植物细胞次生壁形成的研究进展

综述了近年有关植物细胞次生壁形成的研究进展 ,包括目前次生壁研究采用的主要实验系统、研究技术与方法、次生壁分子结构、次生壁形成与细胞骨架变化、激素、钙离子和钙调素等在次生壁形成中的作用以及次生壁形成与细胞程序性死亡的关系等     

杜仲次生木质部分化过程中木质素与半纤维素组分在细胞壁中分布的动态变化

利用紫外光显微镜、透射电子显微镜结合免疫胶体金标记,研究了杜仲（Eucommia ulmoides Oliv.)次生木质部分化过程中木质素与半纤维素组分（木葡聚糖和木聚糖）在细胞壁分布的动态变化。在形成层及细胞伸展区域,细胞壁具有木葡聚糖的分布,而没有木聚糖和木质素沉积,随着次生壁S1层的形成,木质素出现在细胞角隅和胞间层,木聚糖开始出现在S1层中,此时木葡聚糖则分布在初生壁和胞间层;随着次生,壁S2层及S3层的形成和加厚,木质逐逐步由细胞角隅和胞间层扩展到S1、S2和S3层,其沉积呈现出不均匀的块状或片状沉积模式,在次生壁各层形成与其木质化的同时,木聚糖逐渐分布于整个次生壁中,而木糖聚糖仍局限分布于初生壁和胞间层。结果表明,随着细胞次生壁的形成与木质化,细胞壁结构发生较大变化。细胞壁的不同区域,如细胞角隅、胞间层、初生壁和次生壁各层,具有不同的半纤维素组成,其与木质等细胞壁组分结构构成不同的细胞壁分子结构。     

毛竹茎秆纤维细胞发育过程中的细胞程序性死亡

利用TUNEL检测、细胞学及细胞化学方法,对毛竹茎秆纤维细胞发育过程中的细胞程序性死亡进行了研究。在次生壁形成的早期,纤维细胞出现染色质凝聚、细胞器膨胀、液泡膜解体和细胞质泡状化等典型的细胞程序性死亡形态学特征;TUNEL检测反应呈阳性,显示此时的纤维细胞核DNA发生了片段化。此时,在纤维细胞裂解的液泡膜、降解的细胞质和凝聚的染色质上具有ATPase活性。纤维细胞质的Ca^2＋水平会随着次生壁的形成而逐渐升高,随后Ca^2＋聚集成块状。在初生壁形成后期,纤维细胞染色质上的酸性磷酸酶（APase）活性增强。随着纤维次生壁的持续增厚,ATPase、酸性磷酸酶和Ca^2＋将在裂解的细胞质和凝聚的染色质上持续存在多年。结果表明,毛竹茎秆纤维细胞的次生壁形成过程是一个主动自溶的细胞程序性死亡过程。初生壁形成后期染色质上酸性磷酸酶活性增强及次生壁形成期胞质Ca^2＋的聚集,与纤维细胞的程序性死亡密切相关。ATPase,Ca^2＋和APase参与了纤维细胞程序性死亡过程中原生质体的降解。     

一种特殊的长寿细胞:毛竹茎秆纤维细胞

利用显微和细胞化学方法,对毛竹(Phyllostachys edulis)茎秆纤维次生壁形成过程中超微结构变化以及ATP酶、Ca2 -ATPase和酸性磷酸酶的超微细胞化学定位进行了研究.研究发现,次生壁形成早期,细胞核具有双层核膜,染色质凝聚,可见大量的线粒体、粗面内质网和高尔基体等细胞器存在于纤维细胞中;随后,双层核膜消失,细胞器将逐渐解体,多泡体开始出现在纤维细胞的细胞质;随着年龄的增加,纤维细胞壁逐渐增厚,并出现多层结构现象,而运输小泡、细胞膜、胞间连丝和凝聚的染色质将持续存在.在次生壁形成的整个过程中,ATP酶、Ca2 -ATPase和酸性磷酸酶在运输小泡、细胞膜、质膜内陷、胞间连丝和凝聚的染色质中将持续存在.结果表明,毛竹茎秆纤维细胞是一种不同于木本双子叶植物的长寿细胞,纤维原生质体中ATP酶和酸性磷酸酶的持续存在与次生壁的持续增厚密切相关.     

毛竹茎秆纤维发育过程的超微结构观察

利用透射和扫描电镜观察了毛竹(Phyllostachys edulis(Carr.)H.De Lehaie)茎秆纤维发育过程中的超微结构变化.在纤维细胞初生壁形成期,细胞质中线粒体、内质网、高尔基体等细胞器数量有明显的增加,出现大量的由内质网与高尔基体分泌形成的运输小泡,周质微管平行分布于质膜内侧,出现环状片层结构,并在细胞壁与质膜之间出现壁旁体结构.随着次生壁的逐渐形成,细胞质中细胞器逐渐地解体并出现多泡小体;纤维细胞核出现染色质凝聚并边缘化,但在8年生的纤维中可以持续存在;在纤维次生壁形成的整个阶段都存在与周围细胞相联系的胞间连丝和运输小泡;次生壁在前4年加厚明显,以后加厚程度减缓,但可以持续很长一段时间,并随着加厚出现宽窄交替的多层结构.结果表明,线粒体、内质网、高尔基体和壁旁体等细胞器与周质微管一起参与了初生壁和次生壁早期的形成;纤维细胞次生壁的形成过程就是一个漫长的程序性细胞死亡(PCD),而PCD的产物与胞间连丝一起参与了次生壁的形成与加厚;染色质凝聚并边缘化的细胞核与胞间连丝的持续存在,证明毛竹茎秆纤维细胞是一种典型的长寿细胞.     

毛竹茎秆纤维发育过程的超微结构观察

利用透射和扫描电镜观察了毛竹（Phyllostachys edulis (Carr.) H. De Lehaie）茎秆纤维发育过程中的超微结构变化。在纤维细胞初生壁形成期,细胞质中线粒体、内质网、高尔基体等细胞器数量有明显的增加,出现大量的由内质网与高尔基体分泌形成的运输小泡,周质微管平行分布于质膜内侧,出现环状片层结构,并在细胞壁与质膜之间出现壁旁体结构。随着次生壁的逐渐形成,细胞质中细胞器逐渐地解体并出现多泡小体;纤维细胞核出现染色质凝聚并边缘化,但在8 年生的纤维中可以持续存在;在纤维次生壁形成的整个阶段都存在与周围细胞相联系的胞间连丝和运输小泡;次生壁 在前4 年加厚明显,以后加厚程度减缓,但可以持续很长一段时间,并随着加厚出现宽窄交替的多层结构。结果表明,线粒体、内质网、高尔基体和壁旁体等细胞器与周质微管一起参与了初生壁和次生壁早期的形成;纤维细胞次生壁的形成过程就是一个漫长的程序性细胞死亡（PCD）,而PCD 的产物与胞间连丝一起参与了次生壁的形成与加厚;染色质凝聚并边缘化的细胞核与胞间连丝的持续存在,证明毛竹茎秆纤维细胞是一种典型的长寿细胞。     

毛竹茎秆纤维细胞发育过程中的细胞程序性死亡(英文)

利用TUNEL检测、细胞学及细胞化学方法,对毛竹茎秆纤维细胞发育过程中的细胞程序性死亡进行了研究。在次生壁形成的早期,纤维细胞出现染色质凝聚、细胞器膨胀、液泡膜解体和细胞质泡状化等典型的细胞程序性死亡形态学特征;TUNEL检测反应呈阳性,显示此时的纤维细胞核DNA发生了片段化。此时,在纤维细胞裂解的液泡膜、降解的细胞质和凝聚的染色质上具有ATPase活性。纤维细胞质的Ca2+水平会随着次生壁的形成而逐渐升高,随后Ca2+聚集成块状。在初生壁形成后期,纤维细胞染色质上的酸性磷酸酶(APase)活性增强。随着纤维次生壁的持续增厚,ATPase、酸性磷酸酶和Ca2+将在裂解的细胞质和凝聚的染色质上持续存在多年。结果表明,毛竹茎秆纤维细胞的次生壁形成过程是一个主动自溶的细胞程序性死亡过程。初生壁形成后期染色质上酸性磷酸酶活性增强及次生壁形成期胞质Ca2+的聚集,与纤维细胞的程序性死亡密切相关。ATPase,Ca2+和APase参与了纤维细胞程序性死亡过程中原生质体的降解。     

木材形成过程中次生壁沉积和细胞程序性死亡的分子调控机制

木材的形成经历了维管形成层细胞的增殖,木质部细胞的分化和扩张,次生细胞壁的沉积和细胞程序性死亡(programmed cell death, PCD)的过程.近年来,基于遗传学和组学分析,人们已经从模式植物拟南芥和杨树中鉴定出许多调控次生壁沉积的的关键转录因子和转录抑制因子.这些转录因子调控层次分明,共同构成次生壁沉积的转录调控网络,不仅在次生壁组分木质素、纤维素、木聚糖等物质的生物合成过程中起重要作用,而且可激活下游调控细胞程序性死亡相关的水解酶,启动木质部细胞的程序化死亡过程.对这些基因的生物学功能和调控网络进行解析,为阐明木材形成的分子生物学机制奠定了理论基础.本文综述了木材形成过程中次生壁沉积的转录调控网络和细胞程序性死亡相关的酶学机制及其最新研究进展.     

一种特殊的长寿细胞: 毛竹茎秆纤维细胞

利用显微和细胞化学方法, 对毛竹( Phyllostachys edulis) 茎秆纤维次生壁形成过程中超微结构变化以及ATP 酶、Ca²⁺ -ATP_ase 和酸性磷酸酶的超微细胞化学定位进行了研究。研究发现, 次生壁形成早期,细胞核具有双层核膜, 染色质凝聚, 可见大量的线粒体、粗面内质网和高尔基体等细胞器存在于纤维细胞中; 随后, 双层核膜消失, 细胞器将逐渐解体, 多泡体开始出现在纤维细胞的细胞质; 随着年龄的增加,纤维细胞壁逐渐增厚, 并出现多层结构现象, 而运输小泡、细胞膜、胞间连丝和凝聚的染色质将持续存在。在次生壁形成的整个过程中, ATP 酶、Ca²⁺ -ATP_ase 和酸性磷酸酶在运输小泡、细胞膜、质膜内陷、胞间连丝和凝聚的染色质中将持续存在。结果表明, 毛竹茎秆纤维细胞是一种不同于木本双子叶植物的长寿细胞, 纤维原生质体中ATP 酶和酸性磷酸酶的持续存在与次生壁的持续增厚密切相关。     

初生纹孔场与纹孔不能混淆

细胞形成次生壁时，在一些位置上不沉积壁物质，因此形成一些间隙，这种在次生壁上的未增厚部分称为纹孔（ｐｉｔ）；细胞生长时，幼细胞的初生壁扩展，并增加表面积和厚度，但是，初生壁的某些区域较薄、明显凹陷，这个区域称为初生纹孔场（ｐｒｉｍａｒｙｐｉｔｓｆｉｅ...     

The gapped xylem mutant identifies a common regulatory step in secondary cell wall deposition

The phenotype of the novel gapped xylem (gpx) mutant is described. gpx plants exhibit gaps in the xylem in positions where xylem elements would normally be located. These gaps are not part of the transpiration stream and result in gpx plants having fewer functional xylem elements. The gaps are due to the absence of a secondary cell wall in developing xylem elements, resulting in complete degradation of these elements during cell death, and illustrate the importance of the secondary cell wall in retaining a functional xylem element following programmed cell death. Consequently the gpx phenotype suggests that the processes of secondary cell wall formation and cell death are independently regulated in developing xylem. gpx plants also exhibit a highly irregular pattern of secondary cell wall thickening in interfascicular cells, with some cells apparently undergoing little or no secondary cell wall deposition. Secondary cell wall deposition in plants involves the co-ordinate regulation of several complex metabolic pathways. The gpx mutant identifies a key step involved in regulating the deposition of secondary cell wall material in both xylem and interfascicular cells, and suggests that a common regulatory step controls secondary cell wall formation in these diverse cell types. The gpx mutant offers a unique opportunity to elucidate the mechanism by which the complex processes involved in secondary cell wall formation are co-ordinately regulated.     

Identification of novel genes in Arabidopsis involved in secondary cell wall formation using expression profiling and reverse genetics

Forward genetic screens have led to the isolation of several genes involved in secondary cell wall formation. A variety of evidence, however, suggests that the list of genes identified is not exhaustive. To address this problem, microarray data have been generated from tissue undergoing secondary cell wall formation and used to identify genes that exhibit a similar expression pattern to the secondary cell wall-specific cellulose synthase genes IRREGULAR XYLEM1 (IRX1) and IRX3. Cross-referencing this analysis with publicly available microarray data resulted in the selection of 16 genes for reverse genetic analysis. Lines containing an insertion in seven of these genes exhibited a clear irx phenotype characteristic of a secondary cell wall defect. Only one line, containing an insertion in a member of the COBRA gene family, exhibited a large decrease in cellulose content. Five of the genes identified as being essential for secondary cell wall biosynthesis have not been previously characterized. These genes are likely to define entirely novel processes in secondary cell wall formation and illustrate the success of combining expression data with reverse genetics to address gene function.     

Genome-wide Expression Profiling in Seedlings of the Arabidopsis Mutant uro that is Defective in the Secondary Cell Wall Formation

Plant secondary growth is of tremendous importance, not only for plant growth and development but also for economic usefulness. Secondary tissues such as xylem and phloem are the conducting tissues in plant vascular systems, essentially for water and nutrient transport, respectively. On the other hand, products of plant secondary growth are important raw materials and renewable sources of energy. Although advances have been recently made towards describing molecular mechanisms that regulate secondary growth, the genetic control for this process is not yet fully understood. Secondary cell wall formation in plants shares some common mechanisms with other plant secondary growth processes. Thus, studies on the secondary cell wall formation using Arabidopsis may help to understand the regulatory mechanisms for plant secondary growth. We previously reported phenotypic characterizations of an Arabidopsis semi-dominant mutant, upright rosette （uro）, which is defective in secondary cell wall growth and has an unusually soft stem. Here, we show that lignification in the secondary cell wall in uro is aberrant by analyzing hypocotyl and stem. We also show genome-wide expression profiles of uro seedlings, using the Affymetrix GeneChip that contains approximately 24 000 Arabidopsis genes. Genes identified with altered expression levels include those that function in plant hormone biosynthesis and signaling, cell division and plant secondary tissue growth. These results provide useful information for further characterizations of the regulatory network in plant secondary cell wall formation.     

Dynamic Changes in Distribution of Lignin and Hemicelluloses in Cell Walls During Differentiation of Secondary Xylem in Eucommia ulmoides (in English)

The dynamic changes in the distribution of lignin and hemicelluloses (xylans and xyloglucans) in cell walls during the differentiation of secondary xylem in Eucommia ulmoides Oliv. were studied by means of ultraviolet light microscopy and transmission electron microscopy combined with immunogold labelling. In the cambial zone and cell expansion zone, xyloglucans were localized both in the tangential and radial walls, but no xylans or lignin were found in these regions. With the formation of secondary wall S1 layer, lignin occurred in the cell corners and middle lamella, while xylans appeared in S1 layer, and xyloglucans were localized in the primary walls and middle lamella. In pace with the formation of secondary wall S2 and S3 layer, lignification extended to S1, S2 and S3 layer in sequence, showing a patchy style of lignin deposition. Concurrently, xylans distributed in the whole secondary walls and xyloglucans, on the other hand, still localized in the primary walls and middle lamella. The results indicated that along with the formation and lignification of the secondary wall, great changes had taken place in the cell walls. Different parts of cell walls, such as cell corners, middle lamella, primary walls and various layers of secondary walls, had different kinds of hemicelluloses, which formed various cell wall architecture combined with lignin and other cell wall components.     

Secondary cell wall patterning during xylem differentiation

Xylem cell differentiation involves temporal and spatial regulation of secondary cell wall deposition. The cortical microtubules are known to regulate the spatial pattern of the secondary cell wall by orientating cellulose deposition. However, it is largely unknown how the microtubule arrangement is regulated during secondary wall formation. Recent findings of novel plant microtubule-associated proteins in developing xylem vessels shed new light on the regulation mechanism of the microtubule arrangement leading to secondary wall patterning. In addition, in vitro culture systems allow the dynamics of microtubules and microtubule-associated proteins during secondary cell wall formation to be followed. Therefore, this review focuses on novel aspects of microtubule dynamics leading to secondary cell wall patterning with a focus on microtubule-associated proteins.