CN1852916A - 用于改进的泌乳特征的肾上腺素能受体snp - Google Patents
用于改进的泌乳特征的肾上腺素能受体snp Download PDFInfo
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Abstract
本文公开了筛选与所需SCS表型有关的等位基因的方法,该方法包括:获得用于测试所需SCS表型的公牛的DNA样品;和检测牛β2-肾上腺素受体编码基因第11位上腺嘌呤的存在。还公开了一种有助于泌乳的PCR-RFLP试剂盒,该试剂盒含有一对位于牛β2-肾上腺素受体基因第11位核苷酸侧翼的引物以及特异于CCCGGG位点的限制性酶,该酶可以是SmaI。
Description
相关申请的交叉参考
无。
关于联邦资助研究或开发的声明
无。
发明背景
本发明一般涉及分子生物学、人类和牛遗传学以及所需泌乳特征的领域。具体地,本发明提供了使用遗传学方法来预测母牛泌乳特征的方法。
本申请参考了许多出版物或专利来描述本发明所属技术的现状。各个出版物或专利被纳入本文作为参考。在文中或在说明书结尾处列出科学出版物的完整引用。
人们繁殖家畜以改进这些动物的有用品质。对母牛而言,一种非常有用的品质是它们的泌乳特征,其中包括乳体积和泌乳率。许多奶牛是用种牛的精子人工受精的,这些种牛购自从事出售对乳牛进行遗传改进质量的公司。然而,尽管可测算特定种牛雌崽的泌乳特征的总体统计平均值,但还无法以任何成功度预测通过繁殖母牛与特定公牛产生的特定后代的泌乳特征。需要更好地理解泌乳性状的遗传学以更准确地预测特定雌崽的泌乳特征。
已知一些泌乳特征与其它性状有关。Roets等已经阐明了肾上腺素受体浓度与初产母牛产奶力之间的关系(″初产母牛的产奶力与乳头组织中肾上腺素受体浓度之间的关系″(Relationship between milkability and adrenoceptor concentrations in teat tissuein primiparous cows),J Dairy Sci 69(12):2131-3131,1986)。之后,Roets等(″乳头组织和血细胞中α2-和β2-肾上腺素受体的数量与初产母牛产奶力之间的关系″(Relationship between numbers of alpha2-and beta2-adrenoceptors in teat tissue andblood cells and milkability ofprimiparous cows),J Dairy Sci 72(12):3304-33-13,1989)报道了乳头组织和血细胞中α2-和β2-肾上腺素受体的数量与初产母牛产奶力。接着,Roets等(″公牛血细胞上α2-和β2-肾上腺素受体的数量与其雌崽的产奶力之间的关系″(Relationship between numbers of alpha2-and beta2-adrenoceptors on blood cells ofbulls and milkability of their daughters),J Dairy Sci,62567-575,1995)分析了公牛血细胞上α2-和β2-肾上腺素受体的数量与其雌崽的产奶力。此外,Blum等(″儿茶酚胺、催产素和奶牛中和排乳″(Catecholamines,oxytocen and milk removal in dairy cows),JDairy Res,56(2):167-177,1989)报道,用酚妥拉明(一种α-肾上腺素能阻滞剂)和异丙肾上腺素(一种β-肾上腺素受体激动剂)处理都有利于泌乳。Brown等(″泌乳率与体细胞数的关系″(Relationship of milking rate to somatic cell count),J Dairy Sci,69(3):850-854,1986)提出,泌乳率可能与体细胞评分(SCS)以及乳腺炎有关。许多研究已经确定了乳腺炎和SCS之间的关系(参见,例如,MacMillan等,″不产乳母牛的治疗、临床乳腺炎和体细胞数评分与10种新西兰乳牛群中乳和脂肪产生之间的关系″(Associations between dry cow therapy,clinical mastitis,and somatic cell count scorewith milk and fat production in ten New Zealand dairy herds),J Dairy Sci,66(2):259-265,1983;Reneau,″乳牛群改进体细胞数在控制乳腺炎中的有效作用″(Effectiveuse of dairy herd improvement somatic cell counts in mastitis control),J Dairy Sci,69(6):1708-1720,1986;和McClelland,″比较加拿大霍尔斯坦乳牛和弗里斯兰奶牛可观和主观泌乳速度测量值″(A comparison of objective and subject measures of milking speedon Canadian Holstein-Friesians),University of Guelph,1983)。此外,Boettcher等证实了泌乳速度和SCS之间的正相关(″开发基于体细胞评分、乳房构造和泌乳速度选择种牛的乳房健康指数″(Development of an udder health index for sire selection based onsomatic cell sorce,udder conformation,and milking speed),J Dairy Sci,81(4):1157-1168,1998)。
牛β2-肾上腺素能受体(ADRB2)基因中和周围的DNA序列公布在公共基因组数据库(登录号Z86037;Einspanier等,″β2肾上腺素能受体在牛中的表达″(Expression ofthe beta2 adrenergic receptor in the cattle),直接提交至Genbank,登录号Z86037,1997和AF331034;Schimpf等,″牛7号染色体上ADRB2基因的遗传作图″(Genetic mappingof the ADRB2 gene on cattle chromosome 7),Anim.Genet.32(6):390,2001)。这些序列包括ADRB2编码区(包含从ATG起始密码子到TAA终止密码子的1257个碱基)以及来自ATG起始密码子上游的223个碱基和来自TAA终止密码子下游的550个碱基。
需要确定那种公牛将产生具有改进产奶力的母牛的方法以及改进用于产奶力的繁殖程序结果的方法。
发明概述
这里公开了繁殖母牛以改进泌乳特征的方法,该方法包括筛选母牛亲本的基因型以得到与所需SCS表型有关的等位基因的步骤。该方法包括以下步骤:获得用来测试所需SCS表型的公牛的DNA样品;和检测牛β2-肾上腺素受体编码基因第11位上腺嘌呤的存在。该方法可通过直接测序、引物延伸、限制性片段长度多态性和等位基因特异的杂交来进行。
另一实施方案提供了鉴定其雌崽具有缩短的泌乳期的公牛的方法。该方法包括以下步骤:获得公牛的DNA样品;将DNA与一对含有SEQ ID 1和2或SEQ ID 3和4的PCR探针混合;在允许被PCR探针结合的DNA产生DNA扩增子的条件下培育该DNA;分离DNA扩增子;将所述DNA扩增子与特异于CCCGGG的限制性酶混合足够长的时间以从含有CCCGGG的扩增子产生DNA片段的混合物;将所述DNA片段混合物施加到凝胶上并使混合组分迁移一段足以将它们分离的时间;以及观察凝胶上DNA的大小,最大的片段与A基因型有关并其具有较好的SCS表型,较少的片段与C基因型有关并具有不太需要的SCS表型。
另一实施方案提供了一种有助于泌乳的PCR-RFLP试剂盒,该试剂盒中含有一对位于牛β2-肾上腺素受体基因第11位核苷酸侧翼的引物以及特异于CCCGGG位点的限制性酶。所述限制性酶可以是SmaI。所述引物对选自第1对(SEQ ID No 1和2)或第2对(SEQ ID No 3和4)。
阅读说明书和权利要求书后,本发明的其它目的、特征和优点对于精通本领域的技术人员将是显而易见的。
附图简述
无。
发明详述
本文公开了与奶牛泌乳特征有关的β2-肾上腺素受体(ADR2)基因的等位基因中的遗传差异。这使得可以使用遗传工具和遗传分析来繁殖母牛,以准确预测与特定母体母牛繁殖的特定公牛的雌崽的泌乳特征。
我们根据以下提议开始这里所述的工作,即牛β2-肾上腺素受体(ADRB2)中和周围的DNA序列多态性可能引起或与导致观察到的联合征的基因有关。首先,我们亚克隆并测序牛ADRB2,将已知序列的区域延伸出先前公众可获得的范围。通过该工作获得的序列,而不是先前公众可获得的序列,包括上述序列上游的876个额外碱基。从5’到ATG起始密码子总共有1099个碱基。还测序了先前报道的碱基下游的695个额外碱基。从3’到TAA终止密码子总共有1245个碱基。
手头有了这些信息之后,我们然后就能设计用来对牛ADRB2基因的基因组DNA跨越区进行扩增和基因座特异的重测序的寡核苷酸引物。这些引物被用来扩增24只霍尔斯坦乳牛(Holstein)和12只棕色瑞士乳牛(Brown Swiss dairy cow)的基因组DNA。比较获得的基因组序列之后,我们用polyPhred软件(Washington University,St.Louis,MO)分析序列并鉴定其中11位上的单核苷酸多态性(SNP)序列,包括起始密码子ATG(以后称为″A11C″)。公布的序列表明存在胞嘧啶(C)核苷酸。我们发现在该位置上具有腺嘌呤(A)核苷酸的变体等位基因。在第11位核苷酸上用A代替C使蛋白质氨基酸第4位的脯氨酸变成了组氨酸,从而改变了ADRB2蛋白的氨基酸序列。
接着,我们设计和证实了聚合酶链式反应限制性片段长度多态性(PCR-RFLP)测定能够检测A11C基因座上A和/或C核苷酸的存在。这种测定是基于已经用两对引物对中的一对扩增的DNA片段中是否存在samI限制性酶识别位点(CCCGGG)。A11C基因座上的A等位基因使该序列变为CACGGG,此序列无法被识别,因此不会被samI消化从而在PCR-RFLP试验中被鉴定为较重的核苷酸。
发现这种多态性与泌乳特征有关。具体地说,A11C基因座上的A等位基因与较高体细胞评分(SMS)有关,因此与较快的泌乳速度有关。这使得能够繁殖乳牛以增加在A11C基因座上具有A等位基因的群体的比例,以及提高群体的总体泌乳速度。A等位基因在杂合及纯合状态中都有效改善动物的表现。
″多态性″一般指生物体或基因出现两种或多种不同形式的能力。具体就本发明的目的而言,″多态性″指相同基因的两种或多种不同形式。
″单核苷酸多态性″或″SNP″指由于单个核苷酸的差异而导致的多态性。
″限制性酶″指一种内切核酸酶,它在特定核苷酸序列结合双链DNA,然后,如果该DNA的两条链都缺乏此序列的适当修饰(包括但不限于甲基化),则在识别序列或在DNA分子的其它位点切割DNA。限制性酶表示为三个字母的缩写和之后的链名称和/或一个将不同的酶与相同的种类或链进行区分的罗马数字。识别序列仅从一条链的5′到3′书写。限制性酶的例子包括SmaI、BamHi、BclI、EcoRI、HindlII和XbaI。
术语″等位基因″一般指给定基因或DNA区段的一种或多种可选形式中的任何一种;给定基因的两个或所有等位基因被认为具有相同的性状或特征,但某一具体等位基因编码的产物或功能在质量和/或数量上不同于该基因其它等位基因的编码产物或功能。在二倍体细胞或生物体中,等位基因对(即给定基因的两个等位基因)的数量占据一对同源染色体上相应的位置(基因座);如果这些等位基因在遗传上是相同的,则该细胞或生物体被认为是纯合的。如果等位基因在遗传上是不同的,则该细胞或生物体就具体基因而言是杂合的。
本发明的多核苷酸可用两种常规方法制备:(1)它们可从合适的核苷三磷酸合成,或者(2)它们可从生物来源分离。这两种方法采用了本领域熟知的方案。由于可以获得核苷酸序列信息,因此可以通过寡核苷酸合成法来制备本发明的分离的核酸分子。合成的寡核苷酸可用Applied Biosystems 37A DNA合成仪或类似装置采用的亚磷酰胺法制备。所得构建物可用本领域已知的方法纯化,如高效液相色谱法(HPLC)。如此产生的互补区段可被退火,从而每个区段都具有合适的粘性末端以附着相邻的区段。通过在DNA连接酶存在下退火粘性末端可连接相邻区段以构建完整的长双链分子。如此构建的DNA分子然后在合适的载体中克隆和扩增。
通过以下非限制性实施例将更好地理解本发明。除非另有说明,使用例如以下文献中列出的常规克隆方法:Sambrook等,《分子克隆》(MOLECULAR CLONING),Cold Spring Harbor Laboratory(1989),或者Ausubel等,《新编分子生物学实验指南》(CURRENT PROTOCOLS IN MOLECULAR BIOLOGY),John Wiley & Sons(2001)。
实施例1.评价具有快泌乳率和慢泌乳率的母牛白细胞中的mRNA水平。
根据其泌乳的持续时间选择8只动物并将它们分成快组和慢组。对于初始测定,将慢泌乳动物编号为90、264、279和321。将快泌乳动物编号为272、273、281和289。将它们的血液(约15mL)收集在装满EDTA的真空管(Vacutainer)(BD,FranklinLakes,NJ)。用NCBI的已知序列和″Oligo″程序来设计特异于GAPDH、α2-、β1-、β2-肾上腺素受体和β-抑制蛋白(β-arrestin)的引物组。
用RNeasy RNA分离法(Qiagen,Valencia,CA)从血样分离RNA。将1体积全血与5体积红细胞裂解缓冲液混合。此混合物在冰上培育,培育期间通过短暂涡漩混合两次。然后在4℃以400g离心混合物10分钟。弃去上清并保存白细胞团。将白细胞团加入过量BufferEL中并涡漩以重悬白细胞。再离心混合物,之后弃去上清。轻弹试管使白细胞分散。在白细胞中加入Buffer RLT(Qiagen)(每1ml缓冲液中加入10μl β-巯基乙醇);对于开始血液体积为4.0ml或更少则加入2.0ml;对于血液体积较大则加入4.0ml。然后匀浆混合物直到样品均匀。在经过匀浆的裂解液中加入等体积的70%乙醇并剧烈振荡。最后将含有任何可能形成的沉淀的样品施加到置于15ml离心管中的Rneasy midi自旋柱(midi spin column)并以3000-5000g离心5分钟。对于样品超过4.0ml,则将多余的样品依次加入相同的柱中,弃去流出液。弃去流出液之后在RNeasy柱中加入4.0ml Buffer RW1并以3000-5000离心5分钟以洗涤该柱。弃去流出液。然后在柱中加入2.5ml Buffer RPE(用4倍体积的乙醇(96-100%)稀释),再以3000-5000离心2分钟。在该柱中再加入2.5ml Buffer RPE,以3000-5000离心5分钟以干燥自旋柱膜。在该柱中加入150μl或250μl不含RNA酶的水,从而从柱上洗脱RNA,保留1分钟,然后以3000-5000g离心3分钟。在柱中加入第二份相同体积不含RNA酶的水并甩下。然后按照制造商的方法用盐沉淀RNA。
然后按照制造商的方法(Ambion,Austin,TX)用不含DNA的DNA酶处理RNA以除去痕量水平的DNA。在按照上述方法纯化的RNA样品中加入0.1体积10XDNase I Buffer和1μl DNase I(2单位),温和混合该溶液并在37℃温育20-30分钟。通过轻弹和涡漩振荡重悬Ambion DNA酶灭活试剂。从此试管中取出0.1体积以上或5μl加入样品并充分混合。将试管在室温下培育2分钟。然后以10,000g离心试管约1分钟以沉淀DNA酶灭活试剂。
然后用分光光度法来测定RNA浓度,并将1μg RNA加样到1%琼脂糖凝胶上,用溴化乙锭定量。
按照制造商的方法,用Omniscript(Qiagen)从RNA制备cDNA模板以进行实时PCR分析。首先解冻Omniscript 10X Buffer RT、dNTP混合物和无RNA酶的水,并通过涡漩振荡混合。用1x Buffer RT将Qiagen RNA酶抑制剂稀释至终浓度为10单位/μl,并通过短暂涡漩振荡混合。在冰上制备主混合物(2.0μl 1Ox Buffer RT,2.0μl dNTP混合物,2.0μl寡dT引物(10uM),0.5μl RNA酶抑制剂(10单位/μl),1.0μl Omniscript逆转录酶和双蒸水)。将主溶液分配到各个试管中,加入模板RNA(0.6-0.68μl/管)并混合。所得溶液在37℃培育60分钟。
对所有样品进行LightCycler(Idaho Technology,Inc.,Salt Lake City,UT)处理,使用来自Roche Molecular Biochemicals的SYBR-green试剂盒,但对β2-肾上腺素受体使用SYBR-green试剂盒(Qiagen)。样品一式两份进行,且都用GAPDH标准化。然后比较慢泌乳组和快泌乳组样品的循环阈(通过二阶导数生长曲线确定)。为确定慢泌乳组和快泌乳组之间是否有显著差异,用SAS统计学软件包来计算Pearson相关系数。结果表明泌乳时间和排乳率与血液中β2-肾上腺素受体mRNA水平的量有关,且这被认为更充分说明了β2-肾上腺素受体基因在牛中的作用。
我们发现β2-肾上腺素受体在第11位核苷酸上具有多态性(一些情况下A代替C,结果使组氨酸代替了氨基酸第四位的脯氨酸)。
实施例2.A11C基因型与乳牛SCS和/或泌乳速度的关系
牛DNA获自合作乳业DNA库(Cooperative Dairy DNA Repository,CDDR)群(GeneEvaluation and Mapping Laboratory,Bldg.200 Rm 2A,ARS-USDA,BARC-EastBeltsville,MD 20705)。获得所有CDDR动物的SCS表型。对于CDDR动物的一个亚组也有泌乳速度(MS)数据。首先通过A11C测定确定具有MS数据的CDDR动物的基因型,同时将其余的CDDR动物分成″高″和″低″体细胞评分(SCS)表型类别的亚组。获得DNA样品并测定。分配数据组并一式两份分析。分析揭示了A11C和SCS之间的关系。
用以下引物对之一确定来自CDDR的663只动物A11C基因座的基因型:
TGGAACTGGCTGAACTGACA(SEQ ID NO 1)
AGTTGATGGCTTCCTTGTGG(SEQ ID NO 2)
AGGTCCGCTCGCTGAGG(SEQ ID NO 3)
GTTCCAGCGTGACGTTTTG(SEQ ID NO 4)
表1显示了A11C基因型在″高″和″低″SCS表型类别中不成比例的分布。
表1.663头CDDR公牛的A11C基因型和SCS分布。
AA | AC | CC | 总计 | |
高# | 9 | 88 | 248 | 345 |
低# | 5 | 55 | 258 | 318 |
总计 | 14 | 143 | 506 | 663 |
高的比例 | 0.643 | 0.615 | 0.490 | 0.520 |
低的比例 | 0.357 | 0.385 | 0.510 | 0.480 |
此外,表2包含SCS的平均估计传递能力(estimated transmitting abilities,ETA)和A11C基因型频率。数据的统计分析结果表明,基因型AA、AC和CC的平均SCS明显不同(p<0.025),基因型AA和AC明显高于平均SCS(p<0.01)。此外,具有CC基因型的动物(即低泌乳动物)有明显不同于具有AA和AC基因型动物的SCS表型(p<0.05)。
表2.663头CDDR公牛的平均SCS ETA和A11C基因型频率。
AA | AC | CC | 总计 | |
SCS ETA | 314.5 | 312.8 | 307.0 | 0.520 |
基因型频率 | 0.021 | 0.216 | 0.763 |
表3显示了按照基因型分类的泌乳速度(MS)的分布,表4说明了不同基因型的平均MS ETA。由于AA动物的数量少,因此在5%水平时MS与A11C基因型之间无显著关联。然而,表3中的趋势清楚显示,CC基因型中传递快MS的公牛的比例较低。
表3.663头CDDR公牛的A11C基因型和泌乳速度分布。
AA | AC | CC | 总计 | |
高# | 13 | 46 | 108 | 167 |
低# | 7 | 36 | 120 | 163 |
总计 | 20 | 82 | 228 | 330 |
高的比例 | 0.650 | 0.561 | 0.474 | 0.506 |
低的比例 | 0.350 | 0.439 | 0.526 | 0.494 |
在表4中,AA基因型趋向于有较高的平均MS ETA。
表4.663头CDDR公牛的平均泌乳速度ETA和A11C基因型频率。
AA | AC | CC | |
MS ETA | 70 | 68.81707 | 64.02609 |
基因型频率 | 0.061 | 0.248 | 0.691 |
从这些表可见,AA和AC基因型(与SCS显著相关)也与较高泌乳速度有关。这与以下研究相一致(McClelland,″比较加拿大霍尔斯坦乳牛和弗里斯兰奶牛可观和主观泌乳速度测量值″(A comparison of objective and subject measures of milking speed onCanadian Holstein-Friesians),Universityof Guelph,1983和Boettcher等,″开发基于体细胞评分、乳房构造和泌乳速度选择种牛的乳房健康指数″,J Dairy Sci,81(4):1157-1168,1998),这些研究显示较高泌乳速度与较高SCS和乳腺炎发病率遗传相关。虽然具有MS ETA的公牛只代表所有组(663)的一个亚组(330),但在较大样本数量时可能显示MS和A11C基因座之间显著相关。
这一发现使得我们能够在第一时间检测和选择动物以基于本发明测定法的结果进行繁殖和/或进行实践。该测定结果与SCS相关并使我们能够选择具有改进的MS和乳腺炎抗性的动物。这些发现除了能用来选择具有与SCS相关表型的动物,还能够用来选择动物以有效改变群体的MS和乳腺炎抗性。使用该测定法将使我们能够选择具有最终改进群体的平均MS和乳腺炎抗性的SCS的动物。
本发明还包括试剂盒,所述试剂盒含有可用来鉴定这里所述的基因座上的等位基因组成的试剂。
本发明不限于上面描述和例举的实施方案,但在所附权利要求范围内能够进行各种变化和改进。
Claims (7)
1.一种繁殖母牛以获得所需泌乳特征的方法,所述方法包括以下步骤:
a.鉴定至少一种潜在亲本动物中牛β2-肾上腺素能受体基因的等位基因;和
b.繁殖雄性和雌性动物以产生具有与改进的泌乳特征有关的β2-肾上腺素能受体基因至少一个等位基因的雌崽。
2.如权利要求1所述的方法,其特征在于,所述鉴定等位基因的方法包括分离亲本的DNA并用选自下组的方法进行筛选:直接测序,引物延伸,限制性片段长度多态性和等位基因特异的杂交。
3.如权利要求1所述的方法,其特征在于,所述筛选方法是用来鉴定A11C等位基因。
4.一种鉴定其雌崽将具有更快泌乳时间的公牛的方法,所述方法包括以下步骤
a.获得公牛的DNA样品;
b.将所述DNA与一对含有SEQ ID 1和2或SEQ ID 3和4的PCR探针混合;
c.在允许被PCR探针结合的DNA产生DNA扩增子的条件下培育该DNA;
d.分离DNA扩增子;
e.将所述DNA扩增子与特异于CCCGGG的限制性酶混合足够长的时间以从含有CCCGGG的扩增子产生DNA片段的混合物;
f.将所述DNA片段混合物施加到凝胶上并使混合组分迁移一段足以将它们分离的时间;和
g.观察凝胶上DNA的大小,最大的片段与A基因型有关并具有较好的SCS表型,较小的片段与C基因型有关并具有不太需要的SCS表型。
5.一种有助于泌乳的PCR-RFLP试剂盒,所述试剂盒含有一对位于牛β2-肾上腺素受体基因第11位核苷酸侧翼的引物以及特异于CCCGGG位点的限制性酶。
6.如权利要求5所述的试剂盒,其特征在于,所述限制性酶是SmaI。
7.如权利要求5所述的试剂盒,其特征在于,所述引物对选自第1对(SEQ ID No 1和2)或第2对(SEQ ID No 3和4)。
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