龙岩山麻鸭蛋品质性状的全基因组关联研究

doi:10.3864/j.issn.0578-1752.2023.03.014

中国农业科学 ›› 2023, Vol. 56 ›› Issue (3): 572-586.doi: 10.3864/j.issn.0578-1752.2023.03.014

• 畜牧·兽医 • 上一篇

龙岩山麻鸭蛋品质性状的全基因组关联研究

孙艳发¹^,²(), 吴琼¹^,², 林如龙³, 陈红萍³, 甘秋云¹, 沈玥¹, 王亚茹¹, 薛鹏飞¹, 陈飞帆¹, 刘健涛¹, 周陈鑫¹, 兰诗诗¹, 潘浩哲¹, 邓凡¹^,⁵, 岳稳³, 江宵兵⁴(), 李焰¹^,²()

¹龙岩学院生命科学学院，福建龙岩 364012
²龙岩学院/福建省家畜传染病防治与生物技术重点实验室/预防兽医学与生物技术福建省高校重点实验室，福建龙岩 364012
³龙岩市新罗区农业局山麻鸭原种场，福建龙岩 364031
⁴福建省畜牧总站，福州 350003
⁵福建农林大学动物科学学院(蜂学学院)，福州 350002

收稿日期:2021-10-20 接受日期:2022-11-16 出版日期:2023-02-01 发布日期:2023-02-14
通信作者: 江宵兵，Tel：13960752743；E-mail：fzjxb@163.com。李焰，Tel：13860217279；E-mail：529783204@qq.com
联系方式: 孙艳发，Tel：18250071633；E-mail：boysun2010@163.com。
基金资助:
福建省种业创新与产业化工程(2021-2025);农业良种重大科研育种攻关与产业化工程项目(zycxny20211014);福建省科技厅对外合作项目(2021I0045);福建省科技厅引导性科技项目(2020N0034);龙岩学院科研博士启动基金(LB2019001)

Genome-Wide Association Study of Egg Quality Traits in Longyan Shan-Ma Duck

SUN YanFa¹^,²(), WU Qiong¹^,², LIN RuLong³, CHEN HongPing³, GAN QiuYun¹, SHEN Yue¹, WANG YaRu¹, XUE PengFei¹, CHEN FeiFan¹, LIU JianTao¹, ZHOU ChenXin¹, LAN ShiShi¹, PAN HaoZhe¹, DENG Fan¹^,⁵, YUE Wen³, JIANG XiaoBing⁴(), LI Yan¹^,²()

¹College of Life Sciences, Longyan University, Longyan 364012, Fujian
²Fujian Provincial Key Laboratory for the Prevention and Control of Animal Infectious Diseases and Biotechnology/Fujian Provincial Universities Key Laboratory of Preventive Veterinary Medicine and Biotechnology/Longyan University, Longyan 364012, Fujian
³Longyan Shan-Ma Duck Original Breeding Farm, Agricultural Bureau of Xinluo District, Longyan 364031, Fujian
⁴Fujian Provincial Animal Husbandry Headquarters, Fuzhou 350003
⁵College of Animal Science (College of Bee Science), Fujian Agriculture and Forestry University, Fuzhou 350002

Received:2021-10-20 Accepted:2022-11-16 Published:2023-02-01 Online:2023-02-14

摘要/Abstract

摘要：

【目的】通过全基因组关联研究（genome-wide association study，GWAS）技术筛选和鉴定鸭蛋品质性状的单核苷酸多态性（single nucleotide polymorphisms，SNPs）位点及候选基因，为龙岩山麻鸭蛋品质性状分子育种提供参考。【方法】试验测定产蛋后期235只龙岩山麻鸭母鸭蛋品质性状，包括蛋重（egg weight，EW）、蛋形指数（egg shaped index，ESI）、蛋壳厚（eggshell thickness，EST）、蛋壳强度（eggshell strength，ESS）、蛋壳颜色L*、a*、b*值（eggshell colour L*, a*, b*，ESCL、ESCA和ESCB）、蛋白高度（albumin height，AH）、哈氏单位（Haugh unit，HU）、蛋黄颜色（egg yolk colour，EYC）、蛋黄重（egg yolk weight，EYW）和蛋黄比例（egg yolk percentage relative to egg weight，EYP）。使用ASReml-R 4.1软件多性状动物模型对蛋品质性状进行遗传参数估计。使用简化基因组测序技术对鸭血液基因组DNA进行SNP分型，分型后进行蛋品质性状与这些SNPs间的GWAS研究。【结果】龙岩山麻鸭蛋品质性状中，EW、ESI、EST、ESL、ESA和AU具有中高等的遗传力，遗传力在0.21—0.70之间。EW与AU存在较强的正遗传相关（r_g= 0.91±0.37）。ESI与EYC存在较强的遗传负相关（r_g= -0.98±1.03）。EST与ESS具有表型正相关（r_p= 0.41±0.06），与ESA具有遗传和表型负相关（r_g= -0.86±0.25和r_p= -0.15±0.07），与ESB具有遗传和表型正相关（r_g= 0.96±0.37和r_p= 0.18±0.07）。ESA与ESB具有遗传和表型负相关（r_g= -0.64±0.28和r_p= -0.31±0.06）。GWAS研究结果表明，7个SNPs位点与ESI、EST和EYC达到5%基因组水平显著关联（P<4.74×10^-6），涉及6个候选基因。与ESI关联的SNP（chr20:11135563:G:C）位点位于20号染色体含有75A富含亮氨酸重复序列（leucine rich repeat containing 75A）基因内。与EST关联的2个SNPs（chr13:5766560:A:G和chrZ:968819:C:T）位点分别位于13号LOC106014427下游6.86 kb处和Z染色体转录因子4（transcription factor 4）基因内。与EYC关联的4个SNPs位点，其中1个（chr2:38155965:G:A）位于2号染色体钾电压门控通道亚家族H成员8（potassium voltage-gated channel subfamily H member 8）基因内；3个SNPs位于9号染色体上的位点，2个（chr9:22623156:G:A和chr9:22623155:T:C）位于胰岛素受体底物1（insulin receptor substrate 1）内、1个（chr9:22490158:A:T）位于LOC106018641内。同时发现81个SNPs位点与蛋品质性状达到基因组水平潜在关联（P<9.48×10^-5）。13个与EYC关联的SNPs位点集中在9号染色体0.84 Mb（22.16—23.00 Mb）区域内。【结论】估计了龙岩山麻鸭蛋品质性状的遗传参数，通过蛋品质性状GWAS研究鉴定了影响ESI、EST和EYC性状的7个显著的SNPs位点、6个候选基因和1个候选基因区域，这些结果为龙岩山麻鸭蛋品质性状分子育种提供参考信息。

关键词: 龙岩山麻鸭, 蛋品质, 遗传参数, 单核苷酸多态性, 全基因组关联研究

Abstract:

【Objective】Single nucleotide polymorphisms (SNPs) and candidate genes for egg quality traits in duck were identified through genome-wide association study (GWAS) technology, so as to provide reference for molecular breeding to improve egg quality traits of Longyan Shan-Ma Duck.【Method】Egg quality traits of 235 female Longyan Shan-Ma Duck were measured, including egg weight (EW), egg shaped index (ESI), eggshell thickness (EST), eggshell strength (ESS), eggshell colour L* (ESCL), a* (ESCA), b* (ESCB), albumin height (AH), Haugh unit (HU), egg yolk colour (EYC), egg yolk weight (EYW), and egg yolk percentage relative to egg weight (EYP). Genetic parameters of these traits were estimated using multi-trait animal model by ASReml-R 4.1 software. Blood genomic DNAs of these ducks were genotyped using genotyping-by-sequencing (GBS) technology. The GWAS between egg quality traits of the late laying period and SNPs were performed. 【Result】 The heritability of EW, ESI, EST, ESL, ESA and AU was higher among the egg quality traits of Longyan Shan-Ma Duck, and ranged from 0.21 to 0.70. There was a strong positive genetic correlation (r_g= 0.91±0.37) between EW and AU, a strong negative genetic correlation (r_g= -0.98±1.03) between ESI and EYC. EST had a positive phenotypic correlation (r_p= 0.41±0.06) with ESS, negative genetic and phenotypic correlations with ESA (r_g= -0.86±0.25 and r_p= -0.15±0.07), and positive genetic and phenotypic correlations with ESB (r_g= 0.96±0.37 and 0.18±0.07). There were negative genetic and phenotypic correlations between ESA and ESB (r_g= -0.64±0.28 and r_p= -0.31±0.06). Results from the GWAS showed that seven SNPs were significantly associated with ESI, EST and yolk color (YC) at 5% Bonferroni-corrected genome-wide significance level (P<4.74×10^-6), involving six candidate genes. One SNP, chr20:11135563: G:C, was associated with ESI, which was in leucine rich repeat containing 75A gene, located on chromosome 20. Two SNPs, chr13:5766560:A:G and chrZ:968819:C:T, were associated with EST, which were located on chromosome 13, downstream 6.86 Kb of LOC106014427 and in transcription factor 4 gene, respectively. Four SNPs were associated with EYC, one SNP chr2: 38155965:G:A in potassium voltage-gated channel subfamily H member 8 gene located on chromosome 2; three SNPs located on chromosome 9, two SNPs, chr9:22623156:G:A and chr9:22623155:T:C, in insulin receptor substrate 1 gene, and one SNP, chr9:22490158:A:T, in LOC106018641 gene. Eighty-one SNPs associated with egg quality traits reached at suggestive genome-wide significance level (P<9.48×10^-5) were also found. Thirteen SNPs associated with YC were distributed in the 0.84 Mb (22.16-23.00 Mb) region of chromosome 9.【Conclusion】In this study, genetic parameters of egg quality traits of Longyan Shan-Ma Duck were estimated. Seven significant SNPs, six candidate genes, and one candidate region affecting ESI, EST and EYC traits were identified through GWAS. The findings from the present study provided a reference for the molecular breeding of egg quality traits in Longyan Shan-Ma Duck.

Key words: Longyan Shan-Ma Duck, egg quality traits, genetic parameter, SNPs, GWAS

孙艳发, 吴琼, 林如龙, 陈红萍, 甘秋云, 沈玥, 王亚茹, 薛鹏飞, 陈飞帆, 刘健涛, 周陈鑫, 兰诗诗, 潘浩哲, 邓凡, 岳稳, 江宵兵, 李焰. 龙岩山麻鸭蛋品质性状的全基因组关联研究[J]. 中国农业科学, 2023, 56(3): 572-586.

SUN YanFa, WU Qiong, LIN RuLong, CHEN HongPing, GAN QiuYun, SHEN Yue, WANG YaRu, XUE PengFei, CHEN FeiFan, LIU JianTao, ZHOU ChenXin, LAN ShiShi, PAN HaoZhe, DENG Fan, YUE Wen, JIANG XiaoBing, LI Yan. Genome-Wide Association Study of Egg Quality Traits in Longyan Shan-Ma Duck[J]. Scientia Agricultura Sinica, 2023, 56(3): 572-586.

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链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2023.03.014

https://www.chinaagrisci.com/CN/Y2023/V56/I3/572

图/表 9

表1

表2

表3

图1

表4

表5

基因组水平潜在关联的SNP位点"

性状 Trait	染色体 Chromosome	标记号 SNP ID	物理位置 Position (bp)	A1 A1	BETA值 BETA	P值 P value	最近的基因 Nearest gene
蛋重EW	2	chr2:10782468:A:G	10782468	G	3.45	2.81×10^-5	U 120.07 kb ADARB2
	2	chr2:10790319:T:C	10790319	T	3.31	5.81×10^-5	U 112.21 kb ADARB2
	5	chr5:5547457:T:C	5547457	T	2.89	3.36×10^-5	U 63.45 kb MIS18BP1
	5	chr5:5547530:A:G	5547530	G	2.80	6.33×10^-5	U 63.38 kb MIS18BP1
蛋形指数ESI	3	chr3:81386471:A:G	81386471	A	-0.05	6.66×10^-5	D 13.01 NT5E
	5	chr5:7009743:G:A	7009743	G	0.06	2.77×10^-5	within CDKL1
	15	chr15:11227397:T:C	11227397	C	0.05	6.39×10^-5	within AXIN1
	15	chr15:11227409:G:A	11227409	A	0.05	7.59×10^-5	within AXIN1
	15	chr15:13294556:C:T	13294556	T	0.05	2.39×10^-5	within UBN1
	15	chr15:13395347:A:G	13395347	A	0.05	3.49×10^-5	within LOC101799012
	15	chr15:14536844:G:A	14536844	A	0.05	1.02×10^-5	U 41.78 kb RBFOX1
	16	chr16:2579629:A:T	2579629	T	-0.04	5.07×10^-5	within EP400
	16	chr16:3098786:G:A	3098786	A	-0.04	1.97×10^-5	D 22.42 kb LOC110352440
	25	chr25:6315960:T:C	6315960	C	-0.06	5.29×10^-5	D 26.17 kb LOC106020386
蛋壳厚度EST	2	chr2:22513511:C:A	22513511	A	0.03	2.51×10^-5	within ZNF804B
	2	chr2:22582514:A:G	22582514	G	0.03	4.43×10^-5	within ZNF804B
	2	chr2:22582670:G:A	22582670	A	0.03	6.51×10^-5	within ZNF804B
	6	chr6:7192663:G:A	7192663	A	0.03	6.46×10^-5	U 18.98 kb KCNK18
	6	chr6:12756206:T:A	12756206	A	0.02	7.06×10^-5	within SH3PXD2A
	12	chr12:140915:T:A	140915	A	-0.02	9.44×10^-5	within THSD4
	12	chr12:198263:G:A	198263	A	-0.02	5.31×10^-5	within THSD4
	13	chr13:5721234:C:T	5721234	T	0.02	7.27×10^-5	within LOC106014427
	19	chr19:4873329:A:G	4873329	G	0.02	2.69×10^-5	within LOC113845541
蛋壳强度ESS	5	chr5:33699890:A:T	33699890	T	0.66	6.76×10^-5	within LOC113843881
蛋壳颜色L*值 ESCL	7	chr7:3182113:T:A	3182113	A	-3.45	6.81×10^-5	within CTR9
	7	chr7:3204268:G:A	3204268	A	-3.31	6.08×10^-5	U 0.65 kb EIF4G2
	7	chr7:3204312:A:G	3204312	G	-3.34	6.05×10^-5	U 0.60 kb EIF4G2
	7	chr7:3223217:A:G	3223217	G	-3.55	1.05×10^-5	U 14.01 kb LOC113844171
	7	chr7:3223320:T:A	3223320	A	-3.37	9.92×10^-6	U 13.91 kb LOC113844171
	14	chr14:9802965:A:G	9802965	G	2.04	5.95×10^-5	within CLINT1
蛋壳颜色a*值 ESCA	1	chr1:160744676:G:C	160744676	G	-1.09	9.13×10^-5	D 3.53 kb PHF5A
	1	chr1:174907611:C:T	174907611	T	-0.89	5.86×10^-5	U 19.87 kb GPR85
	4	chr4:6332903:C:G	6332903	G	-0.93	5.91×10^-5	U 6.34 kb MANBA
	8	chr8:10668874:C:A	10668874	A	-0.97	8.83×10^-5	D 3.43 kb LOC113844346
蛋壳颜色 b*值 ESCB	1	chr1:59043295:C:T	59043295	T	-1.54	7.39×10^-5	U 142.06 kb ARHGEF7
	1	chr1:59706095:T:C	59706095	C	1.03	6.95×10^-5	D 3.76 kb COL4A2
	3	chr3:16497369:T:C	16497369	C	-1.19	2.36×10^-5	U 13.30 kb VSX1
	4	chr4:7424076:C:G	7424076	G	1.83	6.35×10^-5	within RAP1GDS1
	8	chr8:33254427:C:T	33254427	T	1.20	3.93×10^-5	within SLC30A7
蛋白高度AU	2	chr2:102420094:T:G	102420094	G	-0.89	5.83×10^-5	U 14.46 kb RBBP8
	4	chr4:5482597:A:C	5482597	C	1.49	4.59×10^-5	D 21.57 kb LOC106017184
	4	chr4:71726051:A:G	71726051	G	0.90	2.81×10^-5	within LOC106017926
	7	chr7:7414460:C:T	7414460	T	0.80	3.39×10^-5	D 1.45 kb LOC106019248
	20	chr20:6093764:T:C	6093764	T	1.30	4.41×10^-5	U 5.58 kb SGSM2
	20	chr20:10720431:A:G	10720431	G	1.01	3.37×10^-5	U 15.28 kb RTN4RL1
哈氏单位HU	1	chr1:111329343:T:C	111329343	C	12.09	8.38×10^-5	within TAGLN3
	4	chr4:75257160:A:C	75257160	A	-8.15	3.42×10^-5	U 91.88 kb LOC110353528
	7	chr7:7414460:C:T	7414460	T	6.42	5.32×10^-5	D 1.45 kb LOC106019248
	20	chr20:6093764:T:C	6093764	T	10.22	8.41×10^-5	U 5.58 kb SGSM2
	20	chr20:10720431:A:G	10720431	G	8.18	4.19×10^-5	U 15.28 kb RTN4RL1
蛋黄颜色EYC	1	chr1:196541696:G:T	196541696	T	0.75	5.41×10^-5	within CAMK1D
	2	chr2:2433073:A:G	2433073	G	0.94	2.65×10^-5	within GJC2
	2	chr2:37584936:A:G	37584936	G	1.01	1.99×10^-5	D 60.71 kb SATB1
	2	chr2:37585118:C:T	37585118	T	0.77	6.42×10^-5	D 60.53 kb SATB1
	2	chr2:57679783:T:C	57679783	C	0.71	6.82×10^-5	within GLI3
	2	chr2:57679906:A:G	57679906	G	0.70	9.04×10^-5	within GLI3
	2	chr2:115153719:T:C	115153719	C	-0.97	5.50×10^-5	within ZNF521
	2	chr2:115178457:C:T	115178457	T	-1.10	5.57×10^-5	within ZNF521
	3	chr3:51410494:C:T	51410494	T	-0.80	5.15×10^-5	within RMND1
	3	chr3:51410551:A:G	51410551	G	-0.80	5.15×10^-5	within RMND1
	6	chr6:6402767:G:A	6402767	A	-0.85	8.86×10^-5	within CACUL1
	7	chr7:10854039:A:G	10854039	G	-1.28	6.98×10^-5	within MAP2
	9	chr9:22162962:T:G	22162962	G	-1.08	4.62×10^-5	within NYAP2
	9	chr9:22163003:A:G	22163003	G	-1.07	5.51×10^-5	within NYAP2
	9	chr9:22163123:T:C	22163123	C	-1.07	5.51×10^-5	within NYAP2
	9	chr9:22490118:C:T	22490118	C	-0.96	6.18×10^-6	within LOC106018641
	9	chr9:22490249:A:G	22490249	A	-0.83	2.13×10^-5	within LOC106018641
	9	chr9:22628656:T:C	22628656	C	-0.97	5.97×10^-5	within IRS1
	9	chr9:22636905:C:T	22636905	T	-1.04	4.55×10^-5	within IRS1
	9	chr9:22798030:T:C	22798030	C	-0.92	1.94×10^-5	within COL4A3
	9	chr9:22841648:T:C	22841648	C	-0.95	6.18×10^-5	within COL4A3
	9	chr9:23005131:A:G	23005131	G	-1.04	7.05×10^-5	within DAW1
	14	chr14:1139032:G:A	1139032	A	-0.90	2.55×10^-5	within ZIC4
	24	chr24:3527498:A:G	3527498	A	0.74	7.73×10^-5	within EPB41
蛋黄重EYW	2	chr2:2865193:A:G	2865193	A	-2.66	6.56×10^-5	U 2.17 kb SEC22C
	9	chr9:11219039:A:G	11219039	G	2.15	9.42×10^-5	within TP63
	9	chr9:11360616:G:A	11360616	A	2.27	4.28×10^-5	within TP63
	9	chr9:11360684:A:G	11360684	G	2.27	4.43×10^-5	within TP63
蛋黄比例EYP	2	chr2:129285222:C:T	129285222	T	-6.95	6.53×10^-5	within RMDN1
	5	chr5:8855150:G:A	8855150	A	5.41	6.83×10^-5	D 19.17 kb LOC110352963
	6	chr6:708965:T:C	708965	C	6.13	2.71×10^-5	within DPYSL4

表5

图2

图3

图4

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性状 Trait	例数 N	平均值 Mean	最小值 Min	最大值 Max	标准差 SD	变异系数 CV（%）
蛋重EW (g)	235	69.70	51.30	84.30	5.28	7.57
蛋形指数ESI	235	1.33	1.15	1.50	0.06	4.52
蛋壳厚EST (mm)	235	0.33	0.20	0.44	0.04	10.68
蛋壳强度ESS (kg·cm^-2)	233	3.40	0.61	6.00	1.04	30.68
蛋壳颜色L* 值ESCL	235	78.11	65.11	86.29	2.86	3.67
蛋壳颜色a*值ESCA	234	-3.04	-8.44	-0.53	1.52	-50.17
蛋壳颜色b*值ESCB	234	4.81	1.37	13.63	1.66	34.58
蛋白高度 AH (mm)	235	6.36	1.32	20.80	3.33	9.85
哈氏单位HU	235	74.84	38.22	95.47	10.70	14.29
蛋黄颜色EYC	235	7.37	3.00	13.00	1.22	16.52
蛋黄重 EYW (g)	232	24.14	11.70	37.30	3.70	15.33
蛋黄比例 EYP (%)	232	34.77	17.06	57.91	5.52	15.87

	蛋重 EW	蛋形指数 ESI	蛋壳厚 EST	蛋壳强度 ESS	*蛋壳颜色L值 ESCL**	*蛋壳颜色a值 ESCA**	*蛋壳颜色b值 ESCB**	蛋白高度 AH	哈氏单位 HU	蛋黄颜色 EYC	蛋黄重 EYW	蛋黄比例 EYP
蛋重 EW	0.21± 0.16	-	-	-	-	-	-	0.91± 0.37*	-	-	-	-
蛋形指数 ESI	-	0.29± 0.17	-	-	-	-	-	-	-	-0.98± 1.03*	-	-
蛋壳厚 EST	-	-	0.41± 0.17	-	-	-0.86± 0.25**	0.96± 0.37**	-	-	-	-	-
蛋壳强度 ESS	-	-	0.41± 0.06***	0.01± 0.10	-	-	-	-	-	-	-	-
蛋黄颜色L*值 ESCL	-	-	-	-	0.37± 0.18	-	-	-	-	-	-	-
蛋黄颜色a*值 ESCA	-	-0.16± 0.07*	-0.15± 0.07**	-	0.25± 0.07***	0.70± 0.20	-0.64± 0.28*	-	-	-	-	-
蛋黄颜色b*值 ESCB	-	-	0.18± 0.07**	0.18± 0.06*	-0.31± 0.06***	-0.31± 0.06***	0.16± 0.14	-	-	-	-	-
蛋白高度 AH	0.04± 0.07*	-	-	0.20± 0.06**	-	-0.09± 0.07*	-	0.24± 0.18	-	-	-	-
哈氏单位 HU	-0.18± 0.07***	-	-	0.22± 0.06**	-	-0.12± 0.07*	-	-	0.08± 0.14	-		-
蛋黄颜色 EYC	-0.10± 0.07*	0.07± 0.06*	-	-	-	-	-	-	0.18± 0.06**	0.07± 0.14	-	-
蛋黄重 EYW	0.24± 0.06***	-	-	-	-	-	-	-	-	-	0.01± 0.12	-
蛋黄比例 EYP	-0.26± 0.06***	-	-	-	-	-	-	-	-	-	0.87± 0.02***	0.08± 0.12

龙岩山麻鸭蛋品质性状的全基因组关联研究

Genome-Wide Association Study of Egg Quality Traits in Longyan Shan-Ma Duck

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染色体 Chromosome	SNPs数量 No. of SNPs	独立SNPs数量 No. of independent SNPs	染色体 Chromosome	SNPs数量 No. of SNPs	独立SNPs数量 No. of independent SNPs
1	9911	1523	16	1132	172
2	7488	1250	17	30	9
3	6116	988	18	1420	230
4	4026	644	19	989	149
5	4252	706	20	1546	251
6	2311	435	21	1382	242
7	2120	335	22	1072	177
8	2479	402	23	278	63
9	2163	407	24	687	122
10	1242	202	25	974	186
11	2177	369	26	134	48
12	1028	152	27	819	189
13	2066	377	28	766	173
14	1607	307	29	506	80
15	1545	240	Z	440	119
总计Total				62706	10547

性状 Trait	染色体 Chromosome	标记号 SNP ID	物理位置 Position (bp)	A1 A1	BETA值 BETA	P值 P value	最近的基因 Nearest gene
蛋形指数ESI	20	chr20:11135563:G:C	11135563	C	-0.05	1.48×10^-6	within LRRC75A
蛋壳厚EST	13	chr13:5766560:A:G	5766560	A	0.02	1.36×10^-6	6.86 kb D LOC106014427
蛋壳厚EST	Z	chrZ:968819:C:T	968819	T	-0.02	1.96×10^-6	within TCF4
蛋黄颜EYC	2	chr2:38155965:G:A	38155965	A	1.05	1.97×10^-6	within KCNH8
	9	chr9:22490158:A:T	22490158	T	-0.97	3.77×10^-6	within LOC106018641
	9	chr9:22623155:T:C	22623155	C	-0.92	1.91×10^-6	within IRS1
	9	chr9:22623156:G:A	22623156	A	-1.08	8.75×10^-7	within IRS1

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