谷子F-box家族基因的鉴定、分类及干旱响应

引用本文

霍冬英, 郑炜君, 李盼松, 徐兆师, 周永斌, 陈明, 马有志, 闵东红, 张小红. 谷子F-box家族基因的鉴定、分类及干旱响应. 作物学报, 2014, 40(9): 1585-1594[HUO Dong-Ying, ZHENG Wei-Jun, LI Pan-Song, XU Zhao-Shi, ZHOU Yong-Bin, CHEN Ming, MA You-Zhi, MIN Dong-Hong, ZHANG Xiao-Hong. Identification, Classification, and Drought Response of F-box Gene Family in Foxtail Millet. Acta Agronomica Sinica, 2014, 40(9): 1585-1594] 复制到剪切板

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作物学报编辑部

谷子F-box家族基因的鉴定、分类及干旱响应

霍冬英^1,², 郑炜君¹, 李盼松^1,², 徐兆师², 周永斌^1,², 陈明², 马有志², 闵东红^1,^2,^*, 张小红^1,^2,^*

¹旱区作物逆境生物学国家重点实验室 / 西北农林科技大学, 陕西杨凌 712100

²中国农业科学院作物科学研究所 / 农作物基因资源与基因改良国家重大科学工程 / 农业部麦类生物学与遗传育种重点实验室, 北京 100081

^*通讯作者(Corresponding authors): 张小红, E-mail:zhxh2493@126.com; 闵东红, E-mail:mdh2493@126.com

收稿日期:2014-06-16

基金:本研究由国家转基因生物新品种培育重大专项(2014ZX08002-03B, 2011ZX08002002)资助;

摘要

F-box蛋白参与细胞周期调控、细胞凋亡及信号转导等多种生命活动, 对维持植物正常生长发育a和介导非生物胁迫响应等过程发挥重要作用。谷子具有显著的耐旱耐瘠薄等特性, 本研究根据谷子转录组分析结果, 从525个F-box家族成员中鉴定出19个在干旱胁迫下表达量上调的F-box基因, 根据序列相似性将其分为6类, 同一类基因具有相似的内含子-外显子结构。染色体定位分析发现, 这些基因分别分布在谷子的8条染色体上, 其中, 第2条染色体上含F-box基因最多, 有6个。结构域分析结果表明, 19个F-box蛋白均含保守的F-box结构域, 而C端含FBD、WD40、FBA、ZnF、Kelch和LRR等结构域。启动子元件分析表明, 谷子19个F-box基因均含逆境应答元件, 其中, MYB和MYC元件的数量最多(9~78个), 说明这些基因对干旱应答反应可能主要受MYB、MYC转录因子调控。转录组分析结果表明,SiF-box18基因对干旱胁迫的响应远远高于其他F-box成员, 对干旱、高盐、ABA、SA和JA都有响应; 亚细胞定位结果显示, SiF-box18蛋白定位在细胞核中。本研究为进一步深入了解SiF-box18基因的功能提供了依据。

关键词: 谷子; F-box蛋白; 亚细胞定位; 干旱响应; 诱导机制

Identification, Classification, and Drought Response of F-box Gene Family in Foxtail Millet

HUO Dong-Ying^1,², ZHENG Wei-Jun¹, LI Pan-Song^1,², XU Zhao-Shi², ZHOU Yong-Bin^1,², CHEN Ming², MA You-Zhi², MIN Dong-Hong^1,^2,^*, ZHANG Xiao-Hong^1,^2,^*

¹State Key Laboratory of Crop Stress Biology for Arid Areas / Northwest A&F University, Yangling 712100, China

² Institute of Crop Science, Chinese Academy of Agricultural Sciences / National Key Facility for Crop Gene Resources and Genetic Improvement / Key Laboratory of Crop Genetics and Breeding, Ministry of Agriculture, Beijing 100081, China

Fund:

Abstract

F-box proteins play critical roles in plant growth and development and respond to abiotic stresses, involving in cell-cycle regulation, cell apoptosis and signal transduction. Foxtail millet has a character with drought-tolerance. We identified 19 up-regulated F-box genes from 525 F-box families under stress in foxtail millet based on the transcriptome sequencing and divided them into six subgroups, and the genes in the same subgroup had the same intron-exon structure. These genes were unevenly distributed on eight foxtail millet chromosomes, and there were six genes on chromosome 2, that gene number was more than that contained on other chromosomes. The analysis of conserved domain of 19 genes showed that all the protein sequences had a conserved F-box motif, and several other domain also existed in C-terminal including FBD, WD40, FBA, ZnF, and Kelch LRR. Putativecis-acting elements showed that the number of MYB and MYC was the most, which ranged from 9 to 78. Transcriptome sequencing showed thatSiF-box18 was more hypersensitive to the drought stress than other F-box members and response to drought, salt, ABA, GA, SA, and JA based on the real time PCR. The transient expression inArabidopsisprotoplasts displayed that SiF-box18 protein was localized in nuclear section. This study provided experimental data for further studying the functions of theSiF-box18 gene.

Keyword: Foxtail millet; F-box protein; Subcellular localization; Drought response; Induction mechanism

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在适应干旱等非生物逆境胁迫中, 植物不断进化自身防御系统以减少水分损失或增加水分吸收等^{[ 1]}。蛋白质的降解在植物逆境信号的响应、细胞信号转导等过程中发挥重要作用, 对于维持植物正常生长发育具有重要意义^{[ 2]}。大多数蛋白通过泛素/26S蛋白酶体途径降解。泛素介导的蛋白质降解过程主要有3种酶参与, 包括泛素活化酶(E1), 泛素结合酶(E2)及泛素连接酶(E3)。在ATP供能的情况下, E1的半胱氨酸残基与泛素C末端的甘氨酸形成高能硫酯键, 使泛素激活, 被转移到E2的丝氨酸残基上, 随后E2和E3共同识别底物蛋白, 对其进行泛素化修饰。E3在泛素化-酶体途径的底物特异识别过程中起着关键作用。目前已经发现不同类型的E3^{[ 3]}, 主要的一种是Skp1-Cullin1-F-box (SCF)复合体, 由4个亚基组成, 分别为Cullin、SKP1、RBX1和F-box蛋白^{[ 4, 5]}。每个F-box蛋白家族成员的N端都至少含1个大约40~50个保守氨基酸的F-box基序, 并与Skp1结合; C端是结构不同的蛋白-蛋白相互作用结构域, 包括亮氨酸拉链(LRR)、Kelch和WD40等。

F-box蛋白最早在cyclin F的N端被发现而命名^{[ 6]}, 广泛存在于真核生物中, 参与多种植物激素信号的传导、逆境胁迫应答等, 在调节光形态建成、光周期节律、花器官发育及自交不亲和中发挥着重要功能。植物中发现的第一个F-box蛋白是Unusual Floral Organs (UFO), 参与了植物花分生组织与花器官的建成^{[ 7, 8, 9, 10]}。LKP1/ZTL、LKP2/FKL和FKF1可以调控光形态建成、调节生物钟的昼夜周期节律及开花时间^{[ 11, 12, 13, 14]}。F-box蛋白还与植物的自交不亲和性相关, 许多植物的自交不亲和性由S位点控制, 研究表明, 与S位点相关的S-locus-F-box genes (SLF)能在花粉中特异表达^{[ 15, 16, 17, 18]}。

许多F-box蛋白还参与激素的信号传导, TIR1是生长素受体蛋白, 与AUX/IAA发生相互作用, 导致AUX/IAA降解^{[ 19, 20, 21, 22]}; EBF1过量表达引起植物对乙烯不敏感, ebf1和 ebf2突变株则表现出组成型乙烯反应, EBF1/EBF2两者之一发生突变都导致植株对乙烯反应增强, 而双突变体则表现出三重反应^{[ 23, 24]}。赤霉素(GA)的信号通路也可以通过F-box蛋白(SLY1)调节^{[ 25, 26]}。COI1编码F-box蛋白, 在茉莉酸(JA)信号通路中起关键作用^{[ 27, 28, 29]}。

F-box蛋白参与非生物胁迫应答的调控, 如DOR1 (drought tolerance repressor 1)是S位点类F-box蛋白家族的基因, 在保卫细胞中表达, dor使ABA表达量上调, 气孔关闭, 从而增加抗旱性^{[ 30]}。MAX2 (more axillary growth 2)对干旱响应明显, 与野生型相比, max2的保卫细胞对ABA敏感程度降低, 角质层变薄, 在MAX2突变的情况下, 与ABA合成、运输及代谢有关的基因表达量都降低^{[ 31]}。

随着全球水资源短缺和气候变暖形势的日益严峻, 对作物的抗旱节水性进行遗传改良, 提高作物水分利用率, 已成为目前研究的一个热点。谷子具有耐旱、耐贫瘠等特点, 是一种抗逆性很强的作物。然而, 目前, 植物中关于F-box蛋白的研究主要集中在拟南芥、水稻、玉米中。植物已知功能的F-box蛋白只有30个, 其中22个来源于拟南芥(见附表), 谷子中尚未见报道。本研究对谷子进行了转录组测序分析, 筛选出19个在干旱胁迫下表达量上调的F-box基因, 对其基因结构、系统发育关系及染色体定位等进行分析, 并选择对干旱应答最强的基因 SiF-box18作了具体分析, 为进一步研究F-box家族的功能及 SiF-box18基因的功能提供了依据。

1 材料与方法

1.1 试验材料

谷子Tie-8396种子由中国农业科学院作物科学研究所刁现民研究员课题组提供。谷子幼苗在温度22℃、相对湿度65%、光照周期16 h/8 h的条件下培养。生长3周的幼苗经干旱处理1周, 送华大基因进行转录组测序。

1.2 保守域预测

谷子( Setaria italica)基因组数据和cDNA数据均下载于谷子最新数据库版本Phytozome (http://www.phytozome.net/search.php)。利用SMART (http://smart.embl-heidelberg.de/)在线工具分析F-box蛋白结构域, 利用DOG 2.0^{[ 32]}绘制19个F-box家族蛋白的功能结构域示意图。

1.3 进化树与基因结构分析

使用MEGA5对19个谷子F-box基因进行多序列比对分析, 将分析结果用邻接法生成系统进化树, Bootstrap值设置为1000。利用GSDS在线工具(http:// gsds.cbi.pku.edu.cn/)制作谷子F-box家族基因外显子- 内含子结构示意图。

1.4 染色体定位与启动子顺式作用元件分析

通过MapDraw程序显示每个基因在染色体上的位置, 得到各个基因在染色体上分布状况。采用Yang等^{[ 33]}的方法, 判别同源基因, 即2个基因联配部分序列的长度要覆盖2个基因中较长序列的80%和联配部分序列的相似性大于70%。利用DNAMAN分析谷子中F-box家族基因中的重复片段, 并将分析结果标注在染色体上。从谷子基因组数据库 Phytozome (http://www.phytozome.net/search.php)中选取谷子F-box基因起始密码子上游2000 bp序列作为启动子区, 利用植物顺式作用元件数据库PLACE26.0 (http://www.dna.affrc.go.jp/PLACE/)分析基因的顺式作用元件。

1.5 基因的扩增

根据预测的谷子 SiF-box18基因的CDS序列, 用Primer Premier 5.0设计引物F: 5′-ATG GCG GAG CGG AAG-3′和R: 5′-TCA CCA TAC TGG GAA CAA AGG-3′。用TRIzol试剂盒(TIANGEN, 北京)提取叶片总RNA, 用Primer Script (TaKaRa)反转录试剂盒获得cDNA为模板扩增 SiF-box18, 并将其连接到pEASY-Blunt载体上。

1.6 RNA提取和Real-time PCR分析

对生长3周的谷子幼苗分别进行干旱(6% PEG)、NaCl (100 mmol L^-1)、茉莉酸(JA, 100 μmol L^-1)、脱落酸(ABA, 100 μmol L^-1)、赤霉素(GA, 100 μmol L^-1)和水杨酸(SA, 100 μmol L^-1)处理, 并于0、和24 h分别取样。利用TRIzol试剂盒(TIANGEN, 北京)提取总RNA, 分别利用6种胁迫下的谷子总RNA作为模板, 以SYBR Green染料法进行荧光定量Real-time PCR。用实时荧光定量PCR仪(ABI Prism 7300)进行PCR扩增。反应体系包括2× TaqPCR Master Mix (含荧光染料) 9 μL、10 μmol L^-1引物各0.5 μL、ddH₂O 9 μL和cDNA模板1 μL。反应条件为95℃ 5 min; 95℃ 15 s, 56℃ 20 s, 72℃ 30 s, 60℃ 30 s, 35个循环。PCR引物序列为F: 5′-GGA TGG AAA ACG GAG TCG-3′和R: 5′-TCA GGG TGA AAG TAG CGC-3′。以谷子 Actin基因( Si001873)为内标, 引物序列为Actin-F: 5′-GGC AAA CAG GGA GAA GAT GA-3′和Actin-R: 5′-GAG GTT GTC GGT AAG GTC ACG-3′。用2^-∆∆Ct法计算各基因表达量。

1.7 亚细胞定位

采用In-Fusion (TaKaRa)单酶切技术( BamH I)构建SiF-box18::hGFP融合表达蛋白(F: 5′-TAT CTC TAG AGG ATC CAT GGC GGA GCG GAA G -3′, R: 5′-TGC TCA CCA TGG ATC CCC ATA CTG GGA ACA AAG G-3′)。取4周左右的拟南芥叶片切成细条, 放入15 mL新鲜酶解液中, 避光, 23℃抽真空酶解3 h, 制备原生质体。取融合表达的重组质粒转化原生质体^{[ 34]}。12~16 h后于激光共焦显微镜下观察。

2 结果与分析

2.1 F-box家族成员识别及命名

通过NCBI搜索, 谷子F-box基因家族有525个成员。通过谷子干旱转录组测序, 发现有19个F-box基因上调表达。用Pfam进行结构域分析发现这19个基因都具有完整的F-box结构域(图1), 并将这19个基因命名为 SiF-box1~ SiF-box19。这些F-box家族成员中蛋白最长有767个氨基酸, 最短的有155个氨基酸, 其等电点范围在4.90~11.48之间(表1)。

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	图1 谷子19个F-box家族蛋白保守域Fig. 1 Conserved domains of F-box in 19 proteins of foxtail millet

表1 谷子F-box家族基因的基本信息 Table 1 Basic information of F-box family genes in foxtail millet

2.2 谷子F-box家族的基因结构及蛋白结构域分析

通过Pfam结构域分析显示, 19个谷子F-box蛋白家族均含有F-box保守域(图1)。每个F-box蛋白的N端均有一段保守的F-box基序, 而C端包含不同的蛋白-蛋白相互作用结构域(图2)。根据图中显示, 19个F-box蛋白中, 26.3%的蛋白只含有F-box结构域, 其他蛋白含有多个功能域, 包括Fibrinogen- like domain (FBD)、WD40 repeats (WD40)、F-box associated (FBA)、Zinc finger domain (ZnF)、Kelch repeats (Kelch)和Leucine-rich repeat (LRR)。例如, SiF-box1除了含有F-box基序外, 还包括FBA结构域; SiF-box16含有F-box、LRR和ZnF 3个结构域。有些蛋白不只包含1个F-box基序, SiF-box9与SiF- box13分别包含2个F-box基序, 而SiF-box15中有3个F-box基序。

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	图2 谷子F-box家族蛋白的保守结构域Fig. 2 Conserved domain organization of F-box proteins

2.3 进化树与基因结构分析

对谷子19个F-box基因进行多重序列比对和系统进化分析后, 19个F-box基因划分为6类(图3)。内含子-外显子结构分析显示, 同一亚族的基因具有相似内含子-外显子结构。第I类包含 SiF-box6、SiF-box7、SiF-box2、SiF-box3和 SiF-box4, 这5个基因中, 没有或只包含1个内含子; 第II类中, SiF-box13和 SiF-box14含有3个内含子, 都有FBA结构; 第III类包含 SiF-box5、 SiF-box12、 SiF-box17和 SiF-box19, 其中, SiF-box5和 SiF-box12都含有kelch repeats; 第IV类只包含 SiF-box1一个基因, 没有内含子, 包含FBA结构; 第V类中, SiF-box8、 SiF-box9、 SiF-box15和 SiF-box16都包含F-box、LRR和ZnF结构域, 除 SiF-box8外, 其他3个基因都含有2个或2个以上内含子; 第VI类包含 SiF-box11与 SiF-box18, 只含有F-box基序, 没有其他结构域。

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	图3 谷子F-box基因家族进化树以及内含子外显子结构Fig. 3 Phylogenetic tree and intron-exon structures of F-box genes in foxtail millet

2.4 染色体分布

根据基因位置信息绘制了19个F-box家族基因在谷子染色体上的定位图(图4)。19个基因分布在8条谷子染色体上, 其中第8染色体上不含谷子F-box家族基因, 第2染色体上含F-box家族基因最多, 第1和第9染色体上都有3个F-box家族基因, 其他各染色体上含1~2个F-box家族基因。

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	图4 谷子F-box家族基因的染色体分布Fig. 4 Chromosome distribution of F-box genes in foxtail millet genome

为了确定可能的进化机制, 研究了基因的复制现象, 依据系统发育树及序列相似性, 发现了4对旁系同源基因, 分别为 SiF-box10与 SiF-box14, SiF-box15与 SiF-box16, SiF-box17与 SiF-box19, 以及 SiF-box11与 Si-Fbox18基因。

2.5 启动子顺式作用元件分析

目前发现的与干旱胁迫相关的元件主要有ABRE (ABA-responsive element)、DRE (dehydration- responsive element)、LTRE (low temperature responsive element)、MYB和MYC元件。本研究选择起始密码子上游2000 bp序列作为启动子区, 研究19个F-box家族基因启动子区元件。结果显示, 谷子F-box家族基因启动子区域均含多个MYB和MYC元件, 其中, MYB数量最少为9个, 最多为78个, MYC的数量12~40个。MYB和MYC都属于转录因子, 与干旱、ABA的诱导表达相关。此外, 在这19个F-box家族基因中, 除 SiF-box12外, 在启动子区均含ABRE作用元件; 63.2%含DRE作用元件; 84.2%含LTRE作用元件(表2), 说明F-box家族基因与植物的逆境胁迫应答具有很强的关联。

表2 谷子F-box家族基因启动子逆境相关顺式作用元件的数量分布 Table 2 Distribution of cis-acting elements in foxtail millet F-box gene promoters

2.6 谷子F-box转录组结果分析

19个上调表达的谷子 F-box基因中, 表达量差异明显(图5), SiF-box1、 SiF-box4、SiF-box6、SiF- box10、SiF-box12、SiF-box17和 SiF-box19的相对表达量介于1~5倍; SiF-box2、SiF-box3、SiF-box5、SiF- box7、SiF-box8、 SiF-box9、SiF-box11、SiF-box13、SiF- box14、SiF-box15、SiF-box16、SiF-box17、SiF-box19基因的表达量介于5~10倍之间; 高于10倍的只有 SiF-box18基因。与对照相比 SiF-box18基因的表达倍数是16倍, 因此选择 Si-Fbox18基因进一步分析。

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	图5 谷子F-box基因转录组测序结果分析Fig. 5 Transcriptome analysis of F-box genes in foxtail millet1-19: SiF-box1, …, SiF-box19.

2.7 SiF-box18基因在各种逆境胁迫下的表达模式

为了进一步解析非生物胁迫和外源激素对 SiF-box18基因表达的影响, 对谷子幼苗分别进行干旱、NaCl、茉莉酸(JA)、脱落酸(ABA)、赤霉素(GA)和水杨酸(SA)胁迫处理, 通过荧光定量PCR检测不同处理条件下 SiF-box18的表达量。如图6所示, 在不同处理条件下, SiF-box18基因的表达量在24 h均达到最高。干旱胁迫下的表达倍数最高达到42倍, GA处理条件下表达量最低, 其他胁迫条件下的表达倍数, NaCl为18倍, ABA为5倍, JA与SA均为10倍。

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	图6 SiF-box18表达模式Fig. 6 Expression patterns of the SiF-box18 gene under various treatments

2.8 亚细胞定位

将SiF-box18构建到荧光表达蛋白GFP的N端。利用原生质体转化技术, 将SiF-box18-hGFP融合表达载体和空载体分别转入拟南芥原生质体, 在激光共聚焦显微镜下观察。绿色荧光蛋白在拟南芥原生质体中获得高效瞬时表达, 可以观察到对照GFP蛋白在细胞核、细胞膜、细胞质中均有绿色荧光信号; 而SiF-box18融合蛋白只在细胞核中有绿色荧光信号, 表明SiF-box18蛋白定位在细胞核中(图7)。

	Figure Option View Download New Window
	图7 SiF-box18蛋白的亚细胞定位A: 对照GFP定位在整个细胞; B: GFP-SiF-box18定位在细胞核中。Fig. 7 Subcellular localization of SiF-box18 proteinA: control GFP is distributed in the cell; B: GFP-SiF-box18 is localizated on the nuclear.

3 讨论

F-box蛋白是一个庞大的蛋白家族, 广泛存在于真核生物中。酵母、果蝇、人类、老鼠中分别有11、22、68和74个F-box蛋白成员; 而线虫中包含326个F-box蛋白家族成员^{[ 35]}。植物中有大量的F-box蛋白存在, 拟南芥中大约有600~700个F-box基因^{[ 36, 37]}, 水稻中包含687个F-box基因^{[ 38]}; 最近研究表明, 玉米中有359个F-box蛋白成员^{[ 39]}。根据C端蛋白-蛋白相互作用结构域的不同, 可以把拟南芥F-box家族分为19个亚族^{[ 36, 37]}; 水稻F-box家族分为10个亚族^{[ 38]}; 玉米中分为13个亚族^{[ 39]}。谷子F-box家族通过NCBI搜索, 发现有525个F-box基因, 进一步研究了其中19个受干旱诱导表达的F-box基因。

除N端保守的F-box基序外, 谷子F-box蛋白C端含有不同的蛋白-蛋白相互作用结构域, 例如, FBD、FBA、LRR、WD40、PPR和ZnF。本研究根据序列相似性, 将响应干旱胁迫的谷子19个F-box蛋白分为6类(图3)。第II类中, SiF-box13和SiF-box14包含FBA结构, FBA在泛素化中起重要作用, 可以识别靶蛋白; 第III类中SiF-box5和SiF-box12都含有Kelch repeats, Kelch是植物F-box蛋白特有的结构域^{[ 38]}; 第IV只有SiF-box1一个蛋白, 包含FBA结构; 第V类中, 4个成员都包含F-box、LRR和FBD结构域, LRRs是Leu重复单元, 含有LRRs结构的蛋白一般与信号传导, DNA修复、重组、转录以及抗病等功能相关^{[ 40]}; 第VI类含有F-box基序。此外, SiF-box6和SiF-box15蛋白还包含ZnF结构; 在谷子19个F-box家族中, 3个蛋白包含WD40结构, WD40结构存在于所有真核生物中, 在信号传导、细胞周期调控、细胞凋亡方面起着重要作用^{[ 41]}。说明受干旱诱导表达的谷子F-box蛋白可能同样发挥着调控细胞周期、细胞信号传导等功能。

基因重复是基因组进化及基因家族扩大的重要因素。基因重复后会产生3种结果: (1)基因功能的缺失, 基因的2个拷贝中的一个拷贝由于突变失去功能; (2)产生新的功能, 基因的2个拷贝中的一个拷贝获得新的功能并通过自然选择被保留下来; (3)基因功能互补, 基因的2个拷贝都发生退化突变, 使得2个拷贝功能产生互补作用。本研究发现19个F-box基因有4对旁系同源基因, 这4对基因位于不同的染色体上, 可能是复制后功能产生了分化。

据报道, F-box基因不仅参与信号传导等生命活动^{[ 42]}, 还介导干旱等非生物胁迫响应。例如, 干旱条件下, dor转基因拟南芥植株表现出比野生型更强的抗性, 证明 DOR对干旱胁迫起负调控作用^{[ 30]}; 在渗透胁迫和高盐胁迫下, DEL3 (EID₁-like protein3)高效、快速表达^{[ 43]}。本研究发现19个受干旱诱导表达的谷子F-box基因, 启动子区都包含多种非生物逆境相关元件, 而MYB和MYC的数量远远大于其他元件的数量, 说明F-box基因对干旱胁迫的响应主要通过MYB和MYC转录因子调控。

植物对干旱的响应是一个复杂的过程, 受不同分子和细胞通路的调控。根据干旱胁迫信号转导过程中, 胁迫相关基因的表达是否依赖ABA, 存在两种途径, 其中, ABA依赖型途径通过ABRE结合bZIP转录因子或者通过结合MYB/MYC转录因子激活下游基因的表达; 非ABA依赖型途径则由信号激活传感蛋白, 传递给DREB后结合DRE元件启动下游抗逆基因的表达。研究表明, 与干旱胁迫相关的F-box基因家族基因, 对ABA有不同程度的响应, 例如, ABA处理条件下, DOR表达量降低, 对ABA起负调作用^{[ 30]}; 与对照相比, DEL3的表达量上升, edl3-1表达量下降^{[ 43]}。在ABA胁迫下, SiF-box18基因表达量在24 h达到最高, 是对照的5倍(图6)。推测 SiF-box18基因可能通过依赖ABA途径参与谷子的干旱胁迫。

在干旱和高盐胁迫下, SiF-box18基因的表达量在24 h达到最高, 分别是对照的42倍和18倍(图6)。在植物外源激素GA、SA、JA、ABA处理下, SiF-box18基因表达量均有不同程度的增加(图6), 说明 SiF- box18基因可能参与了GA、SA、JA等植物激素的信号传导过程。 SiF-box18基因的启动子区包含1个ABRE、4个DRE、3个LTRE、45个MYB和14个MYC等多种非生物胁迫响应元件。说明 Si-Fbox18基因可能参与干旱、低温等多种非生物胁迫响应。

4 结论

19个上调表达的谷子F-box基因位于8条染色体上, 其中SiF-box18蛋白被定位在细胞核中, 对干旱、高盐和ABA等都有响应, 推测 SiF-box18基因可能在非生物胁迫响应中起重要的作用。

附表

植物中已知功能的F-box蛋白

Supplemental Table F-box proteins with known function in plants

F-box protein	Gene identifier		Organism	Function	Reference
COI1	At2g39940	Coronatine Insensitive 1	Arabidopsis	Jasmonate-Regulated defense responses, pollen fertility	[ 1] [ 2] [ 3]
ORE9/ MAX2	At2g42620	Oresara 9/ More Axillary 2	Arabidopsis	Regulation of leaf senescence, selective repression of axillary shoots	[ 4] [ 5] [ 6]
TIR1	At3g62980	Transport Inhibitor Response 1	Arabidopsis	Auxin receptor	[ 7] [ 8] [ 9] [ 10]
AFB1/GRH1	At4g03190	Auxin signaling F Box protein 1	Arabidopsis	Auxin receptor, plant growth and development	[ 11] [ 12] [ 13]
AFB2	At3g26810	Auxin signaling F Box protein 2	Arabidopsis	Auxin receptor, plant growth and development, glucose repression	[ 11] [ 12] [ 13]
AFB3	At1g12820	Auxin signaling F Box protein 3	Arabidopsis	Auxin receptor, plant growth and development	[ 11] [ 12] [ 13]
AFB4	At4g24390	Auxin-signaling F-box protein 4	Arabidopsis	Plant growth, development and innate immunity	[ 14]
SON1	At2g17310	Suppressor Of nim1-1	Arabidopsis	Defense responses	[ 15]
UFO	At1g30950	Unusual Floral Organs	Arabidopsis	Determination of floral meristem and floral organ identity	[ 16] [ 17] [ 18] [ 19]
EID1	At4g02440	Empfindlicher Im Dunkelroten Licht 1	Arabidopsis	Negative regulator in phytochrome A-specific light signaling	[ 20] [ 21]
AFR	At2g24540	Attenuated Far-Red Response	Arabidopsis	Positive regulator of phytochrome A-mediated light signaling	[ 22]
EBF1	At2g25490	EIN3-Binding F box protein 1	Arabidopsis	Negative regulator of ethylene-signaling, plant growth	[ 23] [ 24]
EBF2	At5g25350	EIN3-Binding F box protein 2	Arabidopsis	Negative regulator of ethylene-signaling, plant growth	[ 23] [ 24]
ZTL/LKP1	At5g57360	Zeitlupe	Arabidopsis	Circadian rhythm, photomorphogenesis and flowering time	[ 25] [ 26]
LKP2/FKL/ FKF1	At2g18915	Lov Kelch Protein 2	Arabidopsis	Circadian clock regulation and flowering time control	[ 27] [ 28] [ 29] [ 30] [ 31]
SLY1	At4g24210	Sleepy 1	Arabidopsis	Positive regulator of Gibberellic acid signaling	[ 32] [ 33]
SNE	At5g48170	Sneezy	Arabidopsis	Positive regulator of Gibberellic acid signaling	[ 34] [ 35]
CEG (AtSFL61)	At3g22650	Cegenduo	Arabidopsis	Negative regulator of auxin-mediated lateral root formation	[ 36]
DOR	At2g31470	Drought Tolerance Repressor 1	Arabidopsis	Negative regulate drought stress	[ 37]
CFK1	At5g42350	COP9 INTERACTING F-BOX KELCH 1	Arabidopsis	Controlling hypocotyl elongation	[ 38]
FOA1	AT3G17320	F-box overexpressed/ oppressed ABA signaling	Arabidopsis	Play a negative role in ABA signaling.	[ 39]
EDL3	At3g63070	EID1-like protein 3	Arabidopsis	Negative acting factors in ABA nsignalling to degradation via the proteasome.	[ 40]
GID2	AB100246	Gibberellin Insensitive Dwarf 2	Oryza sativa	Positive regulator of Gibberellic acid signaling	[ 41] [ 42]
D3	AK065478	Dwarf 3	O. sativa	Control of tiller bud activity	[ 43] [ 44]
FBK12	OS03g03570	F-box protein containing a kelch repeat motif	O. sativa	Delay in leaf senescence and germination, and increased seed size	[ 45]
DDF1		Dwarf and deformed flower 1	O. sativa	Regulate vegetative growth and floral organ specification	[ 46]
MAIF1	Os02g44990	miRNAs regulated and abiotic stress induced F-box gene	O. sativa	Plays the negative role in response to abiotic stress	[ 47]
FIM	S71192	Fimbriata	Antirrhinum majus	Floral differentiation	[ 48] [ 49]
AhSLF-S2	AJ297974	Self-incompatibility Locus-encoded F-box protein	A. hispanicum	Controls pollen function of S-RNase-based self-incompatibility	[ 50] [ 51] [ 52]
SFB3	AY571665	S haplotype-specific F-box protein	Prunus avium	Male determinant of gametophytic self incompatibility	[ 53] [ 54] [ 55]

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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2006

6.582

0.0

Verslues

P E

, Agarwal

, Katiyar

A S

, Zhu

J K

. Methods and concepts in quantifying resistance to drought, salt and freezing, abiotic stresses that affect plant water status. Plant J, 2006, 45: 523-539

Institute for Integrative Genome Biology and Department of Botany and Plant Sciences, University of California, Riverside, CA 92521, USA * *(fax 951 827 7115; e-mail jian-kang.zhu@ucr.edu ).

> The abiotic stresses of drought, salinity and freezing are linked by the fact that they all decrease the availability of water to plant cells. This decreased availability of water is quantified as a decrease in water potential. Plants resist low water potential and related stresses by modifying water uptake and loss to avoid low water potential, accumulating solutes and modifying the properties of cell walls to avoid the dehydration induced by low water potential and using protective proteins and mechanisms to tolerate reduced water content by preventing or repairing cell damage. Salt stress also alters plant ion homeostasis, and under many conditions this may be the predominant factor affecting plant performance. Our emphasis is on experiments that quantify resistance to realistic and reproducible low water potential (drought), salt and freezing stresses while being suitable for genetic studies where a large number of lines must be analyzed. Detailed protocols for the use of polyethylene glycol-infused agar plates to impose low water potential stress, assay of salt tolerance based on root elongation, quantification of freezing tolerance and the use of electrolyte leakage experiments to quantify cellular damage induced by freezing and low water potential are also presented.

... 在适应干旱等非生物逆境胁迫中, 植物不断进化自身防御系统以减少水分损失或增加水分吸收等^[1] ...

2012

6.736

0.0

Sadanand om

, Bailey

, Ewan

, Lee

, Nelis

. The ubiquitin-proteasome system: central modifier of plant signalling. New Phytol, 2012, 196: 13-28

1 School of Biological and Biomedical Sciences, Durham University, Durham, UK 2 The Scottish Institute for Cell Signalling (SCILLS), Sir James Black Centre, University of Dundee, Dundee, UK * Author for correspondence: Ari Sadanandom Tel: +44 191 334 1263 Email: ari.sadanandom@durham.ac.uk

Summary 13 I. Brief history 13 II. Components of the ubiquitin–proteasome system 14 III. Ubiquitin-mediated degradation: a recurrent theme in the plant life cycle 18 IV. Conclusion and future prospects 25 Acknowledgements 25 References 25

... 蛋白质的降解在植物逆境信号的响应、细胞信号转导等过程中发挥重要作用, 对于维持植物正常生长发育具有重要意义^[2] ...

2000

11.721

0.0

... 目前已经发现不同类型的E3^[3], 主要的一种是Skp1-Cullin1-F-box (SCF)复合体, 由4个亚基组成, 分别为Cullin、SKP1、RBX1和F-box蛋白^[4,5] ...

2004

0.0

Cardozo

, Pagano

. The SCF ubiquitin ligase: insights into a molecular machine. Nat Rev Mol Cell Biol, 2004, 5: 739-751

... 目前已经发现不同类型的E3^[3], 主要的一种是Skp1-Cullin1-F-box (SCF)复合体, 由4个亚基组成, 分别为Cullin、SKP1、RBX1和F-box蛋白^[4,5] ...

2006

2.458

0.0

M S

, Tsai

P I

, Chien

C T

. F-box proteins: the key to protein degradation. J Biomed Sci, 2006, 13: 181-191

The eukaryotic protein degradation pathway involves the ubiquitin (Ub) modification of substrates targeted for degradation by the 26S proteasome. The addition of Ub, a process called ubiquitination, is mediated by enzymes including the E3 Ub ligases which transfer the Ub to targeted substrates. A major type of E3 Ub ligases, the SCF (Skp–Cullin–F-box) complex, is composed of four major components: Skp1, Cul1/Cdc53, Roc1/Rbx1/Hrt1, and an F-box protein. The F-box component of the SCF machineries is responsible for recognizing different substrates for ubiquitination. Interaction with components of the SCF complex is mediated through the F-box motif of the F-box protein while it associates with phosphorylated substrates through its second protein–protein interaction motif such as Trp–Asp (WD) repeats or leucine-rich repeats (LRRs). By targeting diverse substrates, F-box proteins exert controls over stability of proteins and regulate the mechanisms for a wide-range of cellular processes. Here we discuss the importance of F-box proteins by providing a general overview and examples of how F-box proteins function in various cellular settings such as tissue development, cell proliferation, and cell death, in the modeling organism Drosophila .

1.Academia Sinica Institute of Molecular Biology 115 Taipei Taiwan 2.National Taiwan University Institute of Molecular Medicine, Medical College 100 Taipei Taiwan

... 目前已经发现不同类型的E3^[3], 主要的一种是Skp1-Cullin1-F-box (SCF)复合体, 由4个亚基组成, 分别为Cullin、SKP1、RBX1和F-box蛋白^[4,5] ...

1994

9.822

0.0

... F-box蛋白最早在cyclin F的N端被发现而命名^[6], 广泛存在于真核生物中, 参与多种植物激素信号的传导、逆境胁迫应答等, 在调节光形态建成、光周期节律、花器官发育及自交不亲和中发挥着重要功能 ...

1995

9.251

0.0

... 植物中发现的第一个F-box蛋白是Unusual Floral Organs (UFO), 参与了植物花分生组织与花器官的建成^[7,8,9,10] ...

2006

3.347

0.0

Hepworth

S R

, Klenz

J E

, Haughn

G W

. UFO in the Arabidopsis inflorescence apex is required for floral-meristem identity and bract suppression. Planta, 2006, 223: 769-778

The UNUSUAL FLORAL ORGANS ( UFO ) gene of Arabidopsis encodes an F-box protein required for the determination of floral-organ and floral-meristem identity. Mutation of UFO leads to dramatic changes in floral-organ type which are well-characterized whereas inflorescence defects are more subtle and less understood. These defects include an increase in the number of secondary inflorescences, nodes that alternate between forming flowers and secondary inflorescences, and nodes in which a single flower is subtended by a bract. Here, we show how inflorescence defects correlate with the abnormal development of floral primordia and establish a temporal requirement for UFO in this process. At the inflorescence apex of ufo mutants, newly formed primordia are initially bract-like. Expression of the floral-meristem identity genes LFY and AP1 are confined to a relatively small adaxial region of these primordia with expression of the bract-identity marker FIL observed in cells that comprise the balance of the primordia. Proliferation of cells in the adaxial region of these early primordia is delayed by several nodes such that primordia appear “chimeric” at several nodes, having visible floral and bract components. However, by late stage 2 of floral development, growth of the bract generally ceases and is overtaken by development of the floral primordium. This abnormal pattern of floral meristem development is not rescued by expression of UFO from the AP1 promoter, indicating that UFO is required prior to AP1 activation for normal development of floral primordia. We propose that UFO and LFY are jointly required in the inflorescence meristem to both promote floral meristem development and inhibit, in a non-cell autonomous manner, growth of the bract.

1.University of British Columbia Department of Botany Vancouver British Columbia V6T 1Z4 Canada Vancouver British Columbia V6T 1Z4 Canada

... 植物中发现的第一个F-box蛋白是Unusual Floral Organs (UFO), 参与了植物花分生组织与花器官的建成^[7,8,9,10] ...

2012

6.582

0.0

Sasaki

, Yamaguchi

, Aida

, Shikata

, Abe

, Ohtsubo

. Mutation in Torenia fournieri Lind: UFO homolog confers loss of TfLFY interaction and results in a petal to sepal transformation. Plant J, 2012, 71: 1002-1014

1 National Institute of Floricultural Science, Tsukuba, Ibaraki 305-8519, Japan 2 RIKEN Nishina Center, 2-1 Hirosawa, Wako, Saitama 351-0198, Japan * (e-mail nohtsubo@affrc.go.jp ).

> We identified a Torenia fournieri Lind. mutant (no. 252) that exhibited a sepaloid phenotype in which the second whorls were changed to sepal-like organs. This mutant had no stamens, and the floral organs consisted of sepals and carpels. Although the expression of a torenia class B MADS-box gene, GLOBOSA ( TfGLO ), was abolished in the 252 mutant, no mutation of TfGLO was found. Among torenia homologs such as APETALA1 ( AP1 ), LEAFY ( LFY ), and UNUSUAL FLORAL ORGANS ( UFO ), which regulate expression of class B genes in Arabidopsis, only accumulation of the TfUFO transcript was diminished in the 252 mutant. Furthermore, a missense mutation was found in the coding region of the mutant TfUFO . Intact TfUFO complemented the mutant phenotype whereas mutated TfUFO did not; in addition, the transgenic phenotype of TfUFO -knockdown torenias coincided with the mutant phenotype. Yeast two-hybrid analysis revealed that the mutated TfUFO lost its ability to interact with TfLFY protein. In situ hybridization analysis indicated that the transcripts of TfUFO and TfLFY were partially accumulated in the same region. These results clearly demonstrate that the defect in TfUFO caused the sepaloid phenotype in the 252 mutant due to the loss of interaction with TfLFY.

... 植物中发现的第一个F-box蛋白是Unusual Floral Organs (UFO), 参与了植物花分生组织与花器官的建成^[7,8,9,10] ...

2013

0.0

Wang

, Chen

, Weng

, Li

, Cao

, Hu

, Luo

, Yang

. Multiple components are integrated to determine leaf complexity inLotus japonicus. J Integr Plant Biol, 2013, 55: 419-433

1 National Key Laboratory of Plant Molecular Genetics, Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Graduate School of the Chinese Academy of Sciences, the Chinese Academy of Sciences, Shanghai 20032, China 2 State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China * Corresponding author Tel: +86 21 5492 4103; Fax: +86 21 5492 4015; E-mail: dluo@sibs.ac.cn Tel: +86 21 5492 4112; Fax: +86 21 5492 4015; E-mail: jyang@sibs.ac.cn

> Transcription factors and phytohormones have been reported to play crucial roles to regulate leaf complexity among plant species. Using the compound-leafed species Lotus japonicus , a model legume plant with five visible leaflets, we characterized four independent mutants with reduced leaf complexity, proliferating floral meristem (pfm) , proliferating floral organ-2 (pfo-2) , fused leaflets1 (ful1) and umbrella leaflets (uml) , which were further identified as loss-of-function mutants of Arabidopsis orthologs LEAFY ( LFY ), UNUSUAL FLORAL ORGANS ( UFO ), CUP-SHAPED COTYLEDON 2 ( CUC2 ) and PIN-FORMED 1 ( PIN1 ), respectively. Comparing the leaf development of wild-type and mutants by a scanning electron microscopy approach, leaflet initiation and/or dissection were found to be affected in these mutants. Expression and phenotype analysis indicated that PFM/LjLFY and PFO/LjUFO determined the basipetal leaflet initiation manner in L. japonicus . Genetic analysis of ful1 and uml mutants and their double mutants revealed that the CUC2-like gene and auxin pathway also participated in leaflet dissection in L. japonicus , and their functions might influence cytokinin biogenesis directly or indirectly. Our results here suggest that multiple genes were interplayed and played conserved functions in controlling leaf complexity during compound leaf development in L. japonicus .

... 植物中发现的第一个F-box蛋白是Unusual Floral Organs (UFO), 参与了植物花分生组织与花器官的建成^[7,8,9,10] ...

2000

31.957

0.0

... LKP1/ZTL、LKP2/FKL和FKF1可以调控光形态建成、调节生物钟的昼夜周期节律及开花时间^{[11,12,13,14]} ...

2005

0.0

Imaizumi

, Schultz

T F

, Harmon

F G

, Ho

L A

, Kay

S A

. FKF1F-box protein mediates cyclic degradation of a repressor of CONSTANS in Arabidopsis. Science, 2005, 309: 293-297

Department of Cell Biology, The Scripps Research Institute, 10550 North Torrey Pines Road, La Jolla, CA 92037, USA. ↵ * To whom correspondence should be addressed. E-mail: stevek@scripps.edu

... LKP1/ZTL、LKP2/FKL和FKF1可以调控光形态建成、调节生物钟的昼夜周期节律及开花时间^{[11,12,13,14]} ...

2011

6.582

0.0

Tomoyuki

, Yuuki

, Haruna

, Yasunobu

, Yuji

, Yumiko

, Tomohiro

. LOV KELCH PROTEIN2 and ZEITLUPE repress Arabidopsis photoperiodic flowering under non-inductive conditions, dependent on FLAVIN-BINDING KELCH REPEAT F-BOX1. Plant J, 2011, 67: 608-621

1 Department of Life Science, Faculty of Science, Gakushuin University, 1-5-1 Mejiro, Toshima-ku, Tokyo 171-8588, Japan 2 Division of Gene Research, Life Science Research Center, Kagawa University, 2393 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0795, Japan * (fax +81 3 5992 1029; e-mail tomohiro.kiyosue@gakushuin.ac.jp ).

> LOV KELCH PROTEIN2 (LKP2), ZEITLUPE (ZTL)/LOV KELCH PROTEIN1 (LKP1) and FLAVIN-BINDING KELCH REPEAT F-BOX1 (FKF1) constitute a family of Arabidopsis F-box proteins that regulate the circadian clock. Over-expression of LKP2 or ZTL causes arrhythmicity of multiple clock outputs under constant light and in constant darkness. Here, we show the significance of LKP2 and ZTL in the photoperiodic control of flowering time in Arabidopsis. In plants over-expressing LKP2 , CO and FT expression was down-regulated under long-day conditions. LKP2 and ZTL physically interacted with FKF1, which was recruited from the nucleus into cytosolic speckles. LKP2 and ZTL inhibited the interaction of FKF1 with CYCLING DOF FACTOR 1, a ubiquitination substrate for FKF1 that is localized in the nucleus. The Kelch repeat regions of LKP2 and ZTL were sufficient for their physical interaction with FKF1 and translocation of FKF1 to the cytoplasm. Over-expression of LKP2 Kelch repeats induced late flowering under long-day conditions. lkp2 ztl double mutant plants flowered earlier than wild-type plants under short-day (non-inductive) conditions, and both CO and FT expression levels were up-regulated in the double mutant plants. The early flowering of lkp2 ztl was dependent on FKF1 . LKP2 , ZTL or both affected the accumulation of FKF1 protein during the early light period. These results indicate that an important role of LKP2 and ZTL in the photoperiodic pathway is repression of flowering under non-inductive conditions, and this is dependent on FKF1 .

... LKP1/ZTL、LKP2/FKL和FKF1可以调控光形态建成、调节生物钟的昼夜周期节律及开花时间^{[11,12,13,14]} ...

2013

4.134

0.0

... LKP1/ZTL、LKP2/FKL和FKF1可以调控光形态建成、调节生物钟的昼夜周期节律及开花时间^{[11,12,13,14]} ...

2002

3.518

0.0

Lai

, Ma

, Han

, Liang

, Zhang

, Hong

, Xue

. An F-box gene linked to the self-incompatibility locus of Antirrhinum is expressed specifically in pollen and tapetum. Plant Mol Biol, 2002, 50: 29-42

In many flowering plants, self-fertilization is prevented by an intraspecific reproductive barrier known as self-incompatibility (SI), that, in most cases, is controlled by a single multiallelic S locus. So far, the only known S locus product in self-incompatible species from the Solanaceae, Scrophulariaceae and Rosaceae is a class of ribonucleases called S RNases. Molecular and transgenic analyses have shown that S RNases are responsible for pollen rejection by the pistil but have no role in pollen expression of SI, which appears to be mediated by a gene called the pollen self-incompatibility or Sp gene. To identify possible candidates for this gene, we investigated the genomic structure of the S locus in Antirrhinum , a member of the Scrophulariaceae. A novel F-box gene, AhSLF - S 2 , encoded by the S 2 allele, with the expected features of the Sp gene was identified. AhSLF - S 2 is located 9 kb downstream of S 2 RNase gene and encodes a polypeptide of 376 amino acids with a conserved F-box domain in its amino-terminal part. Hypothetical genes homologous to AhSLF - S 2 are apparent in the sequenced genomic DNA of Arabidopsis and rice. Together, they define a large gene family, named SLF ( S locus F-box) family. AhSLF - S 2 is highly polymorphic and is specifically expressed in tapetum, microspores and pollen grains in an allele-specific manner. The possibility that Sp encodes an F-box protein and the implications of this for the operation of self-incompatibility are discussed.

1.Institute of Genetics and Developmental Biology 2.National Center for Gene Research, Chinese Academy of Sciences

... F-box蛋白还与植物的自交不亲和性相关, 许多植物的自交不亲和性由S位点控制, 研究表明, 与S位点相关的S-locus-F-box genes (SLF)能在花粉中特异表达^{[15,16,17,18]} ...

2004

6.582

0.0

Ushijima

, Yamane

, Watari

, Kakehi

, Ikeda

, Hauck

N R

, Iezzoni

A F

, Tao

. The S haplotype-specific F-box protein gene, SFB, is defective in self-compatible haplotypes of Prunus avium and P. mume. Plant J, 2004, 39: 573-586

1 Graduate School of Agriculture, Kyoto University, Kyoto 606-8502, Japan 2 Department of Horticulture, Michigan State University, East Lansing 48824, MI, USA * (fax +81 75 753 6497; e-mail rtao@kais.kyoto-u.ac.jp ).

> Many Prunus species, including sweet cherry and Japanese apricot, of the Rosaceae, display an S-RNase-based gametophytic self-incompatibility (GSI). The specificity of this outcrossing mechanism is determined by a minimum of two genes that are located in a multigene complex, termed the S locus, which controls the pistil and pollen specificities. SFB , a gene located in the S locus region, encodes an F-box protein that has appropriate S haplotype-specific variation to be the pollen determinant in the self-incompatibility reaction. This study characterizes SFBs of two self-compatible (SC) haplotypes, S 4′ and S f , of Prunus . S 4′ of sweet cherry is a pollen-part mutant (PPM) that was produced by X-ray irradiation, while S f of Japanese apricot is a naturally occurring SC haplotype that is considered to be a PPM. DNA sequence analysis revealed defects in both SFB 4′ and SFB f . A 4 bp deletion upstream from the HVa coding region of SFB 4′ causes a frame-shift that produces transcripts of a defective SFB lacking the two hypervariable regions, HVa and HVb. Similarly, the presence of a 6.8 kbp insertion in the middle of the SFB f coding region leads to transcripts for a defective SFB lacking the C-terminal half that contains HVa and HVb. As all reported SFBs of functional S haplotypes encode intact SFB, the fact that the partial loss-of-function mutations in SFB are present in SC mutant haplotypes of Prunus provides additional evidence that SFB is the pollen S gene in GSI in Prunus .

2010

2.07

0.0

Hidenori

, Hiroyuki

, Mai

. Pollen-expressed F-box gene family and mechanism of S-RNase-based gametophytic self- incompatibility (GSI) in Rosaceae. Sex Plant Reprod, 2010, 23: 39-43

Many species of Rosaceae, Solanaceae, and Plantaginaceae exhibit S-RNase-based self-incompatibility (SI) in which pistil-part specificity is controlled by S locus-encoded ribonuclease (S-RNase). Although recent findings revealed that S locus-encoded F-box protein, SLF/SFB, determines pollen-part specificity, how these pistil- and pollen-part S locus products interact in vivo and elicit the SI reaction is largely unclear. Furthermore, genetic studies suggested that pollen S function can differ among species. In Solanaceae and the rosaceous subfamily Maloideae (e.g., apple and pear), the coexistence of two different pollen S alleles in a pollen breaks down SI of the pollen, a phenomenon known as competitive interaction. However, competitive interaction seems not to occur in the subfamily Prunoideae (e.g., cherry and almond) of Rosaceae. Furthermore, the effect of the deletion of pollen S seems to vary among taxa. This review focuses on the potential differences in pollen-part function between subfamilies of Rosaceae, Maloideae, and Prunoideae, and discusses implications for the mechanistic divergence of the S-RNase-based SI.

1.Chiba University Graduate School of Horticulture Matsudo Chiba 271-8510 Japan

2013

3.73

0.0

, Li

. Genetic features of the spontaneous self-compatible mutant, ‘Jin Zhui’ (Pyrus bretschneideri Rehd. ). PLoS One, 2013, 8: e76509

1 Laboratory of Fruit Tree Cell and Molecular Breeding, China Agricultural University, Beijing, China 2 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing, China

‘Jin Zhui’ is a spontaneous self-compatible mutant of ‘Ya Li’ ( Pyrus bretschneideri Rehd. S 21 S 34 ), the latter displaying a typical S -RNase-based gametophytic self-incompatibility (GSI). The pollen-part mutation (PPM) of ‘Jin Zhui’ might be due to a natural mutation in the pollen -S gene ( S 34 haplotype). However, the molecular mechanisms behind these phenotypic changes are still unclear. In this study, we identified five SLF ( S-Locus F-box ) genes in ‘Ya Li’, while no nucleotide differences were found in the SLF genes of ‘Jin Zhui’. Further genetic analysis by S-RNase PCR-typing of selfed progeny of ‘Jin Zhui’ and ‘Ya Li’ × ‘Jin Zhui’ progeny showed three progeny classes ( S 21 S 21 , S 21 S 34 and S 34 S 34 ) as opposed to the two classes reported previously ( S 21 S 34 and S 34 S 34 ), indicating that the pollen gametes of ‘Jin Zhui’, bearing either the S 21 - or S 34 -haplotype, were able to overcome self-incompatibility (SI) barriers. Moreover, no evidence of pollen- S duplication was found. These findings support the hypothesis that loss of function of S -locus unlinked PPM expressed in pollen leads to SI breakdown in ‘Jin Zhui’, rather than natural mutation in the pollen -S gene ( S 34 haplotype). Furthermore, abnormal meiosis was observed in a number of pollen mother cells (PMCs) in ‘Jin Zhui’, but not in ‘Ya Li’. These and other interesting findings are discussed.

2001

38.597

0.0

Gray

W M

, Kepinski

, Rouse

, Leyser

, Estelle

. Auxin regulates SCF(TIR1)-dependent degradation of AUX/IAA proteins. Nature, 2001, 414: 271-276

Nature is the international weekly journal of science: a magazine style journal that publishes full-length research papers in all disciplines of science, as well as News and Views, reviews, news, features, commentaries, web focuses and more, covering all branches of science and how science impacts upon all aspects of society and life.

... 许多F-box蛋白还参与激素的信号传导, TIR1是生长素受体蛋白, 与AUX/IAA发生相互作用, 导致AUX/IAA降解^{[19,20,21,22]} ...

2005

38.597

0.0

Kepinski

, Leyser

. The Arabidopsis F-box protein TIR1 is an auxin receptor. Nature, 2005, 435: 446-451

... 许多F-box蛋白还参与激素的信号传导, TIR1是生长素受体蛋白, 与AUX/IAA发生相互作用, 导致AUX/IAA降解^{[19,20,21,22]} ...

2013

6.555

0.0

... 许多F-box蛋白还参与激素的信号传导, TIR1是生长素受体蛋白, 与AUX/IAA发生相互作用, 导致AUX/IAA降解^{[19,20,21,22]} ...

2013

10.015

0.0

... 许多F-box蛋白还参与激素的信号传导, TIR1是生长素受体蛋白, 与AUX/IAA发生相互作用, 导致AUX/IAA降解^{[19,20,21,22]} ...

31.957

0.0

... EBF1过量表达引起植物对乙烯不敏感, ebf1和ebf2突变株则表现出组成型乙烯反应, EBF1/EBF2两者之一发生突变都导致植株对乙烯反应增强, 而双突变体则表现出三重反应^[23,24] ...

2013

0.0

Iqbal

, Trivellini

, Masood

, Ferrante

, Khan

N A

. Current understand ing on ethylene signaling in plants: The influence of nutrient availability. Plant Physiol Biochem, 2013, 73: 128-138

2003

11.808

0.0

Itoh

, Matsuoka

, Steber

C M

. A role for the ubiquitin-26S proteasome pathway in gibberellin signaling. Trends Plant Sci, 2003, 8: 492-497

... 赤霉素(GA)的信号通路也可以通过F-box蛋白(SLY1)调节^[25,26] ...

2013

6.555

0.0

... 赤霉素(GA)的信号通路也可以通过F-box蛋白(SLY1)调节^[25,26] ...

1998

0.0

Xie

D X

, Feys

B F

, James

, Nieto-Rostro

, Turner

J G

, COI1: an Arabidopsis gene required for jasmonate-regulated defense fertility. Science, 1998, 280: 1091-1094

School of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, UK. ↵ ‡To whom correspondence should be addressed.

... COI1编码F-box蛋白, 在茉莉酸(JA)信号通路中起关键作用^[27,28,29] ...

2005

3.518

0.0

Devoto

, Ellis

, Magusin

, Chang

H S

, Chilcott

, Zhu

, Turner

J G

. Expression profiling reveals COI1 to be a key regulator of genes involved in wound-and methyl jasmonate-induced secondary metabolism, defence, and hormone interactions. Plant Mol Biol, 2005, 58: 497-513

The Arabidopsis gene COI1 is required for jasmonic acid (JA)-induced growth inhibition, resistance to insect herbivory, and resistance to pathogens. In addition, COI1 is also required for transcription of several genes induced by wounding or by JA. Here, we use microarray gene transcription profiling of wild type and coi1 mutant plants to examine the extent of the requirement of COI1 for JA-induced and wound-induced gene transcription. We show that COI1 is required for expression of approximately 84% of 212 genes induced by JA, and for expression of approximately 44% of 153 genes induced by wounding. Surprisingly, COI1 was also required for repression of 53% of 104 genes whose expression was suppressed by JA, and for repression of approximately 46% of 83 genes whose expression was suppressed by wounding. These results indicate that COI1 plays a pivotal role in wound- and JA signalling.

1.University of East Anglia School of Biological Sciences NR4 7TJ Norwich UK 2.John Innes Centre NR4 7UH Norwich UK 3.Torrey Mesa Research Institute, Syngenta Research and Technology 3115 Merryfield Row San Diego CA 92121 USA 4.Syngenta Biotechnology Inc. 3054 Cornwallis Road Research Triangle Park NC 27709 USA 5.University of North Carolina at Chapel Hill Department of Biology Chapel Hill NC 27599 USA 6.Diversa Corporation 4955 Directions Place San Diego CA 92121 USA;

... COI1编码F-box蛋白, 在茉莉酸(JA)信号通路中起关键作用^[27,28,29] ...

2013

6.555

0.0

... COI1编码F-box蛋白, 在茉莉酸(JA)信号通路中起关键作用^[27,28,29] ...

2008

6.555

0.0

... F-box蛋白参与非生物胁迫应答的调控, 如DOR1 (drought tolerance repressor 1)是S位点类F-box蛋白家族的基因, 在保卫细胞中表达, dor使ABA表达量上调, 气孔关闭, 从而增加抗旱性^[30] ...

... 例如, 干旱条件下, dor转基因拟南芥植株表现出比野生型更强的抗性, 证明DOR对干旱胁迫起负调控作用^[30] ...

... 研究表明, 与干旱胁迫相关的F-box基因家族基因, 对ABA有不同程度的响应, 例如, ABA处理条件下, DOR表达量降低, 对ABA起负调作用^[30] ...

2013

6.555

0.0

... MAX2 (more axillary growth 2)对干旱响应明显, 与野生型相比, max2的保卫细胞对ABA敏感程度降低, 角质层变薄, 在MAX2突变的情况下, 与ABA合成、运输及代谢有关的基因表达量都降低^[31] ...

2009

10.526

1.1032

Ren

, Wen

, Gao

, Jin

, Xue

, Yao

. DOG 1. 0: illustrator of protein domain structures cell research. Cell Res, 2009, 19: 271-273

Cell death and differentiation is a monthly research journal focused on the exciting field of programmed cell death and apoptosis. It provides a single accessible source of information for both scientists and clinicians, keeping them up-to-date with advances in the field. It encompasses programmed cell death, cell death induced by toxic agents, differentiation and the interrelation of these with cell proliferation.

... 0^[32]绘制19个F-box家族蛋白的功能结构域示意图 ...

2008

2.881

0.0

Yang

, Zhang

, Yue

J X

, Tian

, Chen

J Q

. Recent duplications dominate NBS-encoding gene expansion in two woody species. Mol Genet Genomics, 2008, 280: 187-198

Most disease resistance genes in plants encode NBS-LRR proteins. However, in woody species, little is known about the evolutionary history of these genes. Here, we identified 459 and 330 respective NBS-LRRs in grapevine and poplar genomes. We subsequently investigated protein motif composition, phylogenetic relationships and physical locations. We found significant excesses of recent duplications in perennial species, compared with those of annuals, represented by rice and Arabidopsis . Consequently, we observed higher nucleotide identity among paralogs and a higher percentage of NBS-encoding genes positioned in numerous clusters in the grapevine and poplar. These results suggested that recent tandem duplication played a major role in NBS-encoding gene expansion in perennial species. These duplication events, together with a higher probability of recombination revealed in this study, could compensate for the longer generation time in woody perennial species e.g. duplication and recombination could serve to generate novel resistance specificities. In addition, we observed extensive species-specific expansion in TIR-NBS-encoding genes. Non-TIR-NBS-encoding genes were poly- or paraphyletic, i.e. genes from three or more plant species were nested in different clades, suggesting different evolutionary patterns between these two gene types.

1.Nanjing University State Key Laboratory of Pharmaceutical Biotechnology, Department of Biology 210093 Nanjing China

... 采用Yang等^[33]的方法, 判别同源基因, 即2个基因联配部分序列的长度要覆盖2个基因中较长序列的80%和联配部分序列的相似性大于70% ...

2013

5.242

0.0

... 取融合表达的重组质粒转化原生质体^[34] ...

2004

12.444

0.0

... 而线虫中包含326个F-box蛋白家族成员^[35] ...

2002

9.737

0.0

Gagne

J M

, Downes

B P

, Shiu

S H

, Durski

A M

, Vierstra

R D

. The F-box subunit of the SCF E3 complex is encoded by a diverse superfamily of genes in Arabidopsis. Proc Natl Acad Sci USA, 2002, 99: 11519-11524

* Cellular and Molecular Biology Program and the Department of Horticulture, 1575 Linden Drive, University of Wisconsin, Madison,WI 53706; and † MIPS/Institute for Bioinformatics, GSF-National Research Center for Environment and Health, 85764 Neuherberg, Germany

The covalent attachment of ubiquitin is an important determinant for selective protein degradation by the 26S proteasome inplants and animals. The specificity of ubiquitination is often controlled by ubiquitin-protein ligases (or E3s), which facilitatethe transfer of ubiquitin to appropriate targets. One ligase type, the SCF E3s are composed of four proteins, cullin1/Cdc53,Rbx1/Roc1/Hrt1, Skp1, and an F-box protein. The F-box protein, which identifies the targets, binds to the Skp1 component ofthe complex through a degenerate N-terminal ≈60-aa motif called the F-box. Using published F-boxes as queries, we have identified694 potential F-box genes in Arabidopsis , making this gene superfamily one of the largest currently known in plants. Most of the encoded proteins contain interactiondomains C-terminal to the F-box that presumably participate in substrate recognition. The F-box proteins can be classifiedvia a phylogenetic approach into five major families, which can be further organized into multiple subfamilies. Sequence diversitywithin the subfamilies suggests that many F-box proteins have distinct functions and/or substrates. Representatives of allof the major families interact in yeast two-hybrid experiments with members of the Arabidopsis Skp family supporting their classification as F-box proteins. For some, a limited preference for Skps was observed, suggestingthat a hierarchical organization of SCF complexes exists defined by distinct Skp/F-box protein pairs. Collectively, the datashows that Arabidopsis has exploited the SCF complex and the ubiquitin/26S proteasome pathway as a major route for cellular regulation and thata diverse array of SCF targets is likely present in plants.

... 植物中有大量的F-box蛋白存在, 拟南芥中大约有600~700个F-box基因^[36,37], 水稻中包含687个F-box基因^[38] ...

... 根据C端蛋白-蛋白相互作用结构域的不同, 可以把拟南芥F-box家族分为19个亚族^[36,37] ...

2002

4.134

0.0

... 植物中有大量的F-box蛋白存在, 拟南芥中大约有600~700个F-box基因^[36,37], 水稻中包含687个F-box基因^[38] ...

... 根据C端蛋白-蛋白相互作用结构域的不同, 可以把拟南芥F-box家族分为19个亚族^[36,37] ...

2007

6.555

0.0

Jain

, Nijhawan

, Arora

, Agarwal

, Ray

, Sharma

, Kapoor

, Tyagi

A K

, Khurana

J P

. F-box proteins in rice. Genomewide analysis, classification, temporal and spatial gene expression during panicle and seed development, and regulation by light and abiotic stress. Plant Physiol, 2007, 143: 1467-1483

F-box proteins constitute a large family in eukaryotes and are characterized by a conserved F-box motif (approximately 40 amino acids). As components of the Skp1p-cullin-F-box complex, F-box proteins are critical for the controlled degradation of cellular proteins. We have identified 687 potential F-box proteins in rice ( Oryza sativa ), the model monocotyledonous plant, by a reiterative database search. Computational analysis revealed the presence of several other functional domains, including leucine-rich repeats, kelch repeats, F-box associated domain, domain of unknown function, and tubby domain in F-box proteins. Based upon their domain composition, they have been classified into 10 subfamilies. Several putative novel conserved motifs have been identified in F-box proteins, which do not contain any other known functional domain. An analysis of a complete set of F-box proteins in rice is presented, including classification, chromosomal location, conserved motifs, and phylogenetic relationship. It appears that the expansion of F-box family in rice, in large part, might have occurred due to localized gene duplications. Furthermore, comprehensive digital expression analysis of F-box protein-encoding genes has been complemented with microarray analysis. The results reveal specific and/or overlapping expression of rice F-box protein-encoding genes during floral transition as well as panicle and seed development. At least 43 F-box protein-encoding genes have been found to be differentially expressed in rice seedlings subjected to different abiotic stress conditions. The expression of several F-box protein-encoding genes is also influenced by light. The structure and function of F-box proteins in plants is discussed in light of these results and the published information. These data will be useful for prioritization of F-box proteins for functional validation in rice.

... 植物中有大量的F-box蛋白存在, 拟南芥中大约有600~700个F-box基因^[36,37], 水稻中包含687个F-box基因^[38] ...

... 水稻F-box家族分为10个亚族^[38] ...

... 第III类中SiF-box5和SiF-box12都含有Kelch repeats, Kelch是植物F-box蛋白特有的结构域^[38] ...

2013

2.881

0.0

Jia

, Wu

, Li

, Huang

, Zheng

. Genome-wide identification and characterization of F-box family in maize. Mol Genet Genomics, 2013, 288: 559-577

Abstract F-box-containing proteins, as the key components of the protein degradation machinery, are widely distributed in higher plants and are considered as one of the largest known families of regulatory proteins. The F-box protein family plays a crucial role in plant growth and development and in response to biotic and abiotic stresses. However, systematic analysis of the F-box family in maize ( Zea mays ) has not been reported yet. In this paper, we identified and characterised the maize F-box genes in a genome-wide scale, including phylogenetic analysis, chromosome distribution, gene structure, promoter analysis and gene expression profiles. A total of 359 F-box genes were identified and divided into 15 subgroups by phylogenetic analysis. The F-box domain was relatively conserved, whereas additional motifs outside the F-box domain may indicate the functional diversification of maize F-box genes. These genes were unevenly distributed in ten maize chromosomes, suggesting that they expanded in the maize genome because of tandem and segmental duplication events. The expression profiles suggested that the maize F-box genes had temporal and spatial expression patterns. Putative cis -acting regulatory DNA elements involved in abiotic stresses were observed in maize F-box gene promoters. The gene expression profiles under abiotic stresses also suggested that some genes participated in stress responsive pathways. Furthermore, ten genes were chosen for quantitative real-time PCR analysis under drought stress and the results were consistent with the microarray data. This study has produced a comparative genomics analysis of the maize ZmFBX gene family that can be used in further studies to uncover their roles in maize growth and development.

1. State Key Laboratory of Crop Biology, College of Life Sciences, Shandong Agricultural University, Taian, Shandong, 271018, People’s Republic of China

... 最近研究表明, 玉米中有359个F-box蛋白成员^[39] ...

... 玉米中分为13个亚族^[39] ...

2001

3.361

0.0

Andrade

M A

, Perez-Iratxeta

, Ponting

C P

. Protein repeats: structures, functions, and evolution. J Struct Biol, 2001, 134: 117-131

... 第V类中, 4个成员都包含F-box、LRR和FBD结构域, LRRs是Leu重复单元, 含有LRRs结构的蛋白一般与信号传导, DNA修复、重组、转录以及抗病等功能相关^[40] ...

1999

13.076

0.0

Smith

T F

, Gaitatzes

, Saxena

, Neer

E J

. The WD repeat: a common architecture for diverse functions. Trends Biochem Sci, 1999, 24: 181-185

... 在谷子19个F-box家族中, 3个蛋白包含WD40结构, WD40结构存在于所有真核生物中, 在信号传导、细胞周期调控、细胞凋亡方面起着重要作用^[41] ...

1999

2.914

0.0

Craig

K L

, Tyers

. The F-box: a new motif for ubiquitin dependent proteolysis in cell cycle regulation and signal transduction. Prog Biophys Mol Biol, 1999, 72: 299-328

... 据报道, F-box基因不仅参与信号传导等生命活动^[42], 还介导干旱等非生物胁迫响应 ...

2011

5.242

0.0

Koops

, Pelser

, Ignatz

, Klose

, Marrocco-Selden

, Kretsch

. EDL3 is an F-box protein involved in the regulation of abscisic acid signalling in Arabidopsis thaliana. J Exp Bot, 2011, 62: 5547-5560

The EID1-like protein 3 (EDL3) shows high similarity to EID1 (Empfindlicher im dunkelroten Licht 1), an F-box protein that functions as a negative regulator in the signalling cascade downstream of the phytochrome A photoreceptor in Arabidopsis thaliana . Analyses revealed a strong and rapid induction of EDL3 gene expression under osmotic stress, high salinity, and upon abscisic acid (ABA) application. Therefore, it was speculated that EDL3 is involved in the regulation of responses controlled by this plant hormone, which not only regulates many aspects of plant development but also integrates responses towards temperature, drought, osmotic, and salt stresses. Physiological data obtained with over-expresser lines and a conditional knock-down mutant demonstrated that EDL3 functions as a positive regulator in ABA-dependent signalling cascades that control seed germination, root growth, greening of etiolated seedlings, and transition to flowering. Results further demonstrate that EDL3 regulates anthocyanin accumulation under drought stress. The observed effects on physiological responses fit to tissue-specific expression patterns obtained with EDL3 -promoter:GUS lines. Bimolecular Fluorescence Complementation assays and yeast two-hybrid analyses showed that EDL3 carries a functional F-box domain. Thus, the protein is presumed to act as a component of a ubiquitin ligase complex that specifically directs negatively acting factors in ABA signalling to degradation via the proteasome.

... 在渗透胁迫和高盐胁迫下, DEL3 (EID₁-like protein3)高效、快速表达^[43] ...

... 与对照相比, DEL3的表达量上升, edl3-1表达量下降^[43] ...