水稻早衰突变体<italic>esl5</italic>的鉴定及其基因精细定位

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桑贤春, 徐芳芳, 朱小燕, 邢亚迪, 何沛龙, 张长伟, 杨正林, 何光华. 水稻早衰突变体esl5的鉴定及其基因精细定位 . 作物学报, 2014, 40(7): 1182-1189
SANG Xian-Chun, XU Fang-Fang, ZHU Xiao-Yan, XING Ya-Di, HE Pei-Long, ZHANG Chang-Wei, YANG Zheng-Lin, HE Guang-Hua. Identification and Gene Fine Mapping of Early Senescent Leaf Mutantesl5 inOryza sativa . Acta Agronomica Sinica, 2014, 40(7): 1182-1189 复制到剪切板

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水稻早衰突变体esl5的鉴定及其基因精细定位

桑贤春, 徐芳芳, 朱小燕, 邢亚迪, 何沛龙, 张长伟, 杨正林, 何光华^*

西南大学水稻研究所 / 转基因植物与安全控制重庆市重点实验室 / 南方山地农业教育部工程研究中心, 重庆400716

^* 通讯作者(Corresponding author): 何光华, E-mail:hegh@swu.edu.cn

第一作者联系方式: E-mail:sangxianchun@163.com

收稿日期:2014-04-16

基金:本研究由中央高校基本科研业务费创新团队项目(XDJK2013A023)和国家自然科学基金项目(31071072;31371597)资助;

摘要

叶片早衰直接影响作物产量和品质, 鉴定早衰突变体、图位克隆调控基因对于研究植物衰老机制具有重要的意义。以甲基磺酸乙酯诱变水稻籼型恢复系缙恢10号, 获得一个早衰突变体esl5(early senescent leaf mutant 5), 本文对其进行了形态鉴定、细胞学观察、理化分析和基因定位等研究。结果表明, 与野生型相比,esl5的苗期叶片正常, 分蘖期呈黄绿色, 孕穗期开始叶片中上部逐渐黄化衰老; 衰老部位的细胞结构异常, 细胞膜降解, 叶绿体基质片层疏松、排列不规则, 光合色素含量和光合速率极显著下降。此外,esl5的·OH和H₂O₂含量极显著升高, SOD和CAT的活性则极显著降低。与野生型相比,esl5的生育期延长20 d左右, 千粒重显著增加, 穗粒数、实粒数和结实率则显著降低。esl5突变性状受1对隐性核基因调控, 精细定位在第3染色体Indel标记Indel03-1和Indel03-2之间83.4 kb的物理范围内, 包含11个注释基因, 这为ESL5的克隆和功能研究奠定了基础, 也有利于水稻品种的遗传改良。

关键词: 水稻(Oryza sativa); 早衰; 基因定位; 活性氧(ROS); 光系统

Identification and Gene Fine Mapping of Early Senescent Leaf Mutantesl5 inOryza sativa

SANG Xian-Chun, XU Fang-Fang, ZHU Xiao-Yan, XING Ya-Di, HE Pei-Long, ZHANG Chang-Wei, YANG Zheng-Lin, HE Guang-Hua^*

Rice Research Institute of Southwest University / Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crops /Engineering Research Center of South Upland Agriculture, Ministry of Education, Chongqing 400716, China

Fund:

Abstract

Premature senility directly influences crop yield and quality in the production. Therefore, it is crucial to identify early senescent leaf mutants and then clone genes associated with senescence by map-based strategy, which is significant in the research of senescence mechanism. An early senescent leaf mutantesl5 has been discovered from the progeny ofindica restorer line Jinhui10 with seeds treated by ethylmethane sulfonate, this paper performed the studies on its morphological identification, cytological observation, physiological analysis and gene mapping. The results showed that theesl5 demonstrated normal phenotype at the seedling stage, displayed yellow green color at the tillering stage, and appeared yellow and senescent leaf blades at booting stage. Cell structure of theesl5 was badly influenced, showing destroyed cell membranes, loose and irregular stroma lamella in the section of senescent leaves. Chlorophyll content and photosynthetic rate of theesl5 were significantly lower than those of the wild type. Besides of the lower activities of SOD and CAT, the higher contents of ·OH and H₂O₂ were detected in theesl5 than in the wild type. The growth period ofesl5 delayed 20 days, grain number per panicle, filled-seed number and seed setting rate decreased significantly, but 1000-seed weight increased significantly at compared with the wild type. Genetic analysis suggested that the mutant trait was controlled by a recessive nuclear gene, and theESL5 was fine mapped on chromosome 3 between Indel markers Indel03-1 and Indel03-2 with 83.4 kb physical distance, containing 11 annotated genes. These results provide a foundation for the cloning ofESL5 by map-based strategy, which is essential in the research of senescence mechanism as well as molecular breeding inOryza sativa.

Keyword: Rice (Oryza sativa); Early senescence; Gene mapping; Reactive oxygen species (ROS); Photosynthetic systems

Show Figures

植物衰老是一个细胞程序性死亡的过程, 涉及叶绿体发育、叶绿素合成与降解、激素和自由基代谢等途径, 既受遗传基因的调控, 也受外界环境的影响^{[ 1]}。植物衰老的主要特征是叶片黄化、叶绿素降解和光合作用下降。自然衰老过程中, 营养物质从叶片转移至种子, 有利于植物的生长繁殖, 是其长期进化过程中形成的一种适应性^{[ 2]}。但在农业生产中, 叶片早衰则会导致作物产量和品质的下降, 如水稻穗中60%~70%的碳是由功能叶合成转运而来的^{[ 3]}。水稻不仅是单子叶模式植物, 也是重要的粮食作物, 克隆水稻早衰基因对探索植物衰老机制和水稻的遗传改良具有重要的意义。如 OsDOS编码一个水稻CCCH-Type锌指蛋白, 功能缺失促进叶片衰老, 过表达则延缓叶片衰老^{[ 4]}。

基因克隆的策略很多, 其中, 转录组和蛋白质组是植物衰老机制研究的常用手段^{[ 5]}。在双子叶模式植物拟南芥中, 利用表达谱芯片技术从衰老的叶片中鉴定到800个上调SAG (senescence-associated genes)^{[ 6]}, 在水稻剑叶自然衰老过程中, 则检测到533个上调SAG^{[ 7]}。全基因组分析获得了大量的SAG, 为从基因组角度阐释叶片衰老机制奠定了基础, 但是这些SAG是否直接调控植物衰老尚需进一步的功能验证。图位克隆是基因分离和功能研究中最直接有效的技术手段, 植物基因组测序的大量完成进一步促进了这一技术的发展, 其关键是优良突变体的获得^{[ 8]}。

目前, 在水稻中仅鉴定到十几个叶片自然早衰突变体, 其中9个基因已定位, 突变体 psl1、 pgl2、 psl3、 pse(t)、 es-t、 esl2、 esl3、 sms1和 lad的调控基因分别定位在第2、第8、第7、第7、第5、第4、第5、第8和第11染色体上^{[ 9, 10, 11, 12, 13, 14, 15, 16, 17]}; 4个基因已克隆, NOE1与 OsDM I3均与过氧化反应有关, NOE1编码过氧化氢酶OsCATC, 基因功能缺失通过激活硝酸还原酶产生大量NO, 从而诱导叶片早衰, noe1突变体内H₂O₂的含量显著高于野生型^{[ 18]}。 OsDMI3位于水稻第5染色体上, 编码Ca²⁺/CaM依赖型蛋白激酶, 在ABA诱导的抗氧化防护过程中具有重要的功能, OsDMI3功能缺陷可造成植株体内H₂O₂的过量积累, 导致叶片衰老^{[ 19]}。 RLS1编码一个C端具有ARM结构域的核苷酸结合蛋白, 参与程序性细胞死亡过程, 叶绿素的快速降解是导致 rls1衰老的主要原因^{[ 20]}; SPL28编码网格相关受体蛋白复合体亚基(AP1M1), 参与高尔基的物质运输途径, 其功能缺失引起植物过敏性反应, 形成叶片类病斑, 导致水稻叶片早衰^{[ 21]}。

EMS诱变籼型水稻恢复系缙恢10号的种子, 在其后代中发现了一个稳定遗传的淡叶早衰突变体, 暂命名为 esl5, 本文对其进行了形态鉴定、理化分析和基因精细定位等研究, 为下一步基因的克隆和功能分析奠定了基础, 也有利于水稻的遗传改良。

1 材料与方法

1.1 突变体材料

缙恢10号是西南大学水稻研究所采用花粉管介导法将高粱DNA导入水稻品系明恢63, 利用分子标记辅助选择获得的一个晚籼恢复系。以甲基磺酸乙酯(EMS)诱变缙恢10号的种子, 从其后代中鉴定到一个淡叶早衰突变体 esl5, 连续6代种植, 突变性状均稳定遗传。以粳稻品种日本晴为母本, esl5为父本, 配制杂交组合, 利用F₁和F₂群体进行遗传分析, 并利用F₂隐性群体进行基因定位。

1.2 测定项目与方法

1.2.1 形态鉴定田间种植 esl5及野生型, 株行距25 cm×30 cm, 3次重复, 全生育期调查植株表型, 成熟后取小区中间5株, 测定有效穗、穗粒数、穗实粒数、结实率、千粒重、籽粒长和籽粒宽等主要农艺性状。

1.2.2 光合色素含量和光合作用测定田间种植情况下, 参照文献^{[ 22]}描述的方法在分蘖期和抽穗期分别测定 esl5和缙恢10号的光合色素含量, 并于抽穗期利用LI-6400型便携式光合作用仪测定净光合速率、气孔导度、蒸腾速率和胞间CO₂浓度。

1.2.3 生理指标检测参照文献^{[ 14]}描述的方法测定超氧阴离子(O₂^-)含量; 利用南京建成生物工程研究所生产的试剂盒, 参照其说明书测定过氧化氢(H₂O₂)、·OH、丙二醛(MDA)、可溶性蛋白质(SP)和脯氨酸(PRO)含量以及超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)活性。

1.2.4 细胞超微结构观察抽穗期, 取 esl5和野生型叶片, 以戊二醛和锇酸双重固定, 不同梯度的乙醇逐级脱水, 置换和包埋, 制超薄切片, 经醋酸双氧铀和柠檬酸铅液双重染色后, 利用H600型透射电镜观察并照相。

1.3 DNA的提取

从日本晴/ esl5的F₂群体中分别取10株典型的正常单株和突变单株, 剪取等量叶片混合后构建正常池和突变池。采用改良的CTAB法提取亲本、正常池和突变池的基因组DNA, 利用碱煮法提取F₂定位单株的基因组DNA^{[ 23]}。

1.4 SSR分子标记分析

参照http://www.gramene.org/microsat, 由上海英骏生物公司合成SSR引物序列。PCR总体系12.5 μL, 包括1.25 μL的10×PCR buffer、0.75 μL的25 mmol L^-1 MgC1₂、0.5 μL的2.5 mmol L^-1 dNTPs、7.9 μL的ddH₂O、1.0 μL的10 μmol L^-1引物、1.0 μL的模板DNA和0.1 μL的5 U μL^-1 Taq DNA聚合酶。PCR程序为94℃预变性3 min; 94℃ 30 s, 56℃ 30 s、72℃ 1 min, 35个循环; 再72℃延伸10 min。PCR产物经10%非变性聚丙烯酰胺凝胶电泳, 0.1%硝酸银染色10 min, 去离子水漂洗2次, 1.5%氢氧化钠和0.2%甲醛显色后, 观察并照相。

1.5 精细定位

利用NTI vector 11.0软件对初步定位区间内93-11和日本晴的DNA序列进行比对, 在差异位点两侧利用Primer Premier5.0 软件设计引物, 开发新的Indel标记。引物的合成、PCR反应和电泳分析同上。

1.6 遗传图谱的构建

定位群体中, 具有 esl5带型的单株记为B, 具有日本晴带型的单株记为A, 具有日本晴/ esl5 F₁杂合带型的单株记为H。根据公式[( H+2 A)/2 n]×100计算遗传距离并构建连锁图谱, 其中 H表示定位群体中出现杂合体带型单株的数量, A表示出现正常株带

型的单株数, n表示用于定位的隐性群体总株数。

2 结果与分析

2.1 形态鉴定

野生型缙恢10号自然生理衰老前叶片一直保持绿色, esl 5的叶片在苗期呈绿色, 分蘖期逐渐发育成黄绿色, 孕穗期开始叶尖黄化衰老, 逐渐扩展至整个叶上部, 一直持续到成熟。 esl5的抽穗期与野生型相比延迟了14 d左右, 野生型处于开花状态时, esl5才刚进入抽穗期, 整个生育期延长了20 d左右(图1)。与野生型相比, 分蘖期 esl5的叶绿素a和b的含量显著下降、类胡萝卜素含量极显著下降; 抽穗期叶绿素a、叶绿素b和类胡萝卜素的含量均极显著下降, 净光合速率、气孔导度和蒸腾速率也极显著下降, 胞间CO₂浓度则无差异(图2)。农艺性状分析表明, esl5的株高略低于野生型, 但差异不显著, 有效穗显著降低, 穗粒数、实粒数和结实率极显著降低, 尤其是结实率, 仅为野生型的44.57%; esl5的千粒重为30 g, 极显著高于野生型的26.5 g, esl5的千粒重增加是由籽粒宽和籽粒厚极显著增加造成的(表1, 图1-E, F)。

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	图1 突变体 esl5及野生型缙恢10号的形态特征Fig. 1 Plant morphology of the esl5 and the wild type Jinhui 10

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	图2 esl5及野生型(WT)的光合参数Fig. 2 Photosynthetic parameters of the wild type (WT) and esl5

表1 野生型(WT)和 esl5农艺性状分析 Table 1 Agronomic trait of the wild type (WT) and the esl5

2.2 细胞学观察

野生型缙恢10号剑叶的细胞呈卵圆形, 细胞膜紧贴细胞壁, 内含叶绿体、线粒体、高尔基体、核糖体、内质网等细胞器(图3-A); 叶绿体由叶绿体膜、类囊体和基质构成, 基质排列规则、致密(图3-B)。与野生型相比, esl5的细胞形态没有太大变化, 细胞膜严重破坏, 液泡变大, 线粒体、高尔基体、核糖体等细胞器部分或完全降解(图3-C); 叶绿体膜变薄、破裂, 基质片层松散、呈线性、排列不规则(图3-D)。

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	图3 野生型(WT)和 esl5剑叶的细胞结构Fig. 3 Cell structure of flag leaf in the wild type and the esl5

2.3 理化分析

丙二醛和可溶性蛋白是植物衰老的重要生理指标, 与野生型相比, esl5的丙二醛和可溶性蛋白含量分别下降了32.00%和39.83%, 均达极显著差异水平(图4-A, B), 进一步表明 esl5是一个叶片早衰突变体。超氧阴离子(O₂^-)、羟基自由基(·OH)和H₂O₂等活性氧(reactive oxygen species, ROS)在生物体内过量累积, 可诱发氧化应激反应, 引起细胞内膜脂过氧化以及蛋白/DNA的损伤等, 导致器官甚至机体的死亡^{[ 24]}。

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	图4 抽穗期野生型(WT)和突变体 esl5的生理特性Fig. 4 Physiological characteristics of the wild type (WT) and esl5 at heading stage

esl5中的O₂^-含量极显著下降, 仅为野生型的35.0%, H₂O₂和·OH的含量则极显著升高, 其中, 突变体·OH的含量为95.95 U mg^-1 prot, 与野生型的58.698 U mg^-1prot相比, 升高了1.63倍。 esl5中POD的活性为48.864 OD₄₇₀ min^-1g^-1FW, 与野生型的33.325 OD₄₇₀ min^-1g^-1FW相比, 升高了46.63%, 达到极显著差异水平; esl5的SOD和CAT酶活性分别下降了41.42%和25.73%, 与野生型相比也达到极显著差异水平(图4)。推测ROS积累导致过氧化损伤可能是造成 esl5早衰的主要原因。

2.4 遗传分析

突变体 esl5和日本晴杂交组合的F₁植株叶色浓绿, 无早衰表型; F₂群体中, 苗期植株呈正常的绿色, 分蘖后期叶色逐渐分化为绿色和黄绿两类, 绿色植株的叶色一直保持到成熟, 黄绿色植株则在抽穗期叶片黄化早衰, 与突变体 esl5的表型类似。3356株F₂群体中, 正常型2546株, 突变型810株, χ²测验符合3∶1的分离比(χ²= 1.34, χ²_0.05= 3.84), 表明 esl5受1对隐性核基因调控。

2.5 基因定位

均匀分布于水稻12条染色体上的480对SSR引物, 112对在日本晴和 esl5间呈现多态性, 利用这112对引物进一步分析正常基因池和突变基因池, 发现, 位于第3染色体上的SSR标记RM6959和RM3400在基因池之间也表现多态性, 暗示可能的连锁关系。利用70株典型F₂隐性单株进行验证, 初步将 ESL5定位在SSR标记RM6959和RM3400之间, 遗传距离分别为4.29 cM和3.57 cM (图5)。在初步定位的基础上, 进一步合成新的SSR引物并开发设计Indel标记, 利用亲本间呈现多态性的标记进行精细定位。结果发现, 在810株F₂隐性单株中, Indel标记Indel03-1和Indel03-2分别有2个和3个交换株, 且交换单株不同, 标记的引物序列见表2。因此, 最终将 ESL5定位在标记Indel03-1和Indel03-2之间物理距离约83.4 kb的范围内, 位于BAC克隆AC087220.9上(图5)。在定位区间内, 检测到11个注释基因, 其中, 3个编码转座子蛋白, 3个编码表达蛋白, 2个编码F-box蛋白, 其他分别编码蛋白磷酸酶2C、重金属相关蛋白以及LTV1蛋白。

表2 新开发的Indel标记 Table 2 Newly designed Indel markers

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	图5 ESL5基因在水稻第3染色体上的分子定位Fig. 5 Molecular mapping of ESL5 on rice chromosome 3

3 讨论

植物衰老通常表现叶片黄化、叶绿体降解、光合色素和可溶性蛋白含量下降等特性。苗期突变体与野生型无明显差异, 分蘖期, esl5的叶色呈淡绿色, 抽穗期叶尖开始黄化衰老, 然后扩展至整个叶片中上部。 esl5黄化部位的光合色素和可溶性蛋白含量均极显著下降, 细胞结构异常, 主要表现为细胞膜破裂、细胞器溶解或部分溶解、叶绿体基质片层疏松、排列不规则等。

esl5是一个新型淡叶早衰突变体, 与已报道的水稻自然早衰突变体明显不同。在水稻中, 已鉴定的自然早衰突变体除叶片衰老外, 多伴随斑点、不育等性状, 衰老时期各有异同。 psl1苗期叶片即黄化衰老^{[ 9]}; pgl2抽穗前倒四叶开始衰老, 抽穗时倒三叶开始衰老, 穗完全抽出后, 倒二叶开始变黄衰老, 到灌浆期结束时剑叶完全衰老死亡^{[ 10]}; psl3是一个显性苗期早衰突变体^{[ 11]}; pse(t)抽穗期首先在老叶上出现褐色斑点, 接着整个叶片黄化衰老^{[ 12]}; es-t苗期叶片即出现黄化衰老, 并伴有铁锈色斑点^{[ 13]}; esl2在孕穗期叶片中上部开始出现黄化早衰^{[ 14]}; esl3全生育期内叶片均出现褐化枯死现象^{[ 15]}。 Lad分蘖后期表现叶尖卷曲枯死^{[ 16]}, sms1除叶片早衰外, 还表现雄性不育^{[ 17]}。

与野生型相比, esl5的O₂^-含量极显著降低, ·OH和H₂O₂含量则极显著升高。O₂^-、·OH和H₂O₂是植物体内ROS的主要形式, 是线粒体、叶绿体和过氧化物酶体等新陈代谢的次生产物, ROS通常被认为是一种有毒的化学物质, 也是重要的信号传导分子^{[ 25, 26]}。其中, ·OH是一种非选择性氧化剂, 也是氧化性最强的ROS自由基, 可引发不饱和脂肪酸发生脂质过氧化, 核酸断裂、蛋白质和多糖分解, 膜结构损伤及功能丧失。近来发现, ROS不足能导致小鼠精子的不育, 推测生物体内ROS可能存在一种动态的平衡, 过高过低都会对细胞造成损伤^{[ 27, 28]}。SOD是O₂^-的主要清除因子, 可通过歧化反应将O₂^-转化为H₂O₂, 再进一步被CAT等分解成没有毒害的H₂O和O₂^{[ 29, 30]}。推测SOD、CAT等酶活性的降低, 致使 esl5产生的·OH和H₂O₂不能及时清除, 造成ROS含量失衡, 导致 esl5叶片的衰老。

本研究中, ESL5被定位在第3染色体83.4 kb的物理范围内, 包含11个注释基因, 均没有进行功能研究。生物信息学分析发现11个注释基因中, 2个编码F-box蛋白, 1个编码蛋白磷酸酶2C (PP2C)。PP2C是蛋白磷酸酶的一个分支, 蛋白磷酸酶是蛋白质可逆磷酸化调节机制中的关键酶^{[ 31]}。 AtNAP是一个NAC转录因子, 其表达上调促进叶片衰老, 下调则延迟衰老^{[ 32]}。 SAG113编码一个PP2C, 是AtNAP的直接靶蛋白, 通过调控ABA介导的气孔运动和水分流失调控叶片衰老^{[ 33, 34]}。F-box蛋白是植物体内一个大的蛋白家族, 参与乙烯、生长素、赤霉素、茉莉酸等激素的信号传导途径, 涉及花器官发育、光形态建成、胁迫反应等生物学过程^{[ 35]}。例如, ORE9即编码一个F-Box蛋白, 通过泛素依赖的蛋白质水解调控衰老相关蛋白, 从而延长拟南芥的叶片衰老^{[ 36]}。但F-box或PP2C是否是 ESL5的候选基因, 尚需进一步研究。

4 结论

esl5是一个新型淡叶早衰突变体, 苗期叶片正常, 分蘖后期变成黄绿色, 孕穗期叶尖开始黄化衰老, 衰老部位细胞膜降解、细胞器部分或完全解体, 叶绿体基质疏松、排列不规则, 光合色素和光合速率极显著下降。与野生型相比, esl5的O₂^-含量、SOD和CAT的活性极显著降低, ·OH和H₂O₂含量则极显著升高, 保护酶活性降低导致ROS平衡失调可能是导致 esl5早衰的主要原因。 esl5受隐性核基因调控, 基因定位在第3染色体上83.4 kb的物理范围内, 包含11个注释基因。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

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2011

0.0

... 植物衰老是一个细胞程序性死亡的过程, 涉及叶绿体发育、叶绿素合成与降解、激素和自由基代谢等途径, 既受遗传基因的调控, 也受外界环境的影响^[1] ...

2007

25.962

0.0

... 自然衰老过程中, 营养物质从叶片转移至种子, 有利于植物的生长繁殖, 是其长期进化过程中形成的一种适应性^[2] ...

2012

3.75

0.7555

... 但在农业生产中, 叶片早衰则会导致作物产量和品质的下降, 如水稻穗中60%~70%的碳是由功能叶合成转运而来的^[3] ...

2006

6.555

0.0

... 如OsDOS编码一个水稻CCCH-Type锌指蛋白, 功能缺失促进叶片衰老, 过表达则延缓叶片衰老^[4] ...

2013

3.518

0.0

. 2013, 82:519-528 DOI:10.1007/s11103-012-9974-2

Towards systems biological understanding of leaf senescence

<li>1.Tobacco Research Institute, Chinese Academy of Agricultural Sciences, Qingdao, 266101, China</li><li>2.Key Laboratory of Tobacco Biology and Processing, Ministry of Agriculture, Qingdao, 266101, China</li>

Abstract

The application of systems biology approaches has greatly facilitated the process of deciphering the molecular mechanisms underlying leaf senescence. Analyses of the leaf senescence transcriptome have identified some of the major biochemical events during senescence including protein degradation and nutrient remobilization. Proteomic studies have confirmed these findings and have suggested up-regulated energy metabolism during leaf senescence which might be important for cell viability maintenance. As a critical part of systems biology, studies involving transcription regulation networking and senescence-inducing signaling have deepened our understanding on the molecular regulation of leaf senescence. The important next steps towards a systems biological understanding of leaf senescence will be discussed.

... 基因克隆的策略很多, 其中, 转录组和蛋白质组是植物衰老机制研究的常用手段^[5] ...

2005

6.582

0.0

... 在双子叶模式植物拟南芥中, 利用表达谱芯片技术从衰老的叶片中鉴定到800个上调SAG (senescence-associated genes)^[6], 在水稻剑叶自然衰老过程中, 则检测到533个上调SAG ^[7] ...

2008

3.518

0.0

. 2008, 67:37-55 DOI:10.1007/s11103-008-9300-1

Identification of early senescence-associated genes in rice flag leaves

1.Huazhong Agricultural University National Key Laboratory of Crop Genetic Improvement and National Centre of Plant Gene Research Wuhan 430070 P.R. China

Leaf senescence is one of the key stages of plant leaf development. It is a highly complex but ordered process involving expression of large scale senescence associated genes, and its molecular mechanisms still remain unclear. By using suppression subtractive hybridization, 815 ESTs that are up-regulated at the onset of rice flag leaf senescence have been isolated. A total of 533 unigenes have been confirmed by macroarray detection and sequencing. 183 of these unigenes have GO annotations, involved in macromolecule metabolism, protein biosynthesis regulation, energy metabolism, gene expression regulations, detoxification, pathogenicity and stress, cytoskeleton organization and flower development. Another 121 unigenes co-localized with previously reported known stay-green QTLS. RT-PCR analysis on the other novel genes indicated that they can be up-regulated in natural early senescence and induced by hormone. Our results indicate that senescence is closely related to various metabolic pathways, thus providing new insight into the onset of leaf senescence mechanism.

2013

6.126

0.9487

... 图位克隆是基因分离和功能研究中最直接有效的技术手段, 植物基因组测序的大量完成进一步促进了这一技术的发展, 其关键是优良突变体的获得^[8] ...

2006

0.0

0.95

. 2006, 51:2986-2992 DOI:10.1007/s11434-006-2222-x

Genetic analysis and molecular mapping of a presenescing leaf gene psl1 in rice (Oryza sativa L.)

(1)Key Laboratory of the Ministry of Education for Plant Functional Ge-nomics/Jiangsu Key Laboratory for Crop Genetics and Physiology, Yangzhou University ,Yangzhou 225009 ,China

A rice psl1 (presenescing leaf) mutant was obtained from a japonica variety Zhonghua 11 via radiation of ⁶⁰Co-γ in M2 generation. Every leaf of the mutant began to wither after it reached the biggest length, while the leaves of the wild variety could keep green for 25―35 d. In this study, genetic analysis and gene mapping were carried out for the mutant identified. The SSR marker analysis showed that the mutant was controlled by a single recessive gene (psl1) located on chromosome 2. Fine mapping of the psl1 locus was conducted with 34 new STS markers developed around psl1 anchored region based on the sequence diversity between Nipponbare and 93-11. The psl1 was further mapped between two STS markers, STS2-19 and STS2-26, with genetic distances of 0.43 and 0.11 cM, respectively, while cosegregated with STS2-25. A BAC contig was found to span the psl1 locus, the region being delimited to 48 kb. This result was very useful for cloning of the psl1 gene.

... 目前, 在水稻中仅鉴定到十几个叶片自然早衰突变体, 其中9个基因已定位, 突变体psl1、pgl2、psl3、pse(t)、es-t、esl2、esl3、sms1和lad的调控基因分别定位在第2、第8、第7、第7、第5、第4、第5、第8和第11染色体上^{[9,10,11,12,13,14,15,16,17]} ...

... psl1苗期叶片即黄化衰老^[9] ...

2007

0.0

0.3579

. 2007, 14:172-180 DOI:10.1016/S1672-6308(07)60024-9

Identification and fine mapping of a gene related to pale green leaf phenotype near centromere region in rice (Oryza sativa L.)

Abstract

A thermo-insensitive pale green leaf mutant (pgl2) was isolated from T-DNA inserted transgenic lines of rice (Oryza sativa L. subsp. japonica cv. Nipponbare). Genetic analysis indicated that the phenotype was caused by a recessive mutation in a single nuclear-encoded gene. To map the PGL2 gene, an F₂ population was constructed by crossing the mutant with Longtefu (Oryza sativa L. subsp. indica). The PGL2 locus was roughly linked to SSR marker RM331 on chromosome 8. To finely map the gene, 14 new InDel markers were developed around the marker, and PGL2 was further mapped to a 2.37 Mb centromeric region. Analysis on chlorophyll contents of leaves showed that there was no obvious difference between the mutant and the wild type in total chlorophyll (Chl) content, while the ratio of Chl a/Chl b in the mutant was only about 1, which was distinctly lower than that in the wild type, suggesting that the PGL2 gene was related to the conversion between Chl a and Chl b. Moreover, the method of primer design around the centromeric region was discussed, which would provide insight into fine mapping of the functional genes in plant centromeres.

... pgl2抽穗前倒四叶开始衰老, 抽穗时倒三叶开始衰老, 穗完全抽出后, 倒二叶开始变黄衰老, 到灌浆期结束时剑叶完全衰老死亡^[10] ...

2010

0.0

0.95

. 2010, 55:2517-2521 DOI:10.1007/s11434-010-4013-7>

Genetic analysis and gene mapping of dominant presenescing leaf gene PSL3 in rice (Oryza sativa L.)

FANG LiKui, LI YunFeng, GONG XiaoPing, SANG XianChun, LING YingHua, WANG XiaoWen, CONG YunFei & HE GuangHua

Leaf senescence as an active process is essential for plant survival and reproduction. However, premature senility is harmful to agricultural production. In this study, a rice mutant, named as psl3 (presescing leaf 3) isolated from EMS-treated Jinhui 10, displays obvious premature senility features both in morphological and physiological level. Genetic analysis showed that mutant trait was controlled by a single dominant gene (PSL3), which was located on rice chromosome 7 between SSR marker c7sr1 and InDel marker ID10 with an interval of 53.5 kb. The result may be useful for the isolation of the PSL3 gene.

... psl3是一个显性苗期早衰突变体^[11] ...

2005

1.668

0.0

... pse(t)抽穗期首先在老叶上出现褐色斑点, 接着整个叶片黄化衰老^[12] ...

2011

0.0

0.95

. 2011, 56:2437-2443 DOI:10.1007/s11434-011-4587-8

Characterization and fine mapping of an early senescence mutant (es-t) in Oryza sativa L

1. State Key Laboratory of Rice Biology, China National Rice Research Institute, Hangzhou 310006, China; 2. College of Life and Environment Sciences, Hangzhou Normal University, Hangzhou 310036, China

An es-t (early senescence-temporary) mutant, produced by ethylene methylsulfonate treatment of strain Nipponbare, was identified in rice. The leaves of es-t appeared yellow at the seedling stage, and had decreased chlorophyll content. Rust spots were found during growth in es-t, especially at the leaf margin and tip. The plants showed a typical early-senescence phenotype at the milky stage. The leaf surface of es-t appeared smoother than wild-type leaves under a scanning electron microscope, because the leaves lack siliceous protuberances around the stoma. Chloroplasts grow abnormally and are filled with many starch grains in es-t. Paraffin section analysis showed that the development of the sclerenchyma cells and vascular bundles were also abnormal in es-t. Genetic analysis indicated that es-t was controlled by a recessive gene, which was finely mapped to a 42-kb interval on chromosome 5. These results will facilitate the positional cloning and functional studies of the gene.

... es-t苗期叶片即出现黄化衰老, 并伴有铁锈色斑点^[13] ...

2012

0.0

1.667

. 2012, 38:1347-1353 DOI:10.3724/SP.J.1006.2012.01347

Genetic analysis and gene mapping of early senescence leaf mutant esl2 in rice

Rice Research Institute, Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crops, Engineering Research Center of South Upland Agriculture, Ministry of Education, Southwest University, Chongqing 400716, China

An early senescence leaf mutant temporarily designated as esl2was discovered in the progeny of an excellent indica restorer line Jinhui 10 with seeds treated by ethyl methane sulfonate (EMS). Themutant kept normal at seedling stage and showed etiolated senescence leaves from booting sage to the maturity, especially at tip and margin of leaf blade. By contrast with the wild type, photosynthetic pigment contents decreased significantly at both booting stage and heading stage, superoxide dismutase (SOD) activity and soluble protein contents decreased significantly, while the activities of peroxidase (POD) and catalase (CAT), and contents of reactive oxygen species (ROS), malondialdehyde (MDA) and praline increased significantly at heading stage in the esl2 mutant. The assay of Transmission Electronic Microscope (TEM) observation was performed and demonstrated that the most cells appeared irregular and void with endolysis and disrupted organelles at senescence part of the mutational leaf, concomitantly, abnormal chloroplast appeared and showed dissolved membrane, indistinct grana, disorganized lamellar structure and incorrect thylakoid. Genetic analysis suggested that the mutational trait was controlled by one nuclear recessive gene. Xinong 1A was crossed with the esl2 and 1005 mutational F₂single plants were used for gene mapping. Finally, Esl2 locus was mapped between SSR marker RM17122 and swu4-13 on the chromosome 4 with physical distance of 244 kb. This result provides a foundation of Esl2 gene cloning by map-based strategy as well as its functional analysis.

利用EMS诱变水稻籼型恢复系缙恢10号, 从其后代中鉴定出一个早衰突变体esl2, 苗期正常, 孕穗期开始叶尖和叶缘黄化衰老。与野生型相比, 孕穗期和抽穗期光合色素含量均显著下降, 抽穗期倒一叶的超氧化酶歧化酶(SOD)活性、可溶性蛋白(SP)含量降低, 活性氧(ROS)、过氧化物酶(POD)、过氧化氢酶(CAT)、丙二醛(MDA)和脯氨酸(PRO)含量或活性增加。超微结构观察表明, esl2衰老部位的细胞多中空且形状不规则, 伴随细胞裂解、细胞质溶解等特征; 同时, 叶绿体结构异常, 叶绿体膜溶解、基粒模糊, 基质片层疏松, 类囊体发育异常。遗传分析表明, 该突变体受一对隐性核基因调控。利用1 005株西农1A/esl2的F₂隐性定位群体, 最终将Esl2定位在第4染色体SSR标记RM17122和swu4-13之间, 物理距离约244 kb, 这为Esl2基因的克隆和功能研究奠定了基础。

... 3 生理指标检测参照文献^[14]描述的方法测定超氧阴离子(O₂^-)含量 ...

... esl2在孕穗期叶片中上部开始出现黄化早衰^[14] ...

2013

0.0

1.667

. 2013, 39:862-867 DOI:10.3724/SP.J.1006.2013.00862

Identification and gene mapping of rice early senescent leaf (esl3) mutant

<div style="line-height: 150%">Rice Research Institute of Southwest University / Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crops / Engineering Research Center of South Upland Agriculture, Ministry of Education, Chongqing 400716, China</div>

Leaf early senescence can directly decrease crop photosynthesis, yield and quality. Therefore, the identifications of mutants with early senescence and its gene function play key roles in crop genetic improvement. A mutant with early senescent leaf blades, named as esl3, was identified from a EMS-induced progeny in restorer line of Jinhui 10. The middle-upper leaf in mutation displayed a brown and withered phenotype from seedling stage to maturity. Correspondingly, the chlorophyll content and net photosynthetic rate declined in the abnormal leaves while enhanced in the normal green part of leaves compared with the wild type. Agronomic traits, such as panicle number, panicle length, grains per panicle, 1000-grain weight, plant height and weight of dry matter, were reduced significantly or very significantly except for seed setting rate as compared with those of wild type. Genetic analysis showed the senescence was controlled by a recessive nuclear gene. Using 391 mutation individuals in a F₂ generation of Nippomese/esl3, ESL3 was mapped between SSR marker RM19085 and Indel marker Ind05-2 on the chromosome 5 with physical distance 91 kb including 14 annotated genes. All of these results provide a foundation for the further gene cloning and functional analysis of ESL3 gene.

叶片早衰直接降低作物的光合作用、产量和品质。因此，鉴定早衰突变体和研究其基因功能对于作物的遗传改良具有重要的作用。esl3来源于水稻籼型恢复系缙恢10号的EMS诱变库，苗期叶片中上部即呈现褐化枯萎，该特征一直持续到植株成熟。与野生型相比，突变体衰老部位叶绿素和光合速率极显著下降，绿色部位光合色素和光合速率则略有升高。农艺性状分析发现，结实率无显著变化，有效穗、穗长、穗粒数、千粒重、株高和干物质重则显著或极限著下降。遗传分析表明，esl3叶片早衰枯死性状受1对隐性核基因控制。利用391株日本晴/esl3的F₂突变型单株，最终把ESL3基因定位在第5染色体SSR标记RM19085和Indel标记Ind05-2之间，物理距离91 kb，包含14个注释基因，为下一步调控基因的克隆和功能研究奠定了基础。

... esl3全生育期内叶片均出现褐化枯死现象^[15] ...

2012

0.0

1.667

. 2012, 38:168-173 DOI:10.3724/SP.J.1006.2012.00168

Analysis of phenotype and physiology of leaf apex dead mutant (lad) in rice and mapping of mutant gene

Rice Research Institute, Southwest University / Chongqing Key Laboratory of Application and Safety of Genetically Modified Crops, Chongqing 400716, China?

Leaf plays an important role in photosynthesis. Studying the leaf premature senescence is important for increasing the economic yield of crop. A new mutant of rice leaf apex dead reported in the paper derived from a restorer line Jinhui10 treated by EMS. In the mutant, the leaf apexes appeared normal before the 5th leaf unfolded. However, the leaf apexes of all leaves except the 5th leaf exhibited yellow and gradually died when the 5th leaf fully expanded; and afterwards the apexes of new fully expanded leaves became yellow and dead. The physiological and biochemical analysis showed that the lad mutant contained lower contents of chlorophyll and soluble protein compared with the wild type control, and the activity of SOD was abnormal. Again, the plant height, leaf length, grain number and seed setting rate of the lad mutant were significantly lower than those of the control. Thepremature senescence mutant was controlled by a single recessive nuclear gene. And the LAD gene was mapped between the SSR markers SWU11-19 and SWU11-5 on the 11th chromosome, with 13 cM of the genetic distance. Furthermore, the LAD gene was co-segregated with the markers SWU11-25 and SWU11-27. The results laid a favorable foundation for further map-based cloning and functional analysis of the LAD gene.

叶片作为植物的主要光合作用场所，研究其早衰机制对提高作物的经济产量具有非常重要的意义。本研究报道了一个来自EMS诱变优良恢复系缙恢10号的新水稻叶尖早衰突变体lad (leaf apex dead)，其叶尖在第5片叶抽出前呈正常状态，当第5叶完全抽出之后前5叶的叶尖变黄并最终枯死；随后的叶子在完全抽出后，叶尖也逐渐变黄并枯死。对该突变体的生理生化分析发现，其叶绿素含量、可溶性蛋白含量明显下降，SOD酶活性异常。其株高、叶长、粒数等也都显著降低。经遗传分析，该突变性状受一对隐性单基因控制，利用分子标记将该基因定位于第11染色体SSR标记SWU11-19和SWU11-5之间，遗传距离为13 cM，并且与SSR标记SWU11-25和SWU11-27共分离。本研究结果为该基因的进一步克隆和功能研究奠定了良好基础。

... Lad分蘖后期表现叶尖卷曲枯死^[16], sms1除叶片早衰外, 还表现雄性不育^[17] ...

2010

2.076

1.323

. 2010, 37:47-55 DOI:10.1016/S1673-8527(09)60024-2

Characterization and mapping of a novel mutant sms1 (senescence and male sterility 1) in rice

①College of Life and Environment Sciences, Shanghai Normal University, Shanghai 200234, China ②Institute of Crop and Nuclear Technology Utilization, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China ③Institute of Plant Physiology & Ecology Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai 200032, China

Plant senescence plays diverse important roles in development and environmental responses. However, the molecular basis of plant senescence is remained largely unknown. A rice spontaneous mutant with the character of early senescence and male sterility (sms) was found in the breeding line NT10-748. In order to identify the gene SMS1 and the underlying mechanism, we preliminarily analyzed physiological and biochemical phenotypes of the mutant. The mutant contained lower chlorophyll content compared with the wild type control and was severe male sterile with lower pollen viability. Genetic analysis showed that the mutant was controlled by a single reces-sive gene. By the map-based cloning approach, we fine-mapped SMS1 to a 67 kb region between the markers Z3-4 and Z1-1 on chromo-some 8 using 1,074 F2 recessive plants derived from the cross between the mutant sms1 (japonica) × Zhenshan 97 (indica), where no known gene involved in senescence or male sterility has been identified. Therefore the SMS1 gene will be a novel gene that regulates the two developmental processes. The further cloning and functional analysis of the SMS1 gene is under way.

植物的衰老在其生长发育以及对环境的应答方面都起着重要作用。然而，到目前为止，有关植物衰老的分子基础并不清楚。在育种后代NT10-748群体中，发现了一个具有早衰和雄性不育（senescence and male sterility，sms）性状的自然突变体。为了鉴定SMS1基因和相关机制，分析了该突变体的生理生化特征,发现突变体较野生型有较低的叶绿素含量和花粉活力，从而导致严重的雄性不育。遗传分析显示该突变体由一对隐性基因控制。通过图位克隆法，利用1,074株从sms1（粳稻）和Zhenshan 97（籼稻）杂交的F2代定位群体，将SMS1定位于第八号染色体分子标记Z3-4 和Z1-1之间67kb区间内，该区间内目前还没有已知的与衰老或雄性不育相关的基因被鉴定。因此，SMS1将会是一个与调控这两个发育过程有关的新基因，进一步克隆和功能分析正在进行中。

... Lad分蘖后期表现叶尖卷曲枯死^[16], sms1除叶片早衰外, 还表现雄性不育^[17] ...

2012

6.555

0.0

... 4个基因已克隆, NOE1与OsDMI3均与过氧化反应有关, NOE1编码过氧化氢酶OsCATC, 基因功能缺失通过激活硝酸还原酶产生大量NO, 从而诱导叶片早衰, noe1突变体内H₂O₂的含量显著高于野生型^[18] ...

2012

6.126

0.9487

... OsDMI3位于水稻第5染色体上, 编码Ca²⁺/CaM依赖型蛋白激酶, 在ABA诱导的抗氧化防护过程中具有重要的功能, OsDMI3功能缺陷可造成植株体内H₂O₂的过量积累, 导致叶片衰老^[19] ...

2012

6.126

0.9487

... RLS1编码一个C端具有ARM结构域的核苷酸结合蛋白, 参与程序性细胞死亡过程, 叶绿素的快速降解是导致rls1衰老的主要原因^[20] ...

2010

6.736

0.0

... SPL28编码网格相关受体蛋白复合体亚基(AP1M1), 参与高尔基的物质运输途径, 其功能缺失引起植物过敏性反应, 形成叶片类病斑, 导致水稻叶片早衰^[21] ...

1994

2.699

0.0

... 2 光合色素含量和光合作用测定田间种植情况下, 参照文献^[22]描述的方法在分蘖期和抽穗期分别测定esl5和缙恢10号的光合色素含量, 并于抽穗期利用LI-6400型便携式光合作用仪测定净光合速率、气孔导度、蒸腾速率和胞间CO₂浓度 ...

2003

0.0

. 2003, 25:705-707

The simple gain of templates of rice genomes DNA for PCR

用碱处理水稻嫩叶，取碱处理水稻叶片浸出液直接用作PCR扩增的模板，扩增结果稳定、准确，与常规提取的DNA扩增效果相比没有显著差异。用此方法制备的模板室温下2周之内、4℃下3周之内、-20℃下4个月以上扩增结果不变。此方法所需试剂均为常规试剂，具有需要材料少、成本低、简便、快捷等优点，尤其适合材料比较宝贵的转基因水稻的预筛选和大样本量的PCR检测，为分子标记技术的实际应用创造了条件。
Abstract:The solution of alkali-treated fresh rice leaves was used directly as the templates of PCR.The amplified results were stable,reliable,and had no difference compared with that amplified with rice total DNA extracted by common method.The stable results can still be obtained based on the templates kept at 25℃ for tow weeks,at 4℃ for three weeks,at -20℃ for over four months.With this technique,less material and only common reagent are required,which is especially adapted to the screening of the precious trangenic rice in advance and large-scale PCR tests.

... 采用改良的CTAB法提取亲本、正常池和突变池的基因组DNA, 利用碱煮法提取F₂定位单株的基因组DNA^[23] ...

2011

0.0

... OH)和H₂O₂等活性氧(reactive oxygen species, ROS)在生物体内过量累积, 可诱发氧化应激反应, 引起细胞内膜脂过氧化以及蛋白/DNA的损伤等, 导致器官甚至机体的死亡^[24] ...

2012

4.304

0.0

... OH和H₂O₂是植物体内ROS的主要形式, 是线粒体、叶绿体和过氧化物酶体等新陈代谢的次生产物, ROS通常被认为是一种有毒的化学物质, 也是重要的信号传导分子^[25,26] ...

2007

5.372

0.0

2013

25.315

0.0

... 近来发现, ROS不足能导致小鼠精子的不育, 推测生物体内ROS可能存在一种动态的平衡, 过高过低都会对细胞造成损伤^[27,28] ...

2013

14.619

0.0

... 近来发现, ROS不足能导致小鼠精子的不育, 推测生物体内ROS可能存在一种动态的平衡, 过高过低都会对细胞造成损伤^[27,28] ...

2012

2.855

0.0

. 2012, 249:469-481 DOI:10.1007/s00709-011-0308-z

Leaf senescence and abiotic stresses share reactive oxygen species-mediated chloroplast degradation

<li>1.Stress Physiology and Biochemistry Laboratory, Water Technology Centre, Indian Agricultural Research Institute (IARI), New Delhi, 110012, India</li>

Abstract
Leaf senescence is a genetically programmed decline in various cellular processes including photosynthesis and involves the hydrolysis of macromolecules such as proteins, lipids, etc. It is governed by the developmental age and is induced or enhanced by environmental stresses such as drought, heat, salinity and others. Internal factors such as reproductive structures also influence the rate of leaf senescence. Reactive oxygen species (ROS) generation is one of the earliest responses of plant cells under abiotic stresses and senescence. Chloroplasts are the main targets of ROS-linked damage during various environmental stresses and natural senescence as ROS detoxification systems decline with age. Plants adapt to environmental stresses through the process of acclimation, which involves less ROS production coupled with an efficient antioxidant defence. Chloroplasts are a major site of protein degradation, and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) is rapidly and selectively degraded during senescence and stress. The process of protein degradation is initiated by ROS and involves the action of proteolytic enzymes such as cysteine and serine proteases. The mechanism of Rubisco degradation still remains to be elucidated. The molecular understanding of leaf senescence was achieved through the characterization of senescence-associated genes and various senescence mutants of Arabidopsis, which is a suitable model plant showing monocarpic senescence. The regulation of senescence involves many regulatory elements composed of positive and negative elements to fine-tune the initiation and progression of senescence. This review gives an overview on chloroplast protein degradation during leaf senescence and abiotic stresses and also highlights the role of ROS management in both processes.

... SOD是O₂^-的主要清除因子, 可通过歧化反应将O₂^-转化为H₂O₂, 再进一步被CAT等分解成没有毒害的H₂O和O₂^[29,30] ...

2004

2.699

0.0

... SOD是O₂^-的主要清除因子, 可通过歧化反应将O₂^-转化为H₂O₂, 再进一步被CAT等分解成没有毒害的H₂O和O₂^[29,30] ...

2013

4.25

0.0

... PP2C是蛋白磷酸酶的一个分支, 蛋白磷酸酶是蛋白质可逆磷酸化调节机制中的关键酶^[31] ...

2006

6.582

0.0

... AtNAP是一个NAC转录因子, 其表达上调促进叶片衰老, 下调则延迟衰老^[32] ...

2012

6.555

0.0

... SAG113编码一个PP2C, 是AtNAP的直接靶蛋白, 通过调控ABA介导的气孔运动和水分流失调控叶片衰老^[33,34] ...

2012

6.582

0.0

... SAG113编码一个PP2C, 是AtNAP的直接靶蛋白, 通过调控ABA介导的气孔运动和水分流失调控叶片衰老^[33,34] ...

2006

0.0

. 2006, 28:1337-1342

The function of f-box protein in plant growth and development

1. Key Laboratory of Crop Germplasm Resources and Biotechnology, MOA, Institute of Crop Science, Chinese Academy of Agricultural Sciences, Beijing 100081, China; 2. College of Life Science, Hebei Normal University, Shijiazhuang 050016, China

Ubiquitin-mediated proteolysis is involved in many biological processes in eukaryotes. SCF complex is a very important ubiquitin E3 ligase which has been exploited very well in plants. F-box protein characterized by an F-box motif is a subunit of SCF complex, which works as determinant in substrate recognition. Currently, many F-box proteins have been identified in plants which are involving in hormone (e.g., ethylene, auxin, gibberellin and jasmonate ) signal transduction and biological processes, such as self-incompatibility and floral development. F-box proteins may also participate stress response in plants. Recent study suggested that the Arabidopsis F-box protein TIR1 is an auxin receptor. Therefore, F-box protein mediated proteolysis may be an important gene expression mechanism in plants.

在真核生物中, 泛素介导的蛋白降解途径参与了许多生物学过程。SCF复合体是一种非常重要的E3泛素连接酶, 在植物中研究的最为深入。F-box蛋白包含一个F-box 基序, 是SCF复合体的一个亚基, 它决定了底物识别的特异性。目前, 从各种植物中已鉴定出大量的F-box蛋白质, 它们参与了植物激素(乙烯, 生长素, GA, JA)的信号传导以及自交不亲和、花器官发育等生物学过程, F-box蛋白还参与了植物的胁迫反应。最新研究结果显示, 一个F-box蛋白TIR1是生长素的受体。因此, F-box蛋白质介导的泛素化蛋白质降解途径可能是植物基因表达调控的重要机制。

... F-box蛋白是植物体内一个大的蛋白家族, 参与乙烯、生长素、赤霉素、茉莉酸等激素的信号传导途径, 涉及花器官发育、光形态建成、胁迫反应等生物学过程^[35] ...

2001

9.251

0.0

... 例如, ORE9即编码一个F-Box蛋白, 通过泛素依赖的蛋白质水解调控衰老相关蛋白, 从而延长拟南芥的叶片衰老^[36] ...