C	TT (7) 0.35	TC (9) 0.45	CC(4) 0.2	T 0.575	C 0.425	0.489	1.956	0.369	0.126
	g.43796424T>A	TT(6) 0.3	TA (9) 0.45	AA (5) 0.25	T 0.525	A 0.475	0.499	1.995	0.374	0.191
	g.43796425G>T	GG(5) 0.25	GT (11) 0.55	TT (4) 0.2	G 0.525	C 0.475	0.499	1.995	0.374	0.211

SNP位点SNP loci	g.43796318T>C	g.43796424T>A	g.43796425G>T
g.43796318T>C		0.370	0.370
g.43796424T>A	0.132		1.000
g.43796425G>T	0.132	1.000

柯乐猪CRISP3基因SNP鉴定及其对繁殖性状的影响

CRISP3 gene SNP identification and its impact on reproductive traits in Kela pigs

RichHTML

PDF (PC)

可视化

摘要/Abstract

引用本文

使用本文

图/表 7

参考文献 28

相关文章 14

编辑推荐

Metrics

本文评价

类型Type	基因型Genotype	g.43796318T>C	g.43796424T>A	g.43796425G>T	频率Frequency
单倍型	H1(302)	T	T	G	0.604
Haplotype	H2(80)	C	A	T	0.160
	H3(62)	T	A	T	0.124
	H4(56)	C	T	G	0.112
双倍型	H1H1(94)	TT	TT	GG	0.376
Diplotype	H1H2(40)	TC	TA	GT	0.160
	H1H3(46)	TT	TA	GT	0.184
	H1H4(28)	TC	TT	GG	0.112
	H2H2(7)	CC	AA	TT	0.028
	H2H3(8)	TC	AA	TT	0.032
	H2H4(18)	CC	TA	GT	0.072
	H3H3(4)	TT	AA	TT	0.016
	H4H4(5)	CC	TT	GG	0.020

SNP位置 SNP loci	基因型 Genotype	产活仔猪数 Number of piglets born alive	初生窝重 Primary litter weight/kg	初生均重 Average birth weight/kg	断奶仔猪数 Number of weaned piglets	断奶窝重 Weaning litter weight/kg	断奶均重 Average weaning weight/kg
g.43796318T>C	TT(143)	8.923±2.885	8.671±3.312 a	1.086±0.186 a	6.895±2.729	38.703±16.667 ab	5.682±1.209
	TC(76)	8.579±2.041	7.418±2.342 b	0.975±0.226 b	6.553±2.187	35.659±11.298 b	5.501±0.780
	CC(31)	9.258±3.596	9.213±3.312 a	1.131±0.243 a	7.452±3.107	42.108±17.808 a	5.769±1.986
g.43796424T>A	TT(127)	8.583±2.464 b	8.078±2.77 b	1.066±0.195	6.575±2.574 b	37.501±15.548	5.770±0.862 a
	TA(104)	9.539±2.982 a	9.030±3.441 a	1.048±0.227	7.519±2.603 a	40.034±15.125	5.352±1.265 b
	AA(19)	7.000±2.236 c	6.537±2.301 c	1.060±0.256	5.158±2.115 c	32.834±15.904	6.316±2.305 a
g.43796425G>T	GG(127)	8.583±2.464 b	8.078±2.77 b	1.066±0.195	6.575±2.574 b	37.501±15.548	5.770±0.862 a
	GT(104)	9.539±2.982 a	9.030±3.441 a	1.048±0.227	7.519±2.603 a	40.034±15.125	5.352±1.265 b
	TT(19)	7.000±2.236 c	6.537±2.301 c	1.060±0.256	5.158±2.115 c	32.834±15.904	6.316±2.305 a

双倍型 Diplotype	产活仔猪数 Number of piglets born alive	初生窝重 Primary litter weight/kg	初生均重 Average birth weight/kg	断奶仔猪数 Number of weaned piglets	断奶窝重 Weaning litter weight/kg	断奶均重 Average weaning weight/kg
H1H1(94)	8.596±2.665 bc	8.035±2.943 ab	1.077±0.188 bc	6.457±2.781 cd	37.687±17.013 ab	5.918±0.920 b
H1H2(40)	8.450±2.342 bc	7.025±2.497 b	0.970±0.237 c	6.250±2.457 d	33.935±12.475 b	5.514±0.988 b
H1H3(46)	9.848±3.077 ab	10.235±3.500 a	1.114±0.181 b	7.978±2.314 b	41.944±15.629 ab	5.229±1.588 b
H1H4(28)	8.750±1.691 b	8.154±2.214 ab	1.009±0.209 bc	6.964±1.875 c	37.711±10.316 ab	5.426±0.456 b
H2H2(7)	6.143±2.268 c	6.964±3.415 b	1.303±0.119 a	4.143±2.268 d	34.114±24.283 b	7.474±3.608 a
H2H3(8)	8.625±1.685 bc	6.806±1.296 b	0.881±0.224 c	6.625±1.685 cd	37.094±7.339 ab	5.701±0.458 b
H2H4(18)	11.167±3.222 a	10.408±3.101 a	1.053±0.267 bc	9.167±2.407 a	48.708±14.331 a	5.307±0.827 b
H3H3(4)	5.250±0.500 d	5.250±1.300 c	0.990±0.140 c	4.000±0.011 d	22.075±2.450 c	5.518±0.615 b
H4H4(5)	7.400±2.191 bc	8.460±2.630 ab	1.158±0.191 ab	6.6±1.949 cd	32.810±11.763 b	4.914±0.255 b

[1]	袁晔, 刘睿, 王凌云, 沈盟, 叶雪莲, 权新华, 王瑞森, 姚祥坦. 江浙地区菱品种遗传多样性的SLAF-seq分析[J]. 浙江农业学报, 2023, 35(8): 1773-1781.
[2]	杨酸, 郭小江, 杨红文, 熊力, 李晨, 谭元成, 王春源, 张依裕. 柯乐猪PRLR基因多态性与繁殖性状的关联性[J]. 浙江农业学报, 2023, 35(3): 556-564.
[3]	刘鹏程, 张继, 邱淦远, 龚俞, 李雪松, 李维, 张依裕, 刘若余. 关岭牛TBC1D7基因单核苷酸多态性筛查及生物信息学分析[J]. 浙江农业学报, 2022, 34(7): 1402-1411.
[4]	陶鹏, 岳智臣, 赵彦婷, 雷娟利, 李必元. 大白菜BrSPS1Fb基因剪接受体位点变异及其对剪接的影响[J]. 浙江农业学报, 2021, 33(11): 2068-2074.
[5]	喻宗岗, 蒋隽, 姚亚铃, 郭英, 李闯, 燕海峰. 雪峰乌骨鸡HSP70基因多态性与精液品质关联分析[J]. 浙江农业学报, 2020, 32(8): 1378-1384.
[6]	卢鑫, 周靖航, 杨朝云, 张梦华, 叶连萌, 李叔臻, 黄锡霞, 马云, 王兴平, 史远刚. 新疆褐牛产奶和繁殖性状候选基因功能注释[J]. 浙江农业学报, 2019, 31(12): 1987-1995.
[7]	任卫合, 任稳稳, 马博妍, 高成宏, 李宏旭, 刘丽霞, 曹忻, 张丽. 大耳白兔MSTN基因SNPs位点筛查及生物信息学分析[J]. 浙江农业学报, 2018, 30(12): 2018-2023.
[8]	陈黎，李国勤，田勇，沈军达，陶争荣，徐坚，曾涛，卢立志*. 北京鸭腹部脂肪组织的转录组特征分析[J]. 浙江农业学报, 2016, 28(5): 743-.
[9]	王健1，董飚1，2，侯庆永3，殷洁鑫1. 太湖鹅MyoG基因多态性与体重的相关性分析[J]. 浙江农业学报, 2015, 27(1): 28-.
[10]	戴丽荷;胡锦平;褚晓红;徐如海;路伏增;翁经强;黄少珍. FSHβ基因多态性及对不同品种猪繁殖性状的影响[J]. , 2013, 25(3): 0-466.
[11]	曾涛;李国勤;王德前;李进军;田勇;陈黎;黄爱霞;石放雄;;沈军达;陶争荣;卢立志;. 热休克蛋白70基因（HSP70）部分片段的单核苷酸多态性及其与蛋鸭产蛋性能的关联性[J]. , 2013, 25(2): 0-234.
[12]	李文杨;刘远;林仕欣;张晓佩;高承芳;林碧芬;董晓宁. BMPR-IB，BMP15，GDF9基因作为福建省4个主要山羊品种多胎性能候选基因的研究[J]. , 2012, 24(6): 0-979.
[13]	袁群英;金庆生 . 焦磷酸测序技术及其在水稻单核苷酸多态性研究中的应用[J]. , 2007, 19(4): 0-328.
[14]	王永强;徐孟奎;孟智启;曹锦如;姚陆松;叶爱红;何秀玲. 家蚕无性繁殖性状的基因效应分析[J]. , 2002, 14(6): 0-310.