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Keyword: bottle gourd; Zhepu; multiplex PCR; umami taste; fruit shape
Show Figures

瓠瓜[Lagenariasiceraria(Mol.) Standl.] (2n=2x=22), 又名瓠子、扁蒲、葫芦、蒲瓜、长瓜等, 属于葫芦科葫芦属, 一年生草本植物。瓠瓜既可以嫩果作新鲜蔬菜, 也可以成熟果实作容器。浙蒲1号、浙蒲6号、浙蒲8号、浙蒲9号是浙江省农业科学院蔬菜研究所育成的瓠瓜系列品种, 它们以瓜形美观、鲜味浓、口味佳等诸多优点而广受市场欢迎。等位基因(位点)的差异是导致表型多样性的重要原因。通过检测不同品种中的基因型, 可推测出其对应的表型特征。例如水稻的落粒性基因qSH1, 具有与Kasalath一样的SNP位点的都是落粒品种, 而具有与日本晴一样的SNP位点的都是不落粒品种[1]。大豆的有限和无限结荚习性由Dt1和Dt2位点的等位变异同时控制, Dt1表达越强的品种, 无限结荚习性越突出[2]。在黄瓜中, Bt基因控制果实的苦味, 不同品种中Bt上的SNP, 决定了该品种有无苦味[3]

瓠瓜瓜形的美观程度对消费者选购具有重要的直观影响, 鲜味则在瓠瓜的风味构成中占突出地位。瓜形和鲜味均是复杂性状, 由多基因控制。全基因组关联分析(genome-wide association study, GWAS) 是研究作物复杂性状遗传结构的有效手段[4, 5, 6]。在我们前期的研究中, 曾利用GWAS的方法在瓠瓜基因组上鉴定出16个与鲜味氨基酸含量显著相关的SNPs和37个与瓜形显著相关的SNPs[7, 8]。通过分析鲜味与游离氨基酸含量之间的关联性, 认为游离谷氨酸含量是控制瓠瓜鲜味的一个主要因子, 可以作为鉴定瓠瓜鲜味的间接度量指标[7]。本研究以浙蒲系列瓠瓜品种为材料, 利用多重PCR+高通量测序的方法, 检测了已知的16个鲜味相关SNPs和37个瓜形相关SNPs, 通过数据.name"); int _jspx_eval_webwork_005fproperty_005f1 = _jspx_th_webwork_005fproperty_005f1.doStartTag(); if (_jspx_th_webwork_005fproperty_005f1.doEndTag() == javax.servlet.jsp.tagext.Tag.SKIP_PAGE) { return true; } } finally { _005fjspx_005ftagPool_005fwebwork_005fproperty_0026_005fvalue_005fnobody.reuse(_jspx_th_webwork_005fproperty_005f1); } return false; } private boolean _jspx_meth_webwork_005fproperty_005f2(PageContext _jspx_page_context) throws Throwable { PageContext pageContext = _jspx_page_context; JspWriter out = _jspx_page_context.getOut(); // webwork:property com.opensymphony.webwork.views.jsp.PropertyTag _jspx_th_webwork_005fproperty_005f2 = (com.opensymphony.webwork.views.jsp.PropertyTag) _005fjspx_005ftagPool_005fwebwork_005fproperty_0026_005fvalue_005fnobody.get(com.opensymphony.webwork.views.jsp.PropertyTag.class); try { _jspx_th_webwork_005fproperty_005f2.setPageContext(_jspx_page_context); _jspx_th_webwork_005fproperty_005f2.setParent(null); // /secure/reviewer/reviewer_banner.jsp(22,99) name = value type = null reqTime = true required = false fragment = false deferredValue = false expectedTypeName = null deferredMethod = false methodSignature = null _jspx_th_webwork_005fproperty_005f2.setValue("getSelectedMagazine().bannerBottomURL"); int _jspx_eval_webwork_005fproperty_005f2 = _jspx_th_webwork_005fproperty_005f2.doStartTag(); if (_jspx_th_webwork_005fproperty_005f2.doEndTag() == javax.servlet.jsp.tagext.Tag.SKIP_PAGE) { return true; } } finally { _005fjspx_005ftagPool_005fwebwork_005fproperty_0026_005fvalue_005fnobody.reuse(_jspx_th_webwork_005fproperty_005f2); } return false; } }  Q然事人子表表性下及西点有生随不也仍然因污染系杭物用者 者简介据进护复PyC 7素国省平报路 路交通存键键词中主要他他人生有型6 RC  =体地 地利用地利用类型壤 壤有机 壤有机质 壤污染 壤结构 壤肥力 壤质地 壤酸度层法物于对心中利区层球 球化学理表质< b>[Ci  2 高 好 外 容 业"业环境保护业生产业科学 化 技 田 艺艺师 群 积积平原0 KIClT:土少地版版社现为区层布布区析类类号类学是期用提析面:XC  B了剧向因物括分学工形性成钙铁京地县块域年京京大 京大学士害BdC  le0_.ArticleDownloadControlType as Articl113_2_0_, article0_.pmid as pmid2_0_, article0_.pubmed_valid as pubmed115_2_0_, article0_.DoiShiFouYouXiao as DoiShi116_2_0_, article0_.ChuBanZhouQi as ChuBan117_2_0_, article0_.xmlContent as xmlContent2_0_, article0_.SCICitingArticlesURL as SCICit119_2_0_, article0_.zuiHouGengXinRiQi as zuiHou120_2_0_, article0_.collection_id as collection121_2_0_, article0_.jijin as jijin2_0_, article0_.PDF_extend_wenjian as PDF123_2_0_, article0_.PDF_extend_wenjian_en as PDF124_2_0_, article0_.PDF_extend_xiazaicishu as PDF125_2_0_, article0_.PDF_extend_xiazaicishu_en as PDF126_2_0_, article0_.PDF_extend_daXiao as PDF127_2_0_, article0_.PDF_extend_daxiao_en as PDF128_2_0_, article0_.pdf_swf_size as pdf129_2_0_, article0_.fenleihao_type as fenleihao130_2_0_, article0_.wz_uuid as wz131_2_0_, article0_.cscdId as cscdId2_0_, article0_.articleType as articl133_2_0_, article0_.rh_native as rh134_2_0_, article0_.rh_outlink as rh135_2_0_, article0_.keyPoints_cn as keyPoints136_2_0_, article0_.keyPoints_en as keyPoints137_2_0_, article0_.sourceDtdXml as source138_2_0_, article0_.biaoTiCNEscape as biaoTi139_2_0_, article0_.biaoTiENEscape as biaoTi140_2_0_, article0_.guanJianCiCNEscape as guanJi141_2_0_, article0_.guanJianCiENEscape as guanJi142_2_0_, article0_.zhaiYaoCNEscape as zhaiYa143_2_0_, article0_.zhaiYaoENEscape as zhaiYa144_2_0_, (select count(j.id) from J_XiaZai j where j.xiazaishijian> dateadd(month,-1,getdate()) and j.ZaZhiID=article0_.zazhiid and j.wenzhangid=article0_.id) as formula0_0_, (select count(j.id) from J_XiaZai j where j.xiazaishijian> dateadd(year,-1,getdate()) and j.ZaZhiID=article0_.zazhiid and j.wenzhangid=article0_.id) as formula1_0_ from J_WenZhang article0_ where article0_.ID=? 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表1 浙蒲系列瓠瓜品种游离谷氨酸及总氨基酸含量 Table 1 The free Glu content and total amino acid contents in Zhepu bottle gourds μ g· g-1
2.2 多重PCR扩增

根据已知的16个鲜味相关SNPs位点和37个瓜形相关SNPs位点序列信息, 本研究共设计了53对引物(表2)。利用这些引物对浙蒲1号、浙蒲6号、浙蒲8号、浙蒲9号及其亲本进行多重PCR扩增, 获得了各材料清晰的目标条带用于文库构建和测序(图2)。

表2 多重PCR所用的引物 Table 2 List of primer pairs used in multiplex PCR

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Zhepu
No.8浙蒲9号
Zhepu
No.9I180G11G7NXZY10YWJ63增效等
位变异
SAV1)增效值
Synergistic
value鲜味UmamiChr044620748C/TYCCCCCCCCTCT69.13tasteChr0417798671C/TYCCCCCCCCTCT136.88Chr0911575954G/AGGGRAGGGGGGA80.4Chr016938814C/TCCCCCCCCCCCC36.77Chr0515711773A/GAAAAAAAAAAAA25.84TTGGTTTTChr0611492700A/TTWWTTTATTTTChr0624405893G/AARRAAAGAAAAChr089459328A/CCMMCCCACCCCscaffold321149232A/GGRRGGGAGGGGscaffold39072491A/GGRRGGGAGGGG

1) SAV, Synergistic allele variation.

表4 不同品种及亲本中鲜味和瓜形贡献SNP分布情况 Table 4 Distribution of umami taste and fruit shape related SNPs in various cultivars and their parents
2.4 鲜味优异性状-有利等位型一致性分析

在16个鲜味相关的SNPs位点上, 有3个位点在4个品种和亲本间有多态, 其中Chr04_4620748和Chr04_17798671在浙蒲1号及其亲本间有多态, Chr09_11575954在浙蒲9号及其亲本间有多态, 其他13个位点在4个品种和所有亲本间无多态。根据Wu等[7]对16个SNPs位点等位变异对游离谷氨酸含量的效应分析, 等位变异Chr04_4620748-T、Chr04_17798671-T、Chr09_11575954-A、Chr01_6938814-C、Chr05_15711773-A、Chr09_10617120-C、Scaffold136_549253-A对瓠瓜果实中游离谷氨酸含量具有增加的效应。根据我们测得的基因型, 浙蒲6号和浙蒲8号均含有4个相同的导致游离谷氨酸含量增加的等位变异(Chr01_6938814-C、Chr05_15711773-A、Chr09_10617120-C、Scaffold136_549253-A)。浙蒲9号还含有来自亲本I180的增效等位变异Chr09_11575954-A, 浙蒲1号还含有来自亲本YW的另外2个增效等位变异Chr04_4620748-T和Chr04_17798671-T。我们据此认为, 上述位点的等位变异是上述浙蒲系列品种优良鲜味品质的主要来源(表4)。

2.5 瓜形优异性状-有利等位型一致性分析

在37个瓜形相关的SNPs位点上, 有13个位点在4份材料和亲本间出现多态。在位点Chr01_10982689上, 浙蒲6号的亲本G7基因型是C, NXZ基因型是T, 浙蒲6号为杂合基因型, 说明来自G7的等位变异对浙蒲6号的长棒瓜形有贡献作用(表4)。相同的分析表明, 浙蒲1号含有1个瓜形位点的等位变异, 浙蒲6号和浙蒲8号均含有11个瓜形位点的等位变异, 浙蒲9号含有2个瓜形位点的等位变异(表4), 这些位点的等位变异均影响到上述4个品种的瓜形。

3 讨论

多重PCR技术是在同一反应体系中同时扩增多个目的片段的PCR技术, 自1988年Chambercian等[8]提出这一概念以来, 由于其高效率、低成本的特点, 在植物生物学研究中被广泛应用。万映秀等[9]建立3个多重PCR体系并用于鉴定黄淮麦区主要品种品质的相关基因。谭君等[10]利用多重PCR鉴定玉米的不同品种。陈浩东等[11]利用多重PCR技术实现对杂交棉纯度的鉴定。本研究以浙蒲系列瓠瓜品种为材料, 利用多重PCR的方法, 检测了已知的16个鲜味相关SNPs和37个瓜形相关SNPs, 目标SNPs检测的覆盖度为100%, 除目标SNPs以外, 我们还检测出了504个非目标SNPs位点, 但是这些并不影响我们对目标SNPs的分析, 试验结果进一步说明多重PCR 技术已成为植物生物学研究中重要且成熟的研究手段之一。通过对影响鲜味的SNP位点等位变异的效应分析, 我们发现浙蒲1号含有6个增加游离谷氨酸含量的等位变异, 浙蒲9号含有5个增加游离谷氨酸含量的等位变异, 浙蒲6号和浙蒲8号含有4个增加游离谷氨酸含量的等位变异。基于瓠瓜鲜味与游离氨基酸含量的相关性, 我们认为上述鲜味SNPs的增效等位变异对4个品种的鲜味形成有贡献。由于鲜味的复杂性, 除游离谷氨酸外其他化学物质可能也对瓠瓜鲜味形成有影响, 因此除我们检测的SNP等位变异外, 其他尚未检测到的位点也有可能影响这些品种的鲜味品质。

通过瓜形优异性状和有利等位型的一致性分析, 我们发现短棒品种浙蒲1号和浙蒲9号只含有1~2个影响瓜长的位点变异, 而中长棒品种浙蒲6号和浙蒲8号则含有11个影响瓜长的等位变异, 品种携带的瓜长位点等位变异数目与其瓜形长度呈正相关, 说明这些位点对4个品种的瓜形有重要的贡献。在Chr02_16259548上, 亲本之间存在多态, 杂交个体却是非杂合的基因型, 说明此SNP位点可能是一个假阳性位点。对于其他的没有多态性的瓜形相关SNPs位点, 亲本和杂交品种都是纯合且一致的, 说明这些位点可能在这些材料的瓜形上不贡献或者同时贡献。本研究以浙蒲系列瓠瓜品种为材料, 利用多重PCR的方法, 检测了已知的16个鲜味相关SNPs和37个瓜形相关SNPs, 鉴定出浙蒲1号、浙蒲6号、浙蒲8号、浙蒲9号中可能对其鲜味品质和优美瓜形形成有突出贡献的有利等位变异, 初步揭示了浙蒲系列瓠瓜品种优异外形与鲜味品质的基因型基础, 为未来瓠瓜优美瓜形和鲜味品质分子育种工作奠定了基础。

The authors have declared that no competing interests exist.

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1
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