类受体蛋白激酶与植物非生物胁迫应答

类受体蛋白激酶(receptor-like protein kinase,RLK)是植物信号转导网络中的重要成员,参与介导生长、发育以及逆境胁迫应答等多种细胞代谢过程．在植物细胞中已发现和克隆了富含亮氨酸重复区型(LRR)、凝集素型(lectin-like)和细胞壁相联型(WAK)等不同的RLK亚家族．这些RLK能够感受多种发育和外界环境胁迫信号, 并在植物对非生物胁迫的响应过程中发挥重要的调控作用．本文结合当今国内外研究进展,简述植物RLK的典型结构域特征,详细介绍多种RLK在植物逆境信号识别与转导中发挥的作用,同时对RLK在非生物胁迫应答中的具体作用机制进行了探讨．     

水稻NAC转录因子家族的研究进展

NAC转录因子家族是一类重要的转录调控因子,在植物中普遍存在。在水稻(Oryza sativa L.)生命历程中,NAC家族参与其细胞生长、组织发育、器官衰老等过程,且在应对外界环境刺激的响应过程中起重要作用。本研究介绍了水稻NAC转录因子家族的结构特点,并综述了水稻NAC转录因子家族参与调控植物生长发育的过程,以及在低温、高盐、病原菌等逆境胁迫中的作用与功能,并对水稻NAC家族今后的研究方向进行了展望。     

AP2功能基因在植物花发育中的重要作用

AP2基因作为调控植物花发育的功能基因,参与花分生特性建立、花器官的特性特化以及形成调控。所编码的AP2/EREBP转录因子的主要特征是都至少含有一个由60到70个左右的氨基酸组成高度保守的DNA结合区,称作AP2结合域。按其所含的AP2结构域的数目分为3个亚族,即AP2亚家族、EREBP亚家族和RAV亚家族,每个亚家族都有各自的作用。AP2基因不但自身调控着花、胚珠的发育,而且与其他因子相互协作,参与到复杂的花发育调控网络。将对AP2基因的特征和分类及其在花发育中的作用进行概述。     

谷子 MADS-box基因家族的鉴定和表达分析

MADS-box基因是真核生物中一类重要的转录因子,参与调控多项植物的生长发育过程。然而关于谷子穗发育的MADS-box基因研究比较少。本研究使用序列相似性检索,在Phytozome 13.0数据库中筛选并且鉴定出了68个谷子MADS家族成员,并对这些家族成员的物理化学性质、系统发育树、染色体定位、表达谱等进行了全面的分析。结果表明,谷子MADS家族成员在染色体上分布不均匀,可以分为5个亚族。通过组织特异性表达谱分析得到,多数MADS基因在穗中表达量要高于其他器官。此外利用转录组测序技术对发育初期的谷穗和成熟期的谷穗进行了转录组测序分析,筛选到数个与谷穗分生组织发育相关MADS-box基因。为进一步揭示MADS-box基因在谷子穗发育过程中的作用奠定了重要的基础。     

植物HD-Zip转录因子的生物学功能

HD-Zip转录因子属于Homeobox蛋白家族, 是植物特异转录因子, 由高度保守的HD(Homeodomain)结构域和Leu zipper(Zip)元件组成, 前者与DNA特异结合, 后者介导蛋白二聚体的形成。HD-Zip转录因子家族包括4个亚家族(HD-Zip Ⅰ-Ⅳ), 其成员通过与其他蛋白互作、参与激素介导的信号途径, 从而调控植物生长发育、光形态建成、花发育、果实发育和植物对逆境应答等生物学过程。文章对近几年关于植物HD-Zip转录因子参与上述生物学功能方面的研究进行了综述, 以期对新功能基因的挖掘和应用研究以及HD-Zip调控机制的阐明奠定基础。     

雷蒙德氏棉HSP70基因家族的进化分析及其同源基因在陆地棉中的表达分析

热激蛋白70家族(HSP70)是一类在植物中高度保守的分子伴侣蛋白,在细胞中协助蛋白质正确折叠。文章利用隐马可链夫模型(HMM)在雷蒙德氏棉(Gossypium raimondii L.)全基因组范围内进行HSP70基因家族成员进化分析,共得到30个HSP70家族成员。利用生物信息学对雷蒙德氏棉HSP70基因的结构、染色体分布、基因倍增模式以及系统进化进行分析,结果表明,HSP70基因家族根据亚细胞定位结果可分为不同的基因亚家族,各亚家族中HSP70基因具有相对保守的基因结构;染色体片段重复和串联重复是雷蒙德氏棉HSP70基因家族扩增的主要方式。通过对不同物种的HSP70基因家族进行系统进化分析可知,HSP70亚组的分化发生在单细胞植物形成前,且细胞质型HSP70成员大量扩增。比较陆地棉棉纤维发育不同时期的深度测序表达谱,发现HSP70基因可能参与棉纤维的生长发育。本研究结果有助于了解棉属植物HSP70基因家族的功能,以期为深入研究棉纤维发育过程中的分子调控机理提供基础。     

SERKs,拟南芥中一组参与多条细胞信号传导途径的共受体

植物体细胞胚胎发生类受体激酶(SERKs)属于富含亮氨酸重复序列类受体激酶(LRR-RLK)第二亚家族,在拟南芥(Arabidopsis thaliana)中共有5个成员,是一组国际植物学界的热点研究蛋白,广受关注.近年来的研究表明,SERKs作为共受体参与包括油菜素内酯信号传导、花药绒毡层细胞分化、气孔和维管束发育及花器官脱落在内的诸多生长发育过程;还参与细胞死亡调控、植物对病原菌的先天免疫反应等过程.本文综述了拟南芥中SERKs在这些过程中的功能     

转录调控因子Fox的功能及分子机制研究进展

Fox (Forkhead box)蛋白家族有19个亚族, 它们通过结合DNA, 激活或抑制目的基因的转录活性, 同时还能参与细胞信号转导、 细胞周期调控和新陈代谢的调节, 在生物体发育及其成熟的组织器官中均能发挥重要作用, 目前, 有关Fox蛋白家族的功能及分子机制已逐步成为免疫学、 遗传学、 医学以及肿瘤学领域的研究热点。本文综述了Fox家族成员的命名及分类、 蛋白结构及其DNA识别机制以及该家族成员如何参与Hh, TGF-β/SMAD, MAPK, Wnt/β-catenin和IGF信号通路的调控。Fox家族可调控线虫的咽、 果蝇的唾液腺以及哺乳动物的肝脏和眼睛等器官的发育, 能够影响细胞周期, 其家族成员FoxA可以和CREB、 GR结合调控新陈代谢。不同物种的Fox家族成员个数存在差异, 并且受到严格的进化选择。对其功能和分子进化机制进一步研究可为阐明生物的发育机理和人类疾病的防治提供新的思路。     

植物类受体激酶CrRLK1-L亚家族及其生物学功能

植物CrRLK1-L亚家族类受体激酶的胞外域具有新颖结构基序,但功能大都未知.该家族成员广泛存在于被子植物中,但在动物和微生物中不存在其同源物.CrRLK1-L家族成员相对较少,但组织表达非常广泛.它们定位于细胞质膜上,并且部分成员的定位还具有极性,这与其参与雌雄配子体的识别和受精作用密切相关.该家族成员普遍具有激酶活性,该活性对其功能的发挥至关重要.目前仅报道在拟南芥中参与助细胞与花粉的识别和调控营养组织的细胞伸长,但参与这些生物学过程的作用机制似乎独立于已知的信号通路之外,可能有自身独特的信号传导机制.所以对这一类具特有结构基序的类受体激酶基因的功能研究,将有助于解析植物特有生物学过程的分子作用机制,特别是在植物有性生殖过程中,合理利用这些分子开展育种实践对未来农业生产具有潜在的应用价值.     

梨Ⅲ型聚酮合成酶家族的比较基因组学研究及表达模式分析

植物Ⅲ型聚酮合成酶(PKS Ⅲ)在植物次生代谢产物生物合成中起着非常重要的作用。本研究从7种蔷薇科果树中共鉴定出57个PKS家族成员,随后进行种间比较基因组学分析。基因结构和保守基序分析表明,57个PKS多数由两个外显子和一个内含子组成,都具有PKS特有的保守结构域Chal-sti-synt-N和Chal-sti-synt-C。进化分析显示,57个PKS可明显分为4个亚家族,其中I亚家族成员数目最多。染色体定位及基因复制事件表明,PKS不均匀地分布在2~5条染色体上,且PKS家族进化过程主要受纯化选择作用。荧光定量分析发现,PbPKS4和PbPKS5表达模式与‘砀山酥梨’(Pyrus bretschneideri cv.‘Dangshan Su’)不同发育时期果实中木质素及石细胞含量变化趋势类似,因此推测这2个基因在梨果实木质素合成及石细胞发育过程中发挥一定作用。本研究为今后研究蔷薇科果树PKS家族建立了基础,也为从分子水平改善梨果实品质提供了理论依据。     

植物M/MLD类受体蛋白激酶及其生物学功能的研究进展北大核心CSCD

植物受体蛋白激酶通过与胞外信号结合感知和接收外部信号传递,在植物各个生理过程及生物代谢中发挥着重大的作用。其中M/MLD类受体蛋白激酶是一类植物特有的具有Malectin-like结构域的受体蛋白激酶。研究表明,M/MLD-RLKs亚家族参与植物发育过程及生物/非生物胁迫调控。该研究对近年来国内外有关植物M/MLD-RLKs的发现、结构特点以及生物学功能等方面的研究进展进行综述,并重点阐述其在调控植物根系、叶片、花发育及响应多种胁迫过程中的作用,为深入研究M/MLD-RLKs在植物生长发育过程中的生理功能提供参考。     

Challenges in understanding RLK function

Plants use receptor-like kinases (RLKs) to transduce extracellular signals into the cell. Recent advancements in RLK research include the cloning of the BRASSINOSTEROID INSENSITIVE 1 and CLAVATA 1 genes, revealing RLK roles in development. Our understanding of RLK function has also been broadened by transgenic approaches in the study of the RLKs pollen receptor kinase 1, and wall associated kinase 1. These results extend the observations that RLKs function in developmental processes and plant defense responses. Additionally, expression based studies suggest roles for other newly reported RLKs in development and light responses. Taken together, the studies confirm the importance of RLKs in diverse plant processes, yet major challenges remain. These include identifying ligands that activate RLKs and characterizing downstream pathways. These challenges can be conquered by coordinated efforts from investigators using molecular, genetic, and biochemical approaches.     

Multiple receptor-like kinase cDNAs from liverwort Marchantia polymorpha and two charophycean green algae, Closterium ehrenbergii and Nitella axillaris: Extensive gene duplications and gene shufflings in the early evolution of streptophytes

Plant receptor-like kinases (RLKs) comprise a large family with more than several hundred members in vascular plants. The RLK family is thought to have diverged specifically in the plant kingdom, and no family member has been identified in other lineages except for animals and Plasmodium, both of which have RLK related families of small size. To know the time of divergence of RLK family members by gene duplications and domain shufflings, comprehensive isolations of RLK cDNAs were performed from a nonvascular plant, liverwort Marchantia polymorpha and two charophycean green algae, Closterium ehrenbergii, and Nitella axillaris, thought to be the closest relatives to land plants. We obtained twenty-nine, fourteen, and thirteen RLK related cDNAs from M. polymorpha, C. ehrenbergii, and N. axillaris, respectively. The amino acid sequences of these RLKs were compared with those of vascular plants, and phylogenetic trees were inferred by GAMT, a genetic algorithm-based maximum likelihood (ML) method that outputs multiple trees, together with best one. The inferred ML trees revealed ancient gene duplications generating subfamilies with different domain organizations, which occurred extensively at least before the divergence of vascular and nonvascular plants. Rather it remains possible that the extensive gene duplications occurred during the early evolution of streptophytes. Multicellular-specific somatic embryogenesis receptor kinase (SERK) involved in somatic embryogenesis was found in a unicellular alga C. ehrenbergii, suggesting the evolution of SERK by gene recruitment of a unicellular gene.     

Flourishing in water: the early evolution and diversification of plant receptor-like kinases

Receptor-like kinases (RLKs) play significant roles in mediating innate immunity and development of plants. The evolution of plant RLKs has been characterized by extensive variation in copy numbers and domain configurations. However, much remains unknown about the origin, evolution, and early diversification of plant RLKs. Here, we perform phylogenomic analyses of RLKs across plants (Archaeplastida), including embryophytes, charophytes, chlorophytes, prasinodermophytes, glaucophytes, and rhodophytes. We identify the presence of RLKs in all the streptophytes (land plants and charophytes), nine out of 18 chlorophytes, one prasinodermophyte, and one glaucophyte, but not in rhodophytes. Interestingly, the copy number of RLKs increased drastically in streptophytes after the split of the clade of Mesostigmatophyceae and Chlorokybophyceae and other streptophytes. Moreover, phylogenetic analyses suggest RLKs from charophytes form diverse distinct clusters, and are dispersed along the diversity of land plant RLKs, indicating that RLKs have extensively diversified in charophytes and charophyte RLKs seeded the major diversity of land plant RLKs. We identify at least 81 and 76 different kinase-associated domains for charophyte and land plant RLKs, 23 of which are shared, suggesting that RLKs might have evolved in a modular fashion through frequent domain gains or losses. We also detect signatures of positive selection for many charophyte RLK groups, indicating potential functions in host–microbe interaction. Taken together, our findings provide significant insights into the early evolution and diversification of plant RLKs and the ancient evolution of plant–microbe symbiosis.     

An autophosphorylation site database for leucine‐rich repeat receptor‐like kinases in Arabidopsis thaliana

Leucine‐rich repeat receptor‐like kinases (LRR RLKs) form a large family of plant signaling proteins consisting of an extracellular domain connected by a single‐pass transmembrane sequence to a cytoplasmic kinase domain. Autophosphorylation on specific Ser and/or Thr residues in the cytoplasmic domain is often critical for the activation of several LRR RLK family members with proven functional roles in plant growth regulation, morphogenesis, disease resistance, and stress responses. While identification and functional characterization of in vivo phosphorylation sites is ultimately required for a full understanding of LRR RLK biology and function, bacterial expression of recombinant LRR RLK cytoplasmic catalytic domains for identification of in vitro autophosphorylation sites provides a useful resource for further targeted identification and functional analysis of in vivo sites. In this study we employed high‐throughput cloning and a variety of mass spectrometry approaches to generate an autophosphorylation site database representative of more than 30% of the approximately 223 LRR RLKs in Arabidopsis thaliana. We used His‐tagged constructs of complete cytoplasmic domains to identify a total of 592 phosphorylation events across 73 LRR RLKs, with 497 sites uniquely assigned to specific Ser (268 sites) or Thr (229 sites) residues in 68 LRR RLKs. Multiple autophosphorylation sites per LRR RLK were the norm, with an average of seven sites per cytoplasmic domain, while some proteins showed more than 20 unique autophosphorylation sites. The database was used to analyze trends in the localization of phosphorylation sites across cytoplasmic kinase subdomains and to derive a statistically significant sequence motif for phospho‐Ser autophosphorylation.     

Genome scanning for identification and mapping of receptor-like kinase (RLK) gene superfamily in <Emphasis Type="Italic">Solanum tuberosum</Emphasis>

Receptor-like kinases (RLKs) are a key class of genes that contribute to diverse phenomena from plant development to defense responses. The availability of completed potato genome sequences provide an excellent opportunity to identify and characterize RLK gene superfamily in this lineage. We identified 747 non-redundant RLK genes in the potato genome that were classified into 52 subfamilies, of which 58% members organized into tandem repeats. Nine of potato RLK subfamilies organized into tandem repeats. Also, six subfamilies exhibited lineage-specific expansion compared to Arabidopsis. The majority of RLK genes were physically organized within heterogeneous and homogeneous clusters on chromosomes and were unevenly distributed on the genome. Chromosome 2, 3 and 7 contained the highest number of RLK genes and the most underrepresented chromosomes were chromosome 8, 10 and 11. Taken together, our results provide a framework for future efforts on comparative, evolutionary and functional studies of the members of RLK superfamily.     

Soybean receptor-like protein kinase genes: paralogous divergence of a gene family

Receptor-like protein kinases (RLKs) in plants play major roles in cellular processes and stress responses. Three soybean (Glycine max) orthologs of Arabidopsis thaliana RLK were isolated and designated GmRLK1, GmRLK2, and GmRLK3. GmRLK1, GmRLK2, and GmRLK3 are similar in sequence, with GmRLK2 and GmRLK3 being nearly identical. The deduced amino acid sequences of GmRLK1, GmRLK2, and GmRLK3 possess characteristics of a transmembrane leucine-rich repeat RLK, AtCLV1. DNA fingerprinting and PCR analyses of a bacterial artificial chromosome library identified five GmRLK contigs (I-V): three for GmRLK1 (I, II, and V), one for GmRLK2 (III), and one for both GmRLK2 and GmRLK3 (IV). Phylogenetic analysis of the soybean RLKs together with other plant RLKs indicates that soybean and A. thaliana CLV1s generate a CLV1 branch, while soybean, A. thaliana, and rice RLKs generate an RLK branch. Thus, the AtCLV1 orthologs may have evolved later than the other pathogen-, environmental stress-, plant hormone-, and development-associated RLKs. A common ancestral GmRLK gene may have duplicated to give rise to GmRLK1, GmRLK2, and GmRLK3, or GmRLK2 and GmRLK3 may have resulted from a recent duplication event(s). Several amino acid replacements in the kinase domain of GmRLK1 compared with those of GmRLK2 and GmRLK3 may reflect evolutionary divergence of individual family members.     

RLKs orchestrate the signaling in plant male-female interaction

Different from animals, sessile plants are equipped with a large receptor-like kinase (RLK) superfamily. RLKs are a family of single trans-membrane proteins with divergent N-terminal extracellular domains capped by a signal peptide and C-terminal intracellular kinase. Researches in the last two decades have uncovered an increasing number of RLKs that regulate plant development, stress response and sexual reproduction, highlighting a dominant role of RLK signaling in cell-to-cell communications. Sexual reproduction in flowering plants is featured by interactions between the male gametophyte and the female tissues to facilitate sperm delivery and fertilization. Emerging evidences suggest that RLKs regulate almost every aspect of plant reproductive process, especially during pollination. Therefore, in this review we will focus mainly on the function and signaling of RLKs in plant male-female interaction and discuss the future prospects on these topics.     

Expansion of the receptor-like kinase/Pelle gene family and receptor-like proteins in Arabidopsis

Receptor-like kinases (RLKs) are a family of transmembrane proteins with versatile N-terminal extracellular domains and C-terminal intracellular kinases. They control a wide range of physiological responses in plants and belong to one of the largest gene families in the Arabidopsis genome with more than 600 members. Interestingly, this gene family constitutes 60% of all kinases in Arabidopsis and accounts for nearly all transmembrane kinases in Arabidopsis. Analysis of four fungal, six metazoan, and two Plasmodium sp. genomes indicates that the family was represented in all but fungal genomes, indicating an ancient origin for the family with a more recent expansion only in the plant lineages. The RLK/Pelle family can be divided into several subfamilies based on three independent criteria: the phylogeny based on kinase domain sequences, the extracellular domain identities, and intron locations and phases. A large number of receptor-like proteins (RLPs) resembling the extracellular domains of RLKs are also found in the Arabidopsis genome. However, not all RLK subfamilies have corresponding RLPs. Several RLK/Pelle subfamilies have undergone differential expansions. More than 33% of the RLK/Pelle members are found in tandem clusters, substantially higher than the genome average. In addition, 470 of the RLK/Pelle family members are located within the segmentally duplicated regions in the Arabidopsis genome and 268 of them have a close relative in the corresponding regions. Therefore, tandem duplications and segmental/whole-genome duplications represent two of the major mechanisms for the expansion of the RLK/Pelle family in Arabidopsis.