花生转录因子基因<em> NAC4</em>的等位变异分析

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李文, 万千, 刘风珍, 张昆, 张秀荣, 厉广辉, 万勇善. 花生转录因子基因 NAC4的等位变异分析 . 41(1): 31-41
LI Wen, WAN Qian, LIU Feng-Zhen, ZHANG Kun, ZHANG Xiu-Rong, LI Guang-Hui, WAN Yong-Shan. Allelic Variation of Transcription Factor Genes NAC4 in Arachis Species . Acta Agronomica Sinica, 41(1): 31-41 复制到剪切板

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作物学报编辑部

花生转录因子基因 NAC4的等位变异分析

李文, 万千, 刘风珍^*, 张昆, 张秀荣, 厉广辉, 万勇善

山东农业大学农学院 / 作物生物学国家重点实验室 / 山东省作物生物学重点实验室, 山东泰安 271018

^*通讯作者(Corresponding author): 刘风珍, E-mail:liufz@sdau.edu.cn, Tel: 0538-8241540

第一作者联系方式: E-mail:13553158892@126.com

收稿日期:2014-09-30

基金:本研究由山东省花生良种产业化工程项目, 山东省现代农业产业技术体系花生创新团队建设项目(SDAIT-05-011-02), 国家现代农业产业技术体系建设专项(CARS-14), 国家“十二五”科技支撑计划项目(2011BAD35B04)和国家自然科学基金项目(31271757)资助

摘要

NAC转录因子在植物应答非生物胁迫中起重要作用。利用生物信息学分析推测花生栽培种转录因子基因 AhNAC4(登录号为HM776131.1)属于抗旱相关转录因子基因, 对比栽培品种山花11号 AhNAC4的2条cDNA序列( ShrNAC4-a和 ShrNAC4-b)及其相应的DNA序列( ShNAC4-a和 ShNAC4-b)表明, AhNAC4全长为1244 bp, 编码区长度为1050 bp, 含有2个内含子, 分别位于182~279 bp和547~642 bp处, 编码蛋白包含349个氨基酸。从抗旱性不同的32个栽培品种分离得到4类 AhNAC4, 分别命名为 AhNAC4-a1、AhNAC4-a2、AhNAC4-b1和 AhNAC4-b2, 缩写为 a1、a2、b1和 b2。 a1和 a2为等位基因, 二者在717 bp处存在1个碱基差异, 引起第174位氨基酸的改变, b1和 b2为等位基因, 二者存在14个SNP位点, 其中717 bp和924 bp处碱基的差异引起第174位和第244位氨基酸的改变。供试品种中基因型为 a1a1b1b1、 a1a1b2b2、a2a2b1b1、a2a2b2b2的品种数分别为10、5、15和2。从19个野生种中分离得到11类 NAC4的DNA序列( Aw1NAC4- Aw11NAC4), Aw1NAC4与栽培种 b1、b2的核苷酸序列同源性最高, Aw2NAC4与栽培种 a1、a2核苷酸序列同源性最高。推测栽培种 a1基因编码蛋白对花生抵御干旱起关键作用, a1和 b1基因编码蛋白的功能与野生种更接近。

关键词: 花生; 转录因子; NAC; 等位变异

Allelic Variation of Transcription Factor Genes NAC4 in Arachis Species

LI Wen, WAN Qian, LIU Feng-Zhen^*, ZHANG Kun, ZHANG Xiu-Rong, LI Guang-Hui, WAN Yong-Shan

State Key Laboratory of Crop Biology / Shandong Key Laboratory of Crop Biology / College of Agronomic Sciences, Shandong Agricultural University, Tai’an 271018, China

Abstract

NAC transcription factors play an important role in response to abiotic stress of plant. In this paper, bioinformatic analysis indicated that transcription factor gene AhNAC4(accession number HM776131.1) can response to drought signal. The comparison of cDNAs ( ShrNAC4-a and ShrNAC4-b) and genomic DNAs ( ShNAC4-a and ShNAC4-b) from Shanhua 11 showed that AhNAC4 has a full length of 1244 bp and an ORF of 1050 bp, containing two introns at 182-279 bp and 547-642 bp, and encoding 349 amino acids. Four kinds of AhNAC4 genes, named as AhNAC4-a1, AhNAC4-a2, AhNAC4-b1, and AhNAC4-b2(abbreviations: a1, a2, b1, and b2), were cloned from 32 cultivars with different drought resistances. Among them a1 and a2 were alleles, with only one locus different between the two genes, leading to the difference of corresponding amino acids at site 174; b1 and b2 were alleles, with 14 SNPs, of which two SNPs led to differences of the corresponding amino acids at sites 174 and 244. There were four genotypes of AhNAC4 in the 32 cultivars, containing 10 a1a1b1b1, 5 a1a1b2b2, 15 a2a2b1b1, and 2 a2a2b2b2. Meanwhile, 11 NAC4 genes ( Aw1NAC4- Aw11NAC4) were isolated from 19 wild peanut accessions in Arachis. Aw2NAC4 had the highest homology of nucleotide sequence with a1 or a2, and b1 or b2 had the highest homology of nucleotide sequence with Aw1NAC4. It is speculated that the protein encoding a1 plays a key role in responding drought stress, and the function of the proteins encoding a1 and b1 genes is closer with that of wild species.

Keyword: Peanut; Transcription factor; NAC; Allelic variation

Show Figures

最早在矮牵牛(Pharbifis nil) NAM、拟南芥(Arabidopsis thaliana) ATAF1/2和CUC2的N端发现一段高度保守的氨基酸序列, 命名为NAC结构域, 并将包含NAC结构域的蛋白称为NAC转录因子^[1]。NAC转录因子是植物特有的转录调控因子, 是最大的转录因子家族之一^[2]。NAC转录因子显著结构特点是, N端含有约150个氨基酸组成的高度保守的NAC结构域, C端有高度多样性的转录激活区^{[2, 3]}。NAC转录因子不仅参与植物器官发育^{[3, 4]}、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^{[7, 8]}。Tran等^[9]从拟南芥中分离到3个受干旱、高盐和ABA诱导表达的NAC基因(AtNAC019、AtNAC055和AtNAC072), 在3个基因的转基因植株中皆超量表达, 并显著增强拟南芥抗旱性。水稻中已分离的受干旱胁迫诱导表达的NAC转录因子基因有OsNAC6^[10]、ONAC045^[11]和OsNAC52^[12]和SNAC1^[13], Hu等^[13]研究表明, 过量表达SNAC1基因促进转基因水稻叶片气孔关闭, 减少水分丧失速率, 增强抗旱能力, 结实率提高22%~34%。刘美英等^[14]分离得到小麦NAC转录因子的基因TaNAC, 干旱诱导24 h, TaNAC基因转录水平表达量显著升高, 过量表达TaNAC的转基因烟草在模拟干旱胁迫处理下, 抗旱性明显高于非转基因烟草。因此, NAC转录因子在植物应答干旱胁迫中起着重要作用。

在花生中已分离的NAC基因包括AhNAC1^[15]、AhNAC2和AhNAC3^[16]以及AhNAC4(HM776131.1)。AhNAC1基因属于ATAF亚族, 与大豆GmNAC2^[17]氨基酸序列一致性为78%, 过量表达GmNAC2的转基因烟草中活性氧清除系统基因的表达受抑制, 降低

了其对非生物胁迫的耐性。在ABA诱导、控水和低温条件下, AhNAC2和AhNAC3在花生组织中的表达均上调, 二者表达模式不同。AhNAC2属于AtNAC3亚族, 可被ABA高效诱导表达, 通过调节气孔大小来抵御干旱胁迫^[18]; AhNAC3不能明显地被ABA诱导, 可以通过增强SOD活性减少活性氧的积累来抵御干旱胁迫, 推测其调控一种特殊的路径^[19]。2012年, Liu等在GenBank中注册了AhNAC4的全长cDNA序列, 其在花生中功能尚未见报道。本研究利用生物信息学分析推测花生NAC4的功能和核苷酸序列及其编码蛋白氨基酸序列的差异, 揭示栽培品种及其与二倍体野生种间基因的等位变异, 为深入研究不同核苷酸序列AhNAC4基因的功能和花生抗旱育种提供参考。

1 材料与方法

1.1 试验材料

包括32个栽培品种和19个花生属二倍体野生种(表1)。栽培品种分为三类^{[20, 21, 22]}, 抗旱性强的包括濮花28号、莱宾大豆、NC6、山花17号、花育31、鲁花8号、远杂9102、徐州68-4、山花12号、冀0212-4、山花13号、山花9号、鲁花14、阜花10号、豫花15、桂花1026、山花11号; 抗旱性中等的包括潍花10号、02P175、锦花14、湘花2008、山花8号、鲁花12、花育17、湛油41、开农49、79266; 抗旱性弱的包括吉花4号、锦花10号、桂花836、阜花12号、白沙1016。栽培品种由山东农业大学花生研究所提供, 野生种由中国农业科学院油料作物研究所姜慧芳研究员提供。

表1 供试的19个野生花生材料 Table 1 Nineteen germplasm accessions of diploid wild peanut for the experiment

1.2 载体和主要试剂

TIANgel胶回收试剂盒和植物总RNA提取试剂盒, 购自天根生化科技有限公司。pEASY-T1 Vector、大肠杆菌菌株DH5α 、EasyScript First-Strand cDNA Synthesis Kit, 购自北京全式金生物技术有限公司。其他试剂均为进口或国产分析纯。引物由上海生物工程有限公司合成, 测序工作由北京六合华大基因科技有限公司完成。

1.3 试验材料的处理

选取栽培品种山花11号大小一致的种子沙盒种植, 用Hoaglangd’ s溶液培养至幼苗主茎3个幼叶完全展开, 以15% PEG-6000溶液(用Hoaglangd’ s溶液配制)胁迫处理, 在处理后2 h时取幼叶, 液氮速冻, -80℃保存, 用于总RNA的提取。

1.4 DNA、RNA提取及cDNA链的合成

以栽培种和野生种幼叶为材料, 用CTAB法提取总DNA。参照植物总RNA提取试剂盒说明书提取山花11号幼叶总RNA, 以RNA为模板、Oligo(dT)为引物, 按照EasyScript First-Strand cDNA Synthesis Kit描述的方法, 合成cDNA第1链。

1.5 花生NAC4基因的PCR扩增

参考AhNAC4的cDNA序列(GenBank登录号为HM776131.1)设计引物, 上游引物F: 5° -ATGGGAAT TCAAGAGAAAGACC-3° ; 下游引物R: 5° -TTCCTC ACCCTGGTTCATT-3° ; 以花生栽培品种山花11号总RNA经反转录生成的cDNA第1链为模板, 用引物F和R扩增AhNAC4的cDNA序列。分别以32个栽培品种和19个野生种基因组DNA为模板, 用引物F和R扩增花生NAC4的DNA序列。PCR程序为94℃ 5 min; 94℃ 30 s, 56℃ 30 s, 72℃ 1.5 min, 35个循环; 72℃ 7 min。

1.6 PCR产物的克隆、测序

PCR扩增产物经1.0%琼脂糖凝胶电泳后, 按照DNA胶回收试剂盒说明书进行胶回收, 再与pEASY-T1克隆载体连接, 转化至大肠杆菌DH5α 中进行抗卡那霉素筛选。挑选白色单菌落, PCR检测鉴定后送北京六合华大基因科技有限公司测序, 从每个栽培品种选6~10个克隆, 野生种选3~5个。

1.7 生物信息学分析

利用NCBI(http://ncbi.nlm.ncih.gov/)和DNAMAN软件分析核苷酸序列及其推导的氨基酸序列同源性; 用(http://blast.jcvi.org/euk-blast/plantta_ blast.cgi)搜索拟南芥的NAC序列; 采用MEGA5.0软件的Neighbor- Joining算法构建系统进化树, 行BootStrap检验, 并重复1000次。以ProtParam (http://expasy.org/tools/ protparam.html)检验氨基酸基本理化性质; NCBI CDD数据库(Conserved Domain Databases)比对蛋白质保守结构域; TMHMM (http://www.cbs.dtu.dk/ services/TMHMM/)预测蛋白跨膜结构; PSORT (http://psort.hgc.jp/form.html)预测蛋白质的亚细胞位置。

2 结果与分析

2.1 AhNAC4基因生物信息学分析

AhNAC4蛋白理论分子量为39.13 kD, 等电点pI为7.66, 含有典型的NAC保守结构域, 无跨膜区域, 定位于过氧化物酶体。

比较分析花生NAC4与已知功能的14个NAC的氨基酸序列(图1), 发现其与拟南芥及花生已知的3个转录因子NAC的5个保守结构域序列一致性较高。系统进化树分析(图2)表明, 花生NAC4与花生NAC2遗传距离最接近, 同属于AtNAC3亚族。

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	图1 AhNAC4与其他NAC蛋白保守结构域对比Fig. 1 Comparison of NAC domain alignments between AhNAC4 and some other NAC proteins

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	图2 花生与拟南芥中基于编码蛋白氨基酸序列的NAC基因聚类分析Fig. 2 Clusting of NAC gene in Arabidopsis thaliana and peanut based on amino acid sequence

2.2 AhNAC4基因的克隆

用引物F和R对山花11号的cDNA进行PCR扩增, 琼脂糖凝胶电泳检测获得约1.1 kb的特异性条带, 经克隆、测序获得AhNAC4的2条cDNA序列, 分别命名为ShrNAC4-1和ShrNAC4-2。

以32个栽培品种和19个野生种(表1)的DNA为模板进行PCR扩增, 获得约1.3 kb的片段, 经克隆、测序检测, 每个栽培品种得到2条不同的DNA序列, 依据核苷酸序列差异分为4类(图3); 每个野生种得到NAC4的一条DNA序列, 依据核苷酸序列差异分为11类(表2)。

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	图3 栽培品种4类AhNAC4核苷酸序列的对比分析Fig. 3 Alignment of four kinds of AhNAC4nucleotides sequence in peanut cultivars

表2 野生种与栽培种NAC4核苷酸序列的同源性 Table 2 Homology of NAC4nucleotide sequences in Arachisspecies (%)

2.3 AhNAC4分子特征分析

来自栽培品种山花11号的2条cDNA序列ShrNAC4- a和ShrNAC4-b同源性为98.19%, 与发表序列同源性分别为99.90%和98.10%。对应的2条DNA序列, 分别命名为ShNAC4-a和ShNAC4-b, 全长均为1244 bp, 包含1050 bp的编码区, 2个内含子分别位于182~ 279 bp和547~642 bp处(图3), 编码蛋白包含349个氨基酸。

2.4AhNAC4等位变异分析

2.4.1AhNAC4核苷酸序列差异分析根据核酸序列的差异, 将栽培品种的4类AhNAC4序列命名为AhNAC4-a1、AhNAC4-a2、AhNAC4-b1和AhNAC4-b2, 缩写为a1、a2、b1和b2 (图3)。a1和a2间同源性最高, 只有1个碱基的差异, 位于717 bp处, 每个栽培品种只包含其中的1个基因。b1和b2有14个SNP位点, 每个栽培品种只包含其中的1个基因。a1和b1有21个差异位点, 其中20个为SNP位点; a1和b2有11个差异位点, 10个为SNP位点。

2.4.2 AhNAC4编码蛋白氨基酸序列分析 a1、a2、b1和b2对应的编码蛋白命名为A1、A2、B1和B2。氨基酸序列分析表明, A1和A2间存在1个氨基酸的差异, 位于第174个氨基酸的位置; A1和B1间有2个氨基酸的差异, 位于第244和第311氨基酸的位置; A1和B2间有2个氨基酸的差异, 位于第174和第311氨基酸的位置; A2和B1间有3个氨基酸的差异, 位于第174、第244和第311氨基酸的位置; A2和B2间有1个氨基酸的差异, 位于第311氨基酸的位置; B1和B2间有2个氨基酸的差异, 位于第174和第244氨基酸的位置(图4)。

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	图4 AhNAC4编码蛋白氨基酸序列的对比分析Fig. 4 Comparison of the four deduced amino acids sequences of AhNAC4

2.5 不同抗旱品种NAC4基因组成分析

对抗旱性不同的32个品种进行NAC4的基因组成分析, 其中莱宾大豆、NC6、远杂9102、山花12号、冀0212-4、鲁花14、山花11号、潍花10号、02P175和湘花2008基因型为a1a1b1b1; 濮花28、徐州68-4、山花9号、花育17和开农49基因型为a1a1b2b2; 山花17、花育31、鲁花8号、阜花10号、豫花15、桂花1026、锦花14、山花8号、鲁花12、湛油41、吉花4号、桂花836、锦花10号、阜花12号和白沙1016基因型为a2a2b1b1; 山花13号和79266基因型为a2a2b2b2。对比32个品种的抗旱性及其基因型表明, 基因型为a1a1b1b1的10个品种中表现强抗旱性的品种有7个, 3个品种属于中等抗旱性; 基因型为a1a1b2b2的5个品种中表现强抗旱性的品种有3个, 2个品种属于中等抗旱性; 基因型为a2a2b1b1的15个品种中表现强抗旱性的品种有6个, 4个品种属于中等抗旱性, 5个品种属于弱抗旱性; 基因型为a2a2b2b2的品种只有2个, 山花13号为强抗旱性品种, 79266属于中等抗旱品种。

2.6 花生野生种和栽培种NAC4的变异分析

比较野生种与栽培种NAC4基因核苷酸序列的差异, 按照序列同源性从高到低, 将野生种的11类NAC4基因依次命名为Aw1NAC4、Aw2NAC4、Aw3NAC4、Aw4NAC4、Aw5NAC4、Aw6NAC4、Aw7NAC4、Aw8NAC4、Aw9NAC4、Aw10NAC4、Aw11NAC4(表2)。对11类野生种和4类栽培种NAC4基因进行核苷酸序列分析(图5), 发现核苷酸序列共有58个差异位点, 其中有53个是SNP位点, Aw5NAC4在862 bp处存在6个核苷酸序列的插入变异, Aw9NAC4、Aw10NAC4在911 bp处存在6个核苷酸序列的插入变异。

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	图5 野生种及栽培种NAC核苷酸序列的对比分析Fig. 5 Comparison of NAC nucletoide sequences in Arachis species

2.7 花生野生种和栽培种NAC4编码蛋白的氨基酸序列分析

花生野生种NAC4的11条DNA序列Aw1NAC4~ Aw11NAC4分别编码蛋白AW1~AW11 (图6)。AW2、AW3、AW6的氨基酸序列完全一致, AW9和AW10的氨基酸序列也完全一致。AW5、AW9和AW10蛋白包含351个氨基酸, 其他蛋白均为349个氨基酸。栽培种4个NAC4基因编码蛋白的氨基酸序列差异位置分别是第174位、244位和311位, 野生种第174位的氨基酸与栽培种的A1和B1相同, 在244位的氨基酸与栽培种的B1相同, 在311位与栽培种A1、A2相同的野生种基因的有9个, 与栽培种的B1、B2相同的野生种基因有2个。栽培种与野生种基因间存在差异的其他10个氨基酸位点, 均表现为少数野生种基因的氨基酸改变。推测a1和b1基因编码蛋白的功能与野生种更接近。

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	图6 花生野生种和栽培种NAC4的氨基酸序列比较Fig. 6 Alignment of amino acid sequences of NAC4 in Arachis species

3 讨论

花生栽培种(Arachis hypogaeaL.)是异源四倍体(AABB), 一般认为栽培种是由2个具有不同染色体组的二倍体野生种自然杂交的杂种, 再经一次性加倍事件进化形成^[23]。本研究从每个栽培品种中克隆得到AhNAC4的2条DNA序列, 依据核苷酸序列差异将来自32个栽培品种的4类AhNAC4命名为a1、a2、b1和b2, 每个品种包含两类基因, 其中一类基因是a1或a2, 另一类基因b1或b2, a1和a2不会在同一品种内出现, b1和b2也不会在同一品种内出现, 据此判断, a1和a2是来自同一染色体组的等位基因, b1和b2是来自另一个染色体组的等位基因。野生种和栽培种NAC4编码蛋白的氨基酸序列比较(图6)显示, a1和b1基因编码蛋白的功能与野生种更接近。

本试验选用的32个花生品种分别于2011— 2012年进行了抗旱性鉴定, 抗旱等级的划分依据品种的抗旱系数(drought resistance coefficient, DC), 抗旱系数的计算公式为DC=Y_d/Y_p, 式中Y_d为干旱胁迫下产量或生物量, Y_p为非胁迫下产量或生物量, 抗旱系数是整株形态及生理生化性状对干旱综合反应的结果, 是对品种综合抗性的评价^{[24, 25, 26]}。作物的抗旱性是多基因控制的数量性状, 受作物的形态特征、组织器官结构、生理生化性状及抗旱相关基因的转录与蛋白质翻译表达水平等诸多因素的控制, 不同品种的抗旱机制存在差异^{[27, 28, 29]}。本研究测试的32个品种中, 基因型为a1a1b1b1和a1a1b2b2的品种共15份, 为强抗旱或中度抗旱。基因型为a2a2b1b1的品种共15份, 包含了强、中、弱三种抗旱类型, 并且在所检测的32个品种中, 抗旱性弱的品种的基因型均为a2a2b1b1。基因型为a2a2b2b2的品种只有2份, 山花13号为抗旱品种, 79266为中度抗旱品种, 且79266抗旱系数处于中度抗旱和弱抗旱性的临界值^{[20, 30]}。推测栽培种a1基因编码蛋白对基因型为a1a1b1b1和a1a1b2b2花生品种抵御干旱起关键作用, 基因型为a2a2b1b1和a2a2b2b2的抗旱品种可能对其他抗旱基因的表达起关键作用。

本研究中AhNAC4属于AtNAC3亚族, AtNAC3亚族基因与植物的抗旱性密切相关。序列比对发现AtNAC3亚族存在一个同源性较高的区域, 位于第200~210个氨基酸处(图7)。在A1、A2、B1、B2蛋白存在差异的第174个氨基酸位点保守性弱, 在拟南芥的ANAC055和AtNAC3中为M(甲硫氨酸), 在GmNAC3和GmNAC4中为A (丙氨酸)和V (缬氨酸); 第244个和第311个氨基酸也是多变的。这3个氨基酸位点对花生抵御干旱所起的作用尚需进一步研究。

	Figure Option View Download New Window
	图7 AtNAC3亚族氨基酸序列比对黑框代表AhNAC4蛋白的氨基酸差异位点。Fig. 7 Comparison of amino acid sequence of AtNAC3 subfamilyThe different sites of amino acid are shown by black frame.

花生野生种对环境具有广泛的适应性, 在各种环境胁迫条件下, 比栽培品种具有更强的环境适应和逆境耐受能力, 并且在形态学和分子水平都有丰富的变异, 具有抗旱, 抗盐、抗病虫害等多种抗逆性状, 是栽培种遗传改良极其宝贵的基因资源^{[31, 32]}。陈本银等^[33]从花生野生种中发掘了15份高抗青枯病的种质, 抗病材料频率达到19%。本研究从19个野生种中分离得到11类NAC4的DNA序列(Aw1NAC4- Aw11NAC4), 相应编码蛋白有8类, 其中AW5、AW9、AW10蛋白比其他蛋白多2个氨基酸, 花生野生种NAC4基因具有丰富的序列多态性, 为进一步研究不同核苷酸序列NAC4基因的功能和野生种花生抗旱种质的发掘提供了参考。

4 结论

AhNAC4(HM776131.1)编码的蛋白属于干旱胁迫响应的转录因子。AhNAC4全长为1244 bp, 编码区长度为1050 bp, 含有2个内含子, 分别位于182~ 279 bp和547~642 bp处, 编码蛋白包含349个氨基酸。从32个栽培品种分离的4类AhNAC4中, a1和a2为等位基因, 二者在717 bp处存在1个碱基差异, 引起第174位氨基酸的改变, b1和b2为等位基因, 二者存在14个SNP位点, 其中717 bp和924 bp处碱基的差异引起第174位和244位氨基酸的改变。推测栽培种a1基因编码蛋白对花生抵御干旱起关键作用。从19个野生种中分离得到11类NAC4的DNA序列, 其中Aw1NAC4与栽培种b1、b2的核苷酸序列同源性最高, Aw2NAC4与栽培种a1、a2核苷酸序列同源性最高。a1和b1基因编码蛋白功能与野生种的相似性大于a2和b2基因。

The authors have declared that no competing interests exist.

参考文献

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1997

9.251

0.0

... 最早在矮牵牛(Pharbifis nil) NAM、拟南芥(Arabidopsis thaliana) ATAF1/2和CUC2的N端发现一段高度保守的氨基酸序列, 命名为NAC结构域, 并将包含NAC结构域的蛋白称为NAC转录因子^[1] ...

... 水稻中已分离的受干旱胁迫诱导表达的NAC转录因子基因有OsNAC6^[10]、ONAC045^[11]和OsNAC52^[12]和SNAC1^[13], Hu等^[13]研究表明, 过量表达SNAC1基因促进转基因水稻叶片气孔关闭, 减少水分丧失速率, 增强抗旱能力, 结实率提高22%~34% ...

... 在花生中已分离的NAC基因包括AhNAC1^[15]、AhNAC2和AhNAC3^[16]以及AhNAC4 (HM776131 ...

2003

4.425

0.0

... NAC转录因子是植物特有的转录调控因子, 是最大的转录因子家族之一^[2] ...

... NAC转录因子显著结构特点是, N端含有约150个氨基酸组成的高度保守的NAC结构域, C端有高度多样性的转录激活区^[2,3] ...

... AhNAC1基因属于ATAF亚族, 与大豆GmNAC2^[17]氨基酸序列一致性为78%, 过量表达GmNAC2的转基因烟草中活性氧清除系统基因的表达受抑制, 降低 ...

2002

3.518

0.0

Duval

, Hsieh T

, Kim S

, Thomas T

Molecular characterization of AtNAM: a member of the Arabidopsis NAC domain superfamily.

Plant Mol Biol, 2002, 50: 237-248

The petunia NAM and Arabidopsis ATAF1 and CUC2 genes define the conserved NAC domain. In petunia, loss-of-function nam mutants result in embryos that fail to elaborate shoot apical meristems (SAM), and nam seedlings do not develop shoots and leaves. We have isolated a NAC domain gene, AtNAM , from an Arabidopsis developing seed cDNA library. Expression of AtNAM mRNA is restricted primarily to the region of the embryo including the SAM. The AtNAM gene contains three exons and is located on Chromosome 1. In vivo assays in yeast demonstrate that AtNAM encodes a transcription factor and that the NAC domain includes a specific DNA binding domain (DBD). The AtNAM DBD is contained within a 60 amino acid region which potentially folds into a helix-turn-helix motif that specifically binds to the CaMV 35S promoter. The putative transcriptional activation domain is located in the C-terminal region of the protein, a highly divergent region among NAC domain-containing genes. The Arabidopsis genome contains 90 predicted NAC domain genes; we refer to these collectively as the AtNAC superfamily. The first two exons of all members of this superfamily encode the NAC domain. Most AtNAC genes contain three exons with the last exon encoding an activation domain. A subfamily of AtNAC genes contains additional terminal exons coding for protein domains whose functions are unknown.

1.Texas A&M University 2.Kumho Life and Environmental Science Laboratory

... NAC转录因子显著结构特点是, N端含有约150个氨基酸组成的高度保守的NAC结构域, C端有高度多样性的转录激活区^[2,3] ...

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

... 在花生中已分离的NAC基因包括AhNAC1^[15]、AhNAC2和AhNAC3^[16]以及AhNAC4 (HM776131 ...

2005

0.0

Kusano

, Asano

, Shimada

, Kadowaki

Molecular characterization of ONAC300, a novel NAC gene specifically expressed at early stages in various developing tissues of rice.

Mol Genet & Genomics, 2005, 272: 616-626

Members of the plant-specific gene family referred to as the NAC family (for NAM-ATAF-CUC-related) are involved in various functions including the regulation of plant development. However, no detailed molecular characterization of any member of the NAC family has yet been reported from monocots. Here, we report such a characterization of ONAC300 , a novel NAC-family gene identified using a cDNA cloned from microdissected phloem cells of rice. The predicted ONAC300 protein sequence falls into the NAM subgroup, which also contains the proteins CUC1 and CUC2 from Arabidopsis, CUP from snapdragon, CmNACP from pumpkin and NAM from petunia. High levels of ONAC300 mRNA were detected by in situ hybridization in developing shoot apical meristem (SAM) and in the associated young leaves. The use of an ONAC300 :: GUS reporter gene revealed that the ONAC300 promoter was expressed predominantly in developing vascular tissues of the leaves and roots. The construct was also expressed in anther filaments, rachis and carpel styles. RT-PCR analysis further revealed that the levels of ONAC300 transcripts were higher in leaves, roots and culms than in panicles. The observed expression pattern of ONAC300 is quite different from those of the dicot NAC genes previously reported. Thus, ONAC300 is a novel member of the NAC family which is expressed at very early developmental stages in the shoot, root and flower, as well as in the mature phloem of vascular tissues in rice.

1.Genetic Diversity Department National Institute of Agrobiological Sciences Kannondai 2-1-2, Tsukuba Ibaraki 305-8602 Japan 2.Department of Biological Science and Technology Tokyo University of Science Yamazaki 2641, Noda Chiba 278-8510 Japan 3.Division of Plant Biotechnology, Tissue Engineering Research Center Tokyo University of Science Yamazaki 2641, Noda Chiba 278-8510 Japan

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

2006

9.251

0.0

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

2006

0.0

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

2012

0.0

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

2012

0.0

孙利军, 李大勇, 张慧娟, 宋凤鸣. NAC转录因子在植物抗病和抗非生物胁迫反应中的作用. 遗传. 2012, 34: 993-1002

Sun L

, Li D

, Zhang H

, Song F

Functions of NAC transcription factors in biotic and abiotic stress responses in plants.

Hereditas (Beijing), 2012, 34: 993-1002 (in Chinese with English abstract)

NAC transcription factors belong to a unique class of transcription factors in plants. The common characteristics of the NAC proteins are the presence of a conserved NAC domain, comprising of about 150 amino acids in N-terminals and a highly variable transcriptional regulation region in C-terminals. Extensive studies have revealed that NAC transcription factors not only play important roles in plant growth and development, but also have functions in regulation of responses to biotic and abiotic stresses. In this minireview, we summarized the functions and mechanisms of the NAC transcriptional factors in plant abiotic and biotic stress responses. We also discussed future directions towards understanding the biological functions of the members of the NAC transcriptional factors in plants.

State Key Laboratory for Rice Biology, Institute of Biotechnology, Zhejiang University, Hangzhou 310029, China

NAC转录因子是植物特有的一类转录因子, 其共同特点是在N端含有一段高度保守、由约150个氨基酸组成的NAC结构域, 而C端为高度变异的转录调控区。研究表明, NAC转录因子不仅参与植物生长发育的调控, 而且在植物抗逆反应中具有重要的调控作用。文章着重介绍NAC转录因子在植物抗逆反应中的作用及其调控机制, 并简要讨论NAC转录因子生物学功能的研究方向。

... NAC转录因子不仅参与植物器官发育^[3,4]、次生壁的形成^[5]、细胞周期的调控^[6], 而且在植物抗逆境胁迫中具有重要的调控作用^[7,8] ...

2004

9.251

0.0

... Tran等^[9]从拟南芥中分离到3个受干旱、高盐和ABA诱导表达的NAC基因(AtNAC019、AtNAC055和AtNAC072), 在3个基因的转基因植株中皆超量表达, 并显著增强拟南芥抗旱性 ...

2005

1.132

0.0

2009

0.0

2010

9.251

0.0

Gao

, Xiong A

, Peng R

, Jin X

, Xu

, Zhu

, Chen J

, Yao

QH.

OsNAC52, a rice NAC transcription factor, potentially responds to ABA and confers drought tolerance in transgenic plants.

Plant Cell, 2010, 100: 255-262

A novel NAC ( N AM, A TAF1/2, C UC2) homologous gene, OsNAC52 , was isolated from Oryza sativa L using RACE (rapid amplification of cDNA ends). Analysis of the amino acid sequence of OsNAC52 revealed a highly conserved NAC domain and a potential nuclear localization sequence in its N-terminus, and the transcriptional activation motif in the C-terminal region. Transgenic plants over-expressing OsNAC52 were highly sensitive to ABA (Abscisic acid), and the growth of the 35S-OsNAC52 transgenic seedlings was significantly more restrained by ABA treatment than those of the wild-type seedlings. Furthermore, over-expression of OsNAC52 activated the expression of downstream genes in transgenic Arabidopsis, resulting in enhanced tolerance to drought stresses but not growth retardation. The results from this study indicate that this novel rice OsNAC52 gene functions as an important transcriptional activator in ABA-inducible gene expression and may be useful in improving plant tolerance to abiotic stress.

1.Shanghai Academy of Agricultural Sciences Biotechnology Research Institute 2901 Beidi Road 201106 Shanghai The People’s Republic of China 2.Yangzhou University College of Bioscience and Biotechnology 88 S. Daxue Road 225009 Yangzhou China

2006

9.737

0.0

2010

0.0

刘美英, 冶晓芳, 唐益苗, 高世庆, 张朝, 赵昌平, 陈学平. TaNAC提高了转基因烟草的抗旱功能. 中国烟草学报, 2010, 16(6): 82-88

Liu M

, Ye X

, Tang Y

, Gao S

, Zhang

, Zhao C

, Chen X

Effect of TaNAC on drought resistance in transgenic tobaccos.

Acta Tab Sin, 2010, 16(6): 82-88 (in Chinese with English abstract)

NAC(NAM-ATAF-CUC)转录因子在植物逆境胁迫应答反应中发挥重要作用.通过同源克隆的方法从小麦中分离得到一个编码NAC转录因子的基因-TaNAC,该基因编码的蛋白序列结构特殊,N端为保守性差的NAC保守域,只包含A,B,D 3个亚结构域,进化关系上与DREB家族更近,其C端共同享有多个基序.实时荧光定量PCR实验证实TaNAC基因受干旱诱导24 h后强烈表达.将该基因构建到PBI121双元植物表达载体的35S启动子下游,通过农杆菌介导的叶盘转化法转入烟草,获得T1代转基因苗.模拟干旱胁迫处理实验表明,过量表达TaNAC的转基因烟草表现出明显优于野生型对照的抗旱功能,说明TaNAC基因的过量表达能够提高烟草抗旱能力,TaNAC可以作为烟草抗旱育种的优良候选基因资源.

... 刘美英等^[14]分离得到小麦NAC转录因子的基因TaNAC, 干旱诱导24 h, TaNAC基因转录水平表达量显著升高, 过量表达TaNAC的转基因烟草在模拟干旱胁迫处理下, 抗旱性明显高于非转基因烟草 ...

2008

0.0

1.321

邵凤霞, 柳展基, 魏丽奇, 曹敏, 毕玉平. 花生NAC类新基因AhNAC1的克隆及序列分析. 西北植物学报, 2008, 28: 1929-1934

Shao F

, Liu Z

, Wei L

, Cao

, Bi Y

Cloning and sequence analysis of a novel NAC-like gene AhNAC1 in peanut (Arachis hypogaea).

Acta Bot Boreali-Occident Sin, 2008, 28: 1929-1934 (in Chinese with English abstract)

NAC转录因子是近些年来新发现的植物特有的转录调控因子,其N端含有高度保守的NAC结构域,在植物的生长发育、器官建成、逆境胁迫以及作物的品质改良中具有重要作用.通过构建优质出口型大花生'鲁花14'开花后20～60 d不同发育时期的种子混合cDNA文库,克隆到一个花生NAC类基因AhNACl(GenBank登录号为EU669863).序列分析表明,AhNAC1基因的cDNA序列长度为1 453 bp,其开放阅读框长度为912 bp,编码303个氨基酸,预测其分子量为34.1 kD,等电点为7.6.通过与其它植物的NAC转录因子的蛋白序列比对,发现AhNAC1蛋白属于ATAF亚族,可能在植物种子发育及逆境反应中起作用.此花生NAC类基因属首次报道.

... 在花生中已分离的NAC基因包括AhNAC1^[15]、AhNAC2和AhNAC3^[16]以及AhNAC4 (HM776131 ...

2009

0.0

1.667

刘旭, 李玲. 花生NAC转录因子AhNAC2和AhNAC3的克隆及转录特征. 作物学报, 2009, 35: 541-545

Liu

, Li

Cloning and characterization of the NAC-like gene AhNAC2 and AhNAC3 in peanut.

Acta Agron Sin, 2009, 35: 541-545 (in Chinese with English abstract)

NAC transcription factors existed differentially in plant are the new transcription regulatory factors with multiple biological functions. Two NAC-like genes from peanut were cloned by RT-PCR and RACE methods, named AhNAC2 and AhNAC3 (GenBank accession No. EU755023 and EU755022), which contain an ORF of 1 050 bp and 1 008 bp and encode 349 and 335 amino acids, respectively. Gene sequence analysis showed that the putative protein of both two genes contained conserved NAC domain and highly different C terminal which is the typical characteristics of NAC transcription factors. Meanwhile, the transcription levels of the two genes were investigated by semi-quantitative RT-PCR, and the result showed that the expressions of AhNAC2 and AhNAC3 genes were enhanced by ABA, GA 3 , water stress, and cold stress respectively. Furthermore, the two genes expressed constitutively in peanut tissues, and their expression patterns were different in various tissues. In conclusion, AhNAC2 and AhNAC3 genes isolated from peanut were new members of NAC transcription factor family, their comparison to RD26 (AT4G27410) reveals a high amino acid homology. We suggested that they play key roles in ABA signal transduction and drought response in peanut.

Guangdong Key Laboratory of Biotechnology for Plant Development, College of Life Science, South China Normal University, Guangzhou 510631

NAC 转录因子是特异存在于植物中具有多种生物功能的新型转录因子。本实验从花生中克隆了 2 个 NAC 基因 AhNAC2 和 AhNAC3 (GenBank 登录号 EU755023 和 EU755022) ；蛋白序列分析表明，两者具有典型 NAC 转录因子特征， N 端含有保守 NAC 结构域。半定量 RT-PCR 显示，在 ABA 和 GA 3 处理，以及控水和低温条件下， AhNAC2 和 AhNAC3 在花生组织中的表达上调，呈现不同的表达模式。 AhNAC2 和 AhNAC3 为典型的 NAC 转录因子新基因，蛋白序列与拟南芥 RD26 同源性较高，推测与响应干旱和 ABA 信号有关。

... 在花生中已分离的NAC基因包括AhNAC1^[15]、AhNAC2和AhNAC3^[16]以及AhNAC4 (HM776131 ...

2013

5.319

0.0

Jin H

, Huang

, Cheng

, Song H

, Yu D

Overexpression of the GmNAC2 Gene, an NAC transcription factor, reduces abiotic stress tolerance in tobacco.

Plant Mol Biol Rep, 2013, 31: 435-442

Abstract NAC (NAM, ATAF1, 2, and CUC2) proteins play important roles in plant development and stress responses. Glycine max NAC-like gene 2 (GmNAC2) is the first ATAF1-like NAC transcription factor identified in soybean. In this study, GmNAC-2-overexpressing tobacco lines were developed and found to be hypersensitive to drought, high salinity, and cold stress. Under abiotic stress, the transgenic tobacco leaves had higher malondialdehyde (MDA) levels compared with wild-type plants. An elevated MDA level is used frequently as an indicator of reactive oxygen species (ROS) and associated cell membrane degradation or dysfunction. To determine the mechanism of GmNAC2 -mediated stress intolerance, leaf transcriptome analyses were performed comparing GmNAC2 -overexpressing tobacco with wild-type tobacco. Identified downregulated genes included genes related to ROS scavenging in GmNAC2 -overexpressing tobacco. These results indicate that GmNAC2 functions as a negative regulator during abiotic stress, and participates in ROS signaling pathways through modulation of the expression of genes related to ROS-scavenging.

1. National Center for Soybean Improvement, National Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing, 210095, China

... AhNAC1基因属于ATAF亚族, 与大豆GmNAC2^[17]氨基酸序列一致性为78%, 过量表达GmNAC2的转基因烟草中活性氧清除系统基因的表达受抑制, 降低 ...

2010

0.622

0.0

... AhNAC2属于AtNAC3亚族, 可被ABA高效诱导表达, 通过调节气孔大小来抵御干旱胁迫^[18] ...

2013

0.0

... AhNAC3不能明显地被ABA诱导, 可以通过增强SOD活性减少活性氧的积累来抵御干旱胁迫, 推测其调控一种特殊的路径^[19] ...

2014

0.0

1.667

厉广辉, 张昆, 刘风珍, 万勇善. 不同抗旱性花生品种根系形态及生理特性. 作物学报, 40: 531-541

Li G

, Zhang

, Liu F

, Wan Y

Morphological and physiological traits of root in different drought resistant peanut cultivars. Acta Agron Sin, 2014 40: 531-541 (in Chinese with English abstract)

Drought stress is a serious constraint for peanut production worldwide. It is necessary to identify the drought resistance mechanisms of different peanut cultivars in drought-resistance breeding . Under the artificial water control condition, the peanut root morphological and physiological characteristics under drought stress at seedling stage and pod-setting stage were studied using 12 different drought-resistance peanut cultivars as material. The results showed that drought resistance at seedling stage was basically identical with that at pod-setting stage. According to yield-drought resistance coefficient, 12 peanut cultivars were divided into three grades: high-resistance, including A596, Shanhua 11, and Rugaoxiyangsheng; mid-resistance, including Huayu 20, Nongda 818, Haihua 1, Shanhua 9, and 79266; and weak-resistance, including ICG6848, Baisha 1016, Hua 17, and Penglaiyiwohou. In those peanut cultivars, Shanhua11 can be used as the standard cultivar for high drought resistance identification, and 79266 as the standard cultivar for weak one. The root drought resistance mechanism of 12 peanut cultivars were different, Shanhua 9, Shanhua11 and Huayu 20 presented a larger biomass and strong absorption capacity, while A596, Nongda 818, Shanhua 11 and Rugaoxiyangsheng had strong antioxidant capacity and membrane stability under drought stress. Correlations between drought resistance and root weight, volume, total absorption area per plant, root superoxide dismutase (SOD) activity, malondialdehyde (MDA) content under serious drought stress were significant, also under control condition. Therefore, under drought stress of 40% RWC for 10 to 24 d after germination, the root weight, volume, total absorption area per plant, SOD activity and MDA content could be used for identifies the drought resistance ability of peanut roots, the resistances degree also can be reflected by the indices above under normal water condition. Shanhua 11 can be used as a suitable standard cultivar for root morphological and physiological drought resistance characteristics identification in peanut

National Key Laboratory of Crop Biology / College of Agronomy, Shandong Agricultural University, Tai’an 271018, China

以 12 个花生品种为试验材料 , 在人工控水条件下 , 通过苗期及结荚期干旱试验 , 对比分析花生品种苗期根系性状与抗旱性的关系。结果表明 , 花生苗期与结荚期抗旱性基本一致。利用产量抗旱系数可把 12 个花生品种的抗旱性划分为强、中、弱 3 级 , 抗旱性强的品种为 A596 、山花 11 和如皋西洋生 , 中度抗旱品种为花育 20 、农大 818 、海花 1 号、山花 9 号和 79266 , 抗旱性弱的品种有 ICG6848 、白沙 1016 、花 17 和蓬莱一窝猴。山花 11 可作为花生强抗旱性鉴定的标准品种 , 79266 可作为花生弱抗旱性鉴定的标准品种。山花 9 号、山花 11 、花育 20 的根系抗旱机制为较大的根量及根系吸收能力 , 而 A596 、如皋西洋生、农大 818 、山花 11 为较强的根系抗氧化能力及膜稳定性。相关分析表明 , 苗期重度干旱胁迫下的单株根系干重、体积、总吸收面积、超氧化物歧化酶 (SOD) 活性和丙二醛 (MDA) 含量与品种抗旱系数的相关性达极显著水平 , 对照与重度干旱胁迫下的以上性状呈极显著正相关。因此 , 在花生出苗后 10 d 进行 40% 土壤相对含水量的干旱胁迫 , 持续胁迫至出苗 24 d 的单株根系干重、体积、总吸收面积、根尖 SOD 活性和 MDA 含量可鉴定花生品种的根系抗旱能力 , 正常水分下的性状值也能反映根系性状的抗旱级别。山花 11 可作为花生根系形态及生理优异抗旱性状鉴定的标准品种。

... 栽培品种分为三类^[20,21,22], 抗旱性强的包括濮花28号、莱宾大豆、NC6、山花17号、花育31、鲁花8号、远杂9102、徐州68-4、山花12号、冀0212-4、山花13号、山花9号、鲁花14、阜花10号、豫花15、桂花1026、山花11号 ...

... 基因型为a2a2b2b2的品种只有2份, 山花13号为抗旱品种, 79266为中度抗旱品种, 且79266抗旱系数处于中度抗旱和弱抗旱性的临界值^[20,30] ...

2014

0.0

1.889

厉广辉, 张昆, 刘风珍, 刘丹丹, 万勇善. 不同抗旱性花生品种的叶片形态及生理特性. 中国农业科学, 2014, 47: 644-654

Li G

, Zhang

, Liu F

, Liu D

, Wan Y

Morphological and physiological traits of leaf in different drought resistant peanut cultivars.

Sci Agric Sin, 2014, 47: 644-654 (in Chinese with English abstract)

【Objective】The main purpose of this paper was to screen leaf traits related to drought resistance, explore the methods of traits index evaluation, and to reveal drought resistance mechanisms of different peanut cultivars. 【Method】 The drought resistance was studied in twelve peanut cultivars under drought stress and normal irrigation in a pot experiment at seedling stage and a pool culture experiment at pod setting stage. Keeping the water treatments by using weighing at seedling stage and supplemental irrigation based on testing soil moisture at pod setting stage with rain-shedding during drought stress. Leaf morphological and physiological traits including organization structure, thickness, specific leaf weight (SLW), leaf area per plant (PLA), photosynthetic rate (Pn), chlorophyll content under the conditions of normal water supply and drought stress at seedling stage were tested, and the relationship between those traits and drought resistance was studied. Drought resistances of cultivars were scored with drought coefficient of biomass at seedling stage and yield at pod setting stage. Drought resistance mechanisms of leaf were evaluated by traits index. 【Result】 The results by two years experiments showed that, under drought stress, the drought resistance coefficients of different peanut cultivars were significantly different. Drought resistance at seedling and pod-setting stages was basically identical. According to yield-drought resistance coefficient, twelve peanut cultivars could be divided into 3 grads: high-resistance, including A596, Shanhua 11 and Rugaoxiyangsheng; mid-resistance, including Huayu 20, Nongda 818, Haihua 1, Shanhua 9 and 79266; weak-resistance, including ICG6848, Baisha1016, Hua17 and Penglaiyiwohou. Water stress changed organization structure in leaves, functional leaf area, PLA, Gs, Pn and Tr were reduced, but SLW of peanuts was increased under soil drought stress. A significant difference in leaf traits among peanut cultivars with different drought resistances was observed. Leaf thickness, ratio of palisade tissue to spongy tissue (PTT/STT), SLW, PLA and Pn were higher under both drought stress and normal irrigation in cultivars with high drought resistance. The drought resistance mechanism of twelve peanut cultivars was different, under drought stress, Rugaoxiyangsheng and Shanhua 11 presented as higher PTT/STT, SLW and Pn, Shanhua 9 and Huayu 20 had larger PLA, A596 was mainly higher in Pn. Correlation analysis between drought resistance coefficient and leaf PTT/STT, SLW, PLA and Pn under drought stress condition was significant. 【Conclusion】 Shanhua11 could be used as a standard variety for high drought resistance identification, and 79266 could be used as a standard variety for weak resistance identification. Treatment of 40% RWC drought stress at 10 d after germination, the leaf PTT/STT, SLW, PLA and Pn could be used to identify the drought resistance of peanuts leaf. PTT/STT, SLW and PLA also could be used to identify the drought resistance of peanuts leaf under normal water condition at anthesis stage. Shanhua 11 could be used as a suitable standard cultivar for leaf drought resistance traits identification in peanut.

College of Agronomy, Shandong Agricultural University/National Key Laboratory of Crop Biology, Tai’an 271018, Shandong

【目的】筛选叶片抗旱相关性状，探索性状指标大小的评价方法，揭示不同抗旱性花生品种抗旱的叶片形态及生理机制。【方法】以12个花生品种为试验材料，在人工控水条件下，于苗期及结荚期给予干旱胁迫及正常灌水对照2个土壤水分处理，苗期处理以盆栽种植方式进行，称重法控水，结荚期处理在池栽条件下进行，测墒补灌法控水，控水期间以电动防雨棚遮雨。测定苗期干旱及正常供水对照的叶片组织结构、厚度、比叶重、叶面积、光合速率、叶绿素含量等叶片性状指标，对比分析上述叶片性状与花生品种抗旱性的关系，以性状指标大小评价不同花生品种抗旱的叶片机制，以生物量抗旱系数评价苗期抗旱性，以收获期产量抗旱系数评价结荚期抗旱性。【结果】通过2年的试验发现，干旱胁迫下，不同花生品种生物量抗旱系数及产量抗旱系数有显著差异，苗期与结荚期抗旱性基本一致。根据产量抗旱系数可将12个花生品种抗旱性分为强、中、弱3级，抗旱性强的品种为A596、山花11号、如皋西洋生，花育20号、农大818、海花1号、山花9号和79266为中度抗旱品种，抗旱性弱的品种有ICG6848、白沙1016、花17和蓬莱一窝猴。干旱胁迫改变了叶片组织结构，降低了单株叶面积、功能叶面积、气孔导度、光合速率和蒸腾速率，增加了比叶重。不同抗旱性花生品种叶片性状有显著差异，抗旱性强的品种在对照及干旱胁迫下均具有较高的叶片厚度、栅/海比、比叶重、单株叶面积、光合速率。干旱胁迫下，较大的叶片栅/海比值、比叶重和光合速率是如皋西洋生和山花11号抗旱的叶片机制；山花9号与花育20号较大的单株光合面积，A596较大的光合速率是它们重要的抗旱机制。相关分析表明，干旱胁迫下的叶片栅/海比、比叶重、单株叶面积和光合速率与品种抗旱系数的相关性达极显著水平。【结论】山花11号和79266分别可作为花生强、弱抗旱性鉴定的标准品种。花生出苗后10 d进行40%土壤相对含水量的干旱处理，持续胁迫14 d时的叶片栅/海比、单株叶面积、比叶重和光合速率可鉴定花生种质的叶片抗旱能力。正常灌水条件下，始花期的栅/海比、单株叶面积、比叶重也能反映花生叶片的抗旱级别。山花11号可作为花生叶片优异抗旱性状鉴定的标准品种。

2013

0.0

2013

0.657

0.0

... )是异源四倍体(AABB), 一般认为栽培种是由2个具有不同染色体组的二倍体野生种自然杂交的杂种, 再经一次性加倍事件进化形成^[23] ...

1995

0.0

1.889

杨建昌, 王志琴, 朱庆森. 水稻品种的抗旱性及其生理特性的研究. 中国农业科学, 1995, 28(5): 65-72

Yang J

, Wang Z

, Zhu Q

Drought resistance and its physiological characteristics in rice varieties.

Sci Agric Sin, 1995, 28(5): 65-72 (in Chinese)

江苏农学院，扬州 225009

研究了10个中熟水、陆稻品种（含杂交组合）的抗旱性以及对水分胁迫的生理反应。结果表明：不同品种的抗旱系数为：旱-A03＞旱-501＞珍珠矮＞扬稻4号＞汕优63＞六优1号＞亚优2号＞南京11＞盐粳2号＞盐粳235，而抗旱指数和在水分胁迫下的产量则以3个杂交组合汕优63、亚优2号和六优1号最高；在水分胁迫下，抗旱性较好（抗旱系数较高）的品种SOD（超氧化物歧化酶）活性明显增加的叶水势阈值较高，叶片质膜透性明显增加的叶水势阈值较低，抗旱性较差（抗旱系数较低）的品种则相反；水分胁迫下库源比、茎鞘物质的输出率和转换率的相对比率（胁迫处理/对照×100%），抗旱性较好的品种较高，抗旱性较差的品种较低；在水分胁迫下剑叶的气孔密度增加，但抗旱性较好的品种增加较少，抗旱性较差的品种增加较多。

... 2012年进行了抗旱性鉴定, 抗旱等级的划分依据品种的抗旱系数(drought resistance coefficient, DC), 抗旱系数的计算公式为DC=Y_d/Y_p, 式中Y_d为干旱胁迫下产量或生物量, Y_p为非胁迫下产量或生物量, 抗旱系数是整株形态及生理生化性状对干旱综合反应的结果, 是对品种综合抗性的评价^[24,25,26] ...

2008

2.151

0.0

2006

2.474

0.0

1999

2.474

0.0

... 作物的抗旱性是多基因控制的数量性状, 受作物的形态特征、组织器官结构、生理生化性状及抗旱相关基因的转录与蛋白质翻译表达水平等诸多因素的控制, 不同品种的抗旱机制存在差异^[27,28,29] ...

2012

0.0

1.667

张智猛, 戴良香, 丁红, 陈殿绪, 杨伟强, 宋文武, 万书波. 中国北方主栽花生品种抗旱性鉴定与评价. 作物学报, 2012, 38: 495-504

Zhang Z

, Dai L

, Ding

, Chen D

, Yang W

, Song W

, Wan S

Identification and evaluation of drought resistance in different peanut varieties widely grown in northern China.

Acta Agron Sin, 2012, 38: 495-504 (in Chinese with English abstract)

It is necessary to systematically identify and evaluate drought-resistant varieties for stable development of peanut production in arid and semi-arid regions in China. To establish drought resistance index of peanut cultivars, we determined 13 morphological and biochemical indices including plant height, number of branch, biomass, leaf water content and photosynthetic pigment content in the pot experiment at pod setting and pod filling stages with 29 peanut cultivars (strains) widely grown in north China under artificial water stress condition. Drought resistances of varieties were scored with drought coefficient method and subordinate function value under water stresses. The results showed that the same index of drought resistance characterized in different peanut varieties achieved different results, and 29 peanut varieties were divided into four grads: high tolerant, moderate tolerant, weaker tolerant, and drought susceptible. In the 29 cultivars (strains), we screened out the high drought tolerant varieties consisting of Tangke 8, Jihua 2, Jihua 4, Huayu 25, Huayu 17, Huayu 22, Datangyou, and Huayu 21. The cultivars (strains) of Huayu 20, Huayu 24, Weihua 6, Huayu 27, Luhua 11, Fuhua 11, Fuhua 13, Tangyou 4, Fenghua 1, and G2 had moderate drought resistance. The cultivars (strains) with weaker drought resistance were 16-8, Fuhua 10, M5, Huayu 16, and Luhua 14. At pod setting and pod filling stages Huayu 19, Huayu 23, Fenghua 6, Weihua 8, M7, and TE-2 were all drought susceptible. From seedling to pod setting stages under water stress, all morphological and biochemical indices but the number of branch and carotenoid content and their comprehensive D values could be taken as evaluation criteria. From seedling to pod filling stages under water stress, there only existed significanct correlation between leaf area and drought resistance. Either at pod setting or pod filling stages under water stress, comprehensive D value of the indices had significant correlation with drought resistance, which could be used to identify peanuts drought resistance.

1 Peanut Research Institute of Shandong Province, Qingdao 266100, China; 2 Shandong Academy of Agricultural Sciences, Ji’nan, 250100, China

人工控水条件下，通过盆栽试验，在花生植株的结荚期和饱果期, 测定29个品种(系)的株高、分枝数、生物累积量、叶片含水量和光合色素含量等与抗旱性有关的13个表型性状和生理生化指标，采用抗旱系数法和隶属函数值法，鉴定各性状指标水分胁迫下的抗旱性。结果表明,相同指标评价不同花生品种抗旱性时得出的结果不同,依此可将供试品种(系)划分为抗旱性强、中、弱和不抗旱4类,即供试品种中唐科8号、冀花2号、冀花4号、花育25、花育17、花育22、大唐油、花育21具有较强的抗旱能力；花育20、花育24、潍花6号、花育27、鲁花11、阜花11、阜花13、唐油4号、丰花1号和G2具有中等抗旱能力；具有较弱抗旱能力的品种(系)有16-8、阜花10号、M5、花育16和鲁花14；而花育19、花育23、丰花6号、潍花8号、M7和TE-2在结荚期和饱果期均无抗旱能力。水分胁迫至结荚期，除分枝数和类胡萝卜素外，其余各形态型状和生理生化指标及其综合D值均可作为鉴定品种(系)抗旱性的依据；水分胁迫至饱果期，只有叶面积与抗旱系数间的相关极显著。无论在结荚期或饱果期，综合D值可作为鉴定品种抗旱性的指标，单一指标均不能准确判定某品种(系)的抗旱性。

2011

0.0

1.989

张智猛, 万书波, 戴良香, 宋文武, 陈静, 石运庆. 花生抗旱性鉴定指标的筛选与评价. 植物生态学报, 2011, 35: 100-109

Zhang Z

, Wan S

, Dai L

, Song W

, Chen

, Shi Y

Estimating and screening of drought resistance indexes of peanut.

Chin J Plant Ecol, 2011, 35: 100-109 (in Chinese with English abstract)

Aims Our objective was to define an index of peanut ( Arachis hypogaea ) drought resistance and comprehensively evaluate drought resistance of peanut varieties under controlled water conditions. Methods The experiment was conducted on the farm of Qingdao Academy of Agricultural Science with 29 peanut varieties. Two levels of water content (80%–85% and 45%–50% of field moisture capacity) were used in a randomized complete block design with four replications. We measured 13 morphological and physiological characters such as plant height, branch numbers, biomass, leaf water content and photosynthetic pigment content in seedling and pod-pin stages. Important findings The 29 peanut cultivars can divided into four classes: high-, mid- and low-tolerant cultivars and intolerant cultivars. The high-tolerant cultivars included ‘Tangke 8’, ‘Jihua 2’, ‘Datangyou’, ‘Huayu 25’, ‘Huayu 17’, ‘Luhua 14’ and ‘Fenghua 1’. Subordinate function values of the morphological indexes including main stem height, branch number and biomass, and photosynthetic pigments contents of the same cultivars at seedling stage were different. In addition, the subordinate function value of the seedling stage was unrelated to drought resistance, so a single physiological index cannot be used to identify the drought resistance of peanut. But the values drought resistance comprehensive evaluation ( D ) of physiological indexes were significantly related to drought resistance and can be used to identify the drought resistance of peanut. At the pod-pin stage, the D values of morphological indexes and physiological indexes were both significantly related to drought resistance, so the subordinate function value and D value of morphological indexes and physiological indexes at this stage can serve as indexes for identifying peanut drought resistance.

1 Peanut Research Institute of Shandong Province, Qingdao 266100, China; 2 Shandong Academy of Agricultural Sciences, Jinan 250100, China

为确定鉴定花生( Arachis hypogaea )品种(系)抗旱性指标体系, 综合评价花生品种(系)的抗旱性, 在人工控水条件下, 通过盆栽试验, 测定了29个花生品种(系)苗期和花针期的株高、分枝数、生物累积量、叶片含水量和光合色素含量等与抗旱性有关的13个表观形态性状和生理性状的指标, 采用抗旱系数法和隶属函数值法, 对各指标性状进行了水分胁迫下的抗性评价和鉴定。结果表明, 29个花生品种(系)可划分为抗旱性较强、中等、较弱和不抗旱4类, 其中‘唐科8号’、‘冀花2号’、‘大唐油’、‘花育25号’、‘花育17号’、‘鲁花14号’、‘丰花1号’ 7个品种(系)具有较强的抗旱能力; 苗期同一品种(系)的主茎高、分枝数和生物累积量等形态指标和光合色素等生理指标的隶属函数值均有较大差别, 苗期各指标隶属函数值与品种(系)抗旱性无显著相关关系, 苗期单一形态指标不能作为鉴定品种(系)抗旱性的指标; 但苗期抗旱性综合评价值( D )与抗旱系数间存在显著相关关系, D 的大小可作为抗旱性的鉴定指标。花针期形态指标和生理指标 D 值间, 以及各类指标 D 值与抗旱系数间均存在显著或极显著的相关关系, 此期植株形态指标、生理指标隶属函数值以及综合 D 值均可作为鉴定品种(系)抗旱性的指标。

2004

0.0

0.7753

严美玲, 李向东, 矫岩林, 王丽丽. 不同花生品种的抗旱性比较鉴定. 花生学报, 2004, 33(1): 8-12

Yan M

, Li X

, Jiao Y

, Wang L

Identification of drought resistance in different peanut varieties.

J Peanut Sci, 2004, 33(1): 8-12(in Chinese)

通过研究花生种子在盛花期与抗旱性密切相关的形态指标与生理指标,对9个不同的花生品种(系)进行了抗旱性鉴定,并用隶属函数法对花生品种的抗旱性进行了综合评价.结果表明,同一花生品种在干旱与正常供水条件下相比较,其株高、单株叶面积、根冠比、比叶重、根茎叶干重及叶绿素含量差异显著,因此,它们可以作为鉴定花生抗旱性的简易指标.不同花生品种的抗旱性有明显差异,并鉴定出高抗旱品种丰花1号,抗旱品种海花1号、农大142、鲁花14号,不抗旱品种农大818、鲁花5号与79266.

... 基因型为a2a2b2b2的品种只有2份, 山花13号为抗旱品种, 79266为中度抗旱品种, 且79266抗旱系数处于中度抗旱和弱抗旱性的临界值^[20,30] ...

2009

1.668

0.0

... 花生野生种对环境具有广泛的适应性, 在各种环境胁迫条件下, 比栽培品种具有更强的环境适应和逆境耐受能力, 并且在形态学和分子水平都有丰富的变异, 具有抗旱, 抗盐、抗病虫害等多种抗逆性状, 是栽培种遗传改良极其宝贵的基因资源^[31,32] ...

2012

0.0

0.6276

2008

0.0

0.951

陈本银, 姜慧芳, 任小平, 廖伯寿, 黄家权. 野生花生抗青枯病种质的发掘及分子鉴定. 华北农学报, 2008, 23(3): 170-175

Chen B

, Jiang H

, Ren X

, Liao B

, Huang J

Identification and molecular traits of Arachis species with resistance to bacterial wilt.

Acta Agric Boreali-Sin, 2008, 23(3): 170-175 (in Chinese with English abstract)

以花生属5个区组的79份野生花生种质为材料,系统鉴定了野生花生对青枯病的抗性反应,从中发掘高抗青枯病的种质15份,含匍匐区组种质3份、直立区组1份、异形花区组1份、花生区组8份、未命名种质2份,抗病材料频率达到19%,高于栽培种花生资源的抗性频率.通过SSR分析表明,在所获得的抗青枯病野生花生材料中,四倍体野生种A.monticola与栽培种花生的亲缘关系最近,其次为花生区组的二倍体野生种A.duranensis和A.chacoense.根据DNA扩增结果,绘制了抗青枯病种质的指纹图谱,明确了其SSR分子特性.

... 陈本银等^[33]从花生野生种中发掘了15份高抗青枯病的种质, 抗病材料频率达到19% ...