CN109402264B - 一种提高苏尼特羊繁殖力的分子标记及其应用 - Google Patents
一种提高苏尼特羊繁殖力的分子标记及其应用 Download PDFInfo
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Abstract
本发明公开了利用GDF9基因提高苏尼特羊繁殖力的方法，是检测待测苏尼特羊基因组中的GDF9基因中g.41770718T/G位点的核苷酸，从而确定苏尼特羊的基因型,然后通过基因型确定苏尼特羊繁殖力，如果苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为T时，其纯合体的基因型为TT；苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为G时，其纯合体的基因型为GG，它们的杂合体基因型为TG；实验证明GG基因型苏尼特羊的平均产羔数高于GT基因型，基因型GT苏尼特羊的平均产羔数高于TT型。本发明利用PCR‑RFLP迅速、简便的检测分子标记多态性，为利用分子标记辅助育种技术提高苏尼特羊繁殖力，提供了一个准确简便的方法。
Description
技术领域
一种提高苏尼特羊繁殖力的分子标记及其应用
背景技术
GDF9(growth differentiation factor 9，GDF9)即生长分化因子9，与BMP15基因同属于转化生长因子β(TGFβ)超家族成员。绵羊的GDF9基因位于到5号染色体上，全长约2.5kb，其中包含2个外显子和1个内含子，外显子1长397bp，编码134个氨基酸，外显子2长968bp，编码456个氨基酸，内含子长1126bp。(Sadighi M,Bodensteiner K J,Beattie A Eet al.Genetic mapping of ovine growth differentiation factor 9(GDF9)to sheepchromosome 5.[J].Animal Genetics,2002,33(3):244-245.)
苏尼特羊产于内蒙古自治区锡林郭勒盟苏尼特左旗和苏尼特右旗，有戈壁羊之称。其主要饲养方式为纯天然牧场自然放牧，气候特点为春季干旱多风、夏季炎热、冬季寒冷，主要饲养植物种类有沙葱、多根葱、小型针茅草等多达477种。苏尼特羊有很强的适应能力，在终年放牧不补任何伺料的条件下都能很快抓膘，而且肌肉丰满，体格壮大，肉质细嫩，无膻味，口感好，有较高的产肉性能。苏尼特羊肉富含人体所需要的各种氨基酸、脂肪酸、矿物质、维生素等，且蛋白质含量高、脂肪含量低，因此在国内外非常受欢迎。1986年苏尼特羊被锡林郭勒盟技术监督局批准为地方良种，1997年被内蒙古自治区人民政府正式命名为苏尼特羊，在2010年被列入全国优良畜种名录，2015年被列入国家级畜禽遗传保护名录。(高凤明，白乙尔图，刘金，孙彦军，邵志勇。苏尼特羊及羊肉的品质与营养。[J]中国畜牧兽医文摘，2014，30(12)：43-44.)
李碧侠等用PCR-SSCP技术分析了GDF9基因在小尾寒羊、湖羊、多赛特和萨福克4个绵羊品种的多态性，结果表明在湖羊、多赛特和萨福克羊GDF9基因外显子1编码区152bp处发生了A→G的突变，导致第51位氨基酸由天冬酰胺变为天冬氨酸，小尾寒羊没有发生该突变。(李碧侠，储明星，王金玉。绵羊GDF9基因PCR-SSCP分析。[J]遗传学报，2003，30(4)：307-310.)
Hanrahan等用PCR-SSCP和直接测序的方法在Cambridge绵羊和Belclare绵羊GDF9基因中发现了G1～G8共8个突变，其中有3个核苷酸改变(分别位于编码区471bp处的C→T[G2突变]、477bp处的G→A[G3突变]和978bp处A→G[G5突变])没有导致氨基酸变化；4个G→A的突变导致了氨基酸变化，分别是位于编码区260bp处的突变导致第87位由精氨酸变为组氨酸(G1突变)，编码区721bp处的突变导致第241位由谷氨酸变为赖氨酸(G4突变)，编码区994bp处的突变导致第332位由缬氨酸变为异亮氨酸(G6突变)，编码区1111bp处的突变导致第371位由缬氨酸变为甲硫氨酸(G7突变)；另外编码区1184bp处的C→T碱基突变导致第395位由丝氨酸变为苯丙氨酸(G8突变，FecGH)。(Hanrahan J P,Gregan S M,Mulsant P etal.Mutations in the genes for oocyte derived growth factors GDF9 and BMP15are associated with both increased ovulation rate and sterility in Cambridgeand Belclare sheep(Ovisaries).[J].Biology of Reproduction,2004,70(4):900-909.)
管峰等用PCR-RFLP方法研究GDF9基因G8突变在湖羊、夏洛来、多赛特、萨福克、中国美利奴肉用多胎品系、中国美利奴羊和罗姆尼7个绵羊群体中的分布，结果发现仅在湖羊群体中检测到G8纯合突变，但发生率极低(0.645％，2/310)。(管峰，艾君涛，庞训胜等。绵羊GDF9和BMP15基因多态性检测。[J]生命科学研究，2005，9(2)：184-188.)
杨晶采用PCR-SSCP方法分析GDF9基因在小尾寒羊和多赛特羊中的多态性，结果表明在小尾寒羊和多赛特羊GDF9基因外显子2中都检测到了G3突变和编码区729bp处的G→T突变，后者导致243位氨基酸由谷氨酰胺变为组氨酸。(杨晶。小尾寒羊GDF9基因和BMP15基因多态性及其与高繁殖力关系的研究。[D]北京：中国农业科学院，2006.)
高丽霞用PCR-SSCP方法分析GDF9基因外显子2在小尾寒羊、滩羊、同羊和欧拉羊4个绵羊群体中的多态性，结果发现在这4个绵羊品种GDF9基因外显子2编码区558bp和第692bp处都发生了T→C的突变，其中后者导致231位氨基酸由异亮氨酸变为苏氨酸。(高丽霞。小尾寒羊TGF-β1和GDF9基因多态性及其与繁殖力关系的研究。[D]北京：中国农业科学院，2007.)
Nicol等研究发现，FecTT突变是冰岛Thoka绵羊GDF9基因外显子2编码区1279位发生的A→C的突变，FecTT突变纯合子Thoka母羊不育，突变杂合子母羊产羔数比野生型多0.6只。该突变对Thoka绵羊生殖的影响主要表现在三个方面。第一：解剖发现，FecTT突变纯合子(TT)Thoka母羊都存在严重的生殖道畸形现象。表现为子宫没有发育，卵巢小而且未被激活，没有排卵前卵泡和黄体等。第二：TT型母羊卵巢的皮质区域含有大量的原始卵泡，其中许多都被一至两层颗粒细胞包围，且大多数呈现异常形态，也有极少数卵泡被两到四层不对称的颗粒细胞包围。一些TT型卵泡含有相当于正常窦状卵泡大小的卵母细胞，但在高倍镜下观察发现许多卵泡不是已完全瓦解，就是正在瓦解；另一些异常的卵泡中卵母细胞聚集成簇，并被一层颗粒细胞包围。而野生型母羊(++)的卵巢中含有发育到各个阶段的卵泡，在卵泡皮质区域仅可见极少数原始卵泡。第三：与++型母羊相比，FecTT突变使TT型母羊血浆中的促卵泡激素和黄体生成素水平增加，而抑制素A和雌二醇水平下降。TT型母羊颗粒细胞无明显增殖，而在++型母羊中，当卵母细胞增大到直径达80mm时，就会增殖出现许多颗粒和卵泡膜细胞。(Nicol L,Bishop S C,Pong-Wong R et al.Homozygosity for a singlebase-pair mutation in the oocyte-specific GDF9 gene results in sterility inThoka sheep.[J].Reproduction,2009,138(6):921-933.)
畅静桃等采用PCR-SSCP和测序的方法在小尾寒羊、白萨福克、特克塞尔和藏羊中都检测到GDF9基因G2突变。(畅静桃，罗玉柱，胡江。绵羊高繁殖力候选基因GDF9的多态性分析。[J]甘肃农业大学学报，2009，44(2)：30-33.)
陈勇等用PCR-SSCP方法在小尾寒羊，中国美利奴(新疆型)绵羊多胎品系、肉用品系、体大品系，萨福克，无角多赛特，中国美利奴(新疆型)和德国肉用美利奴8个绵羊群体中都检测到了GDF9基因的G4突变。(陈勇，雒秋江，张亚军等。绵羊BMP15和GDF9基因多态性与产羔性能间的关系。[J]新疆农业大学学报，2009，32(5)：10-17.)
Barzegari等在伊朗的Moghani和Ghezel绵羊群体检测GDF9基因的G1突变。(Barzegari A,Atashpaz S,Ghabili K et al.Polymorphisms in GDF9andBMP15associated with fertility and ovulation rate in Moghani and Ghezel sheepin Iran.[J].Reproduction in Domestic Animals,2009,45(4):666-669.)
Polley等在印度的Garole绵羊群体中均检测到GDF9基因的G1突变。(Polley S,DeS,Brahma B et al.Polymorphism of BMPR1B,BMP15 and GDF9 fecundity genes inprolific Garole sheep.[J].Tropical Animal Health and Production,2009,85(2):122-129.)
FecGE是GDF9基因的一种新型突变，是巴西SantaInes绵羊GDF9基因第2外显子编码区1034位发生了G→T改变，从而引起肽链第345位氨基酸残基改变，由半胱氨酸变为苯丙氨酸(C345F)，突变位点正好位于成熟蛋白质内，且该突变能引起排卵数和产羔数的增加，突变纯合子排卵数增加0.82枚，产羔数增加0.58,而突变杂合子及野生型在排卵数和产羔数方面无明显变化。(Silva B D,Castro E A,Souza C J et al.A new polymorphisminthe growth and differentiation factor 9(GDF9)gene is associated withincreased ovulation rate and prolificacy in homozygous sheep.[J].AnimalGenetics,2011,42(1):89-92.)
Vage等在挪威白绵羊中研究发现，FecGF突变纯合子人工授精公羊的雌性后代产羔数增加0.46～0.57只，杂合子公羊的雌性后代产羔数增加0.20～0.25只。(Vage D I,Husdal M,Kent M P et al.A missense mutation in growth differentiation factor9(GDF9)is strongly associated with litter size in sheep.[J].BMC Genetics,2013,14:1)
2013年，Mullen等在Lleyn绵羊中发现FecGH的突变。次年，Mullen等又在在FinnishLandrace绵羊和Belclare绵羊中研究发现，FecGF突变的纯合子和杂合子排卵数都比野生型高[2]。([1]Mullen M P,Hanrahan J P,Howard D J et al.Investigation ofprolific sheep from UK and Ireland for evidence on origin of the mutations inBMP15(FecXG,FecXB)and GDF9(FecGH)in Belclare and Cambridge sheep.[J].PLoS ONE,2013,8(1):e53172.[2]Mullen M P,Hanrahan J P.Direct evidence on thecontribution of a missense mutation in GDF9to variation in ovulation rate ofFinnsheep.[J].PLoS ONE,2014,9(4):e95251.)
Souza等通过巴西绵羊育种协会的三联交付的数据库，确定了多产的法兰西岛母羊群重新测序后鉴定了突变位点(c.943C>T)并定义这个位点为FecGV；并且证明了FecGV的杂合体有更高的排卵率。(Souza C J,McNeilly A S,Benavides M V,Melo E O,Moraes JC.Mutation in the protease cleavage site of GDF9increases ovulation rate andlitter size in heterozygous ewes and causes infertility in homozygous ewes.[J].Animal genetics,2014,45(5):732-739.)
2015年雷梦媛等在6个绵羊品种(湖羊、小尾寒羊、洼地绵羊、杜泊绵羊、黑萨福克和白萨福克)中均未检测到FecGE突变。(雷梦媛，潘章源，狄冉，刘秋月，胡文萍，王翔宇，储明星和李学伟。山羊和绵羊GDF9基因FecGE突变检测。扬州大学学报。[J]2015，36(1)：1-1.)
R.Khodabakhshzadeh利用PCR-SSCP技术检测Kermani绵羊品种GDF9基因的外显子2(634bp和647bp片段)，此外，使用Gen Alex 6.41软件计算每个SNP的群体遗传指数。测序结果显示在研究在羊群中还存在3个突变(SNP)在443,477和721个位置。(Khodabakhshzadeh R,Mohammadabadi M R,Esmailizadeh A K,Shahrebabak M H,Bordbar F,Namin A S.Identification of point mutations in exon 2of GDF9gene inKermani sheep.[J].Polish Journal of Veterinary Sciences,2016,19(2):281-289.)
有研究表明，Saidi绵羊的GDF9基因的1号外显子有A→T的突变。(El Fiky Z A,Hassan G M,Nassar M I.Genetic polymorphism of growth differentiation factor 9(GDF9)gene related to fecundity in two Egyptian sheep breeds.[J].Journal ofAssisted Reproduction and Genetics.2017,34(12):1683-1690.)
多篇国内外文献证实在小尾寒羊、湖羊、多赛特、特克塞尔、德国肉用美利奴[1]，新疆策勒绵羊、多浪绵羊[2]，滩羊[3]，印度的Garole、Malpura、Decanni绵羊[4]，伊朗的Shal绵羊[5]，北非的Barbarine、Queue Fine de L’Ouest、Noire de Thibar、Sicilo-Sarde、D’Man绵羊[6]中均不携带GDF9基因的G8突变。([1]储明星，桑林华，王金玉等。小尾寒羊高繁殖力候选基因BMP15和GDF9研究。[J]遗传学报，2005，32(1)：38-45。[2]白杰，史洪才，刘明军等。BMP15和GDF9基因作为策勒黑羊和多浪羊的多胎主效候选基因的研究。[J]草食家畜，2007(2)：1-4。[3]孙红霞，田秀娥，王永军。BMPR-IB，BMP15和GDF9基因作为滩羊繁殖性状主效候选基因的研究。[J]西北农业学报，2009，18(5)：17-21。[4]Kumar S,Mishra A K,Kolte A Pet al.Screening for Booroola(FecB)and Galway(FecXG)mutations in Indiansheep.[J].Small Ruminant Research,2008,80(1):57-61.[5]Ghaffari M,Nejati-Javaremi A,Rahimi-Mianji G.Lack of polymorphism in the oocyte derived growth factor(GDF9)gene in the Shal breed of sheep.[J].South African Journal of AnimalScience,2009,39(4):355-360.[6]Vacca G M,Dhaouadia A,Rekikb M etal.Prolificacy genotypes at BMPR1B,BMP15and GDF9 genes in North African sheepbreeds.[J].Small Ruminant Research,2010,88(1):67-71.)
发明内容
本发明的内容是提供了一种提高苏尼特羊双羔率的分子标记及使用方法
本发明所提供的提高苏尼特羊双羔率的分子标记，检测待测苏尼特羊基因组中的GDF9基因中g.41770718T/G处是否存在一个T→G的突变，以确定苏尼特羊个体在该位点的基因型，然后通过基因型筛选种用苏尼特羊；所述的GDF9基因的核苷酸序列NCBIReferenceSequence为NC_019462.2的羊5号染色体，第41770600bp到41771399bp区域。
所述确定苏尼特羊的基因型的方法为：如果苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为T时，其纯合体的基因型为TT；苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为G时，其纯合体的基因型为GG，它们的杂合体基因型为TG；
通过基因型提高苏尼特羊繁殖力为：所述GDF9基因中g.41770718T/G位点GG基因型苏尼特羊的平均产羔数高于GT基因型，基因型GT苏尼特羊的平均产羔数高于TT型。
苏尼特羊繁殖力相关基因GDF9的分子标记的特异性引物，其特征在于，所述特异性引物序列如下：
SEQ ID NO.1：5’-AATGCCAGGGGAAAGGAA-3’；
SEQ ID NO.2：5’-TGGCTTGGAAGAATTAGCAAG-3’
本发明方法，采用限制性内切酶片段长度多态性(restriction fragment lengthpolymorphism，RFLP)，方法对在GDF9基因进行单核苷酸多态性(single nucleotidepolymorphism，SNP)检测，以比较GDF9基因在苏尼特羊品种中的多态性，并对具有RFLP多态性的片段进行测序比较分析，寻找到与繁殖力相关的分子标记，数据统计结果表明，该分子标记可辅助提高苏尼特羊的繁殖力。本发明的方法即利用该分子标记，建立RFLP检测该基因型的技术体系，并利用该基因型选育种羊，辅助提高苏尼特羊繁殖力的有效方法。实验证明，该方法可以迅速、简便的检测苏尼特羊的繁殖力。本发明的方法为利用分子标记辅助育种技术提高苏尼特羊的繁殖力，提供了一个准确简便的检测方法。
附图说明
图1为GDF9基因中g.41770718T/G位点的PCR产物图
图2为GDF9基因中g.41770718T/G位点的酶切图
图3为GDF9基因中g.41770718T/G位点的测序结果
具体实施方式
下述实施例中提到的实验方法，如无特别说明均为常规方法。
下列实施例中所用主要试剂Takara、NEB和华大基因公司购买。
实施例1、检测提高苏尼特羊双羔率的分子标记的方法的建立及其效果验证
1.检测提高苏尼特羊繁殖力的分子标记的方法建立
1)模板材料准备
采集苏尼特羊的血样并记录繁殖力和胎次，所采集的苏尼特羊来自于内蒙古锡林郭勒盟苏尼特左旗，将血样保存在抗凝管中，待用。
2)PCR产物测序。
随机抽取20只苏尼特羊(连续产双羔母羊10只+连续单羔母羊10只)。进行PCR产物测序，找到了位于GDF9启动子的SNP位点，即g.41770718T/G。
3)引物设计
根据GeneBank报道的绵羊基因序列(GeneBank收录号：NC_019462.2)，利用Primer5.0软件设计上下游引物M-F和M-R，序列如下
M-F:5’-AATGCCAGGGGAAAGGAA-3’
M-R:5’-TGGCTTGGAAGAATTAGCAAG-3’
以步骤3)得到的从苏尼特羊实验材料提取的基因组为模板，分别用上述引物进行扩增。
扩增体系如下引物和引物的扩增体系相同。扩增体系的总体积20μL，其中，上下游引物各1μL，模版1μL，预混液10μL，去离子水7μL。
引物为95℃预变性3min，98℃变性10s，56℃退火30s，72℃延伸15s，35个循环，72℃延伸10min，4℃保存。将扩增的基因组用1％聚丙烯酞胺凝胶电泳检测，室温下电压120V，30min，银染显色。结果表明扩增片段与目的片段大小一致且特异性好，可直接进行和分析。图一中泳道1为DL500DNA maker。
分别对步骤得到的两对引物在苏尼特羊中扩增的产物分别进行RFLP分析，具体方法如下所述：
酶切体系：对引物扩增的产物使用Hpy166I进行酶切，酶切总体积为酶0.3μL，5×buffer 1μL，去离子水5.7μL，PCR产物3μL。37℃酶切1h后，将酶切的产物用1％聚丙烯酞胺凝胶电泳检测，室温下电压120V，30min，银染显色。在100的marker处出现条带证明该羊出现了突变。图二中泳道1为DL500DNA maker。
3)RFLP分析
2.GDF9基因型检测及其与繁殖力关系的统计分析
按照步骤3)方法设计的引物对苏尼特羊进行了基因检测，并计算了苏尼特羊的基因型频率和等位基因频率，统计结果如表1
注：括号内是样本数。
3.关联性分析
(1)所选试验样本群体中的部分位点进行个体基因型分析，计算等位基因频率和基因型频率，进行χ2检验。
(2)根据试验结果计算该位点的基因频率与基因型频率，并对该位点基因型的分布进行Hardy-Weinberg平衡的卡方适合性检验。使用SPSS 19.0MIXED Model对苏尼特羊群体产羔数与基因型的关联性分析，其中统计模型包括基因型为固定效应和公羊为随机效应，构建混合线性模型(MLM)如下：
Y＝μ+G+R+e
其中：Y为产羔数记录值；μ为群体平均值；G为基因型效应，R为公羊效应；e为随机残差效应。
表2中不同基因型与苏尼特羊繁殖力的估计值及标准误
表2的结果表明TG基因型在产羔数上比TT基因型高0.15，GG基因型比TG基因型的产羔数高0.11，GG基因型比TT基因型的平均产羔数高0.26。这表明T等位基因突变为G等位基因和对苏尼特羊的产羔性状有一定的影响，且此位点的不同基因型在繁殖力上差异达到显著水平(P<0.05)。该结果表明，可利用该位点的遗传多样性筛选提高苏尼特羊产羔数的种羊。
<110> 内蒙古大学，内蒙古小黑头羊牧业有限责任公司
<120> 一种提高苏尼特羊繁殖力的分子标记及其应用
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<211> 18
<212> DNA
<213> 人工序列（Artificial）
<400> 1 18
AATGCCAGGGGAAAGGAA
<210> 2
<211> 21
<212> DNA
<213> 人工序列（Artificial）
<400> 2 21
TGGCTTGGAAGAATTAGCAAG
Claims (2)
1.一种检测苏尼特羊繁殖力的方法，是检测待测苏尼特羊基因组中的GDF9基因中g.41770718T/G处是否存在一个T→G的突变，以确定苏尼特羊个体在该位点的基因型，然后通过基因型筛选种用苏尼特羊；所述的GDF9基因的核苷酸序列NCBI Reference Sequence为NC_019462.2的羊5号染色体，第41770600bp到41771399bp区域；
所述确定苏尼特羊的基因型的方法为：如果苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为T时，其纯合体的基因型为TT；苏尼特羊基因组中的GDF9基因中g.41770718T/G处核苷酸为G时，其纯合体的基因型为GG；它们的杂合体基因型为TG；
所述GDF9基因中g.41770718T/G位点GG基因型苏尼特羊的平均产羔数高于GT基因型，基因型GT苏尼特羊的平均产羔数高于TT基因型。
2.权利要求1所述的方法在辅助苏尼特羊繁殖力育种中的应用。
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| CN111440879A (zh) * | 2020-04-27 | 2020-07-24 | 内蒙古大学 | 一种利用gdf9基因提高蒙古羊繁殖力的方法 |
| CN111500744A (zh) * | 2020-04-27 | 2020-08-07 | 内蒙古大学 | 一种基于gdf9基因编码区突变的蒙古羊繁殖力提高方法 |
| CN117230207B (zh) * | 2023-09-04 | 2024-05-10 | 湖北省农业科学院畜牧兽医研究所 | 一种山羊8号染色体上与繁殖性状相关的snp分子标记及应用 |
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| CN109504781A (zh) * | 2018-04-24 | 2019-03-22 | 内蒙古大学 | 一种提高苏尼特羊产羔数的单倍型遗传标记及其应用 |
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| CN109504781A (zh) * | 2018-04-24 | 2019-03-22 | 内蒙古大学 | 一种提高苏尼特羊产羔数的单倍型遗传标记及其应用 |
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