日本落叶松木质部发育相关基因筛选及其共表达网络构建

【目的】鉴定日本落叶松木质部发育相关基因,构建核心基因与木质部发育相关基因的共表达网络,为后期开展日本落叶松木材形成相关研究提供参考。【方法】对日本落叶松木质部、韧皮部和针叶3个组织进行二代和三代转录组测序,利用R软件的DEseq2包筛选木质部相对韧皮部和木质部相对针叶的差异表达基因,通过整合2组差异基因获得木质部特异表达基因,借助GO、KEGG及BLASTN等生物信息学分析手段探索基因功能,利用WGCNA分析构建木质部特异表达基因共表达网络。【结果】共获得2 596个木质部特异的高表达和低表达基因;GO分析结果显示这些基因在代谢过程、细胞过程、定位膜、细胞、细胞组件、催化活性、位点结合和转运活性等分类中显著富集; KEGG分析结果显示这些基因在淀粉和蔗糖代谢、类黄酮生物合成和代谢途径通路中显著富集,在苯丙烷代谢途径及淀粉和蔗糖代谢途径中分别富集到38个和196个基因;鉴定出木材形成相关基因,包括木质素合成相关基因PAL4、CCR1、C4H、HCT、COMT1、PER12、PER52、CYP98A3、LAC12和LAC17等,纤维素和半纤维素合成相关基因DEC、CEL1、Csl、CTL2和SPS3等; 2 596个木质部特异的高表达和低表达基因经WGCNA分析后筛选出与木质部发育相关基因关联度较高的17个核心基因。【结论】筛选的日本落叶松木质部发育相关基因参与半乳甘露聚糖合成、木葡聚糖合成、纤维素微纤丝形成、细胞壁纤维素合成、次生细胞壁形成过程、纤维伸长过程、调控合成木质素的碳代谢流、木质素生物合成及降解、木质素单体聚合、木质素单体甲基化和细胞程序化死亡等木材形成相关生物学过程;在共表达网络中筛选出的17个核心基因可作为今后研究的重点来探索其在木材形成过程中的具体功能。     

白皮松木质部分化中轴向管胞和射线细胞的微观构造与木质化研究

以白皮松（Pinus bungeana）为研究对象,采用光学显微镜、偏光显微镜和扫描电子显微镜结合细胞壁组织化学染色等木材解剖学方法,探究木质部分化中轴向管胞与射线细胞的微观构造特征变化及其木质化进程。组织离析与形态学分析结果表明：木质部分化中轴向管胞在细胞扩大后其细胞长度基本稳定,随着木质部的分化,射线管胞的细胞壁厚度不断增加,而在当年生木质部中未发现射线薄壁细胞的细胞壁加厚。木质素的自发荧光及其特异性染色分析结果表明：轴向管胞的木质化进程开始于次生壁S1层加厚阶段的胞间层与细胞角隅,射线管胞的木质化进程与轴向管胞类似,在当年生木质部内射线薄壁细胞未发生木质化。木质部分化过程中管胞（轴向、射线）与射线薄壁细胞的细胞壁加厚及木质化进程具有明显差异。     

植物激素与木材形成

综述了植物激素在树木维管组织分化中的作用．生长素是调节木质部细胞分化的主要因子；细胞分裂素在诱导分生组织发端和分生细胞分化上起重要作用；赤霉素促进纤维的形成，赤霉素和生长素结合使用，可有效地促进次生木质部中纤维分化；乙烯能促进树木韧皮部和木质部分化。还讨沦了木材形成过程中的三个主要问题：生长素含量的多少直接控制了沿树轴木质部导管、管胞的大小和密度；外界环境因子影响木质部细胞分化；温带落叶阔叶树环孔材和散孔材形成的机理，生长素控制环孔材树种早材宽导管的形成以及晚材窄导管和纤维形成的假说。对今后改善树木木材的质和量上有一定参考价值。     

活立木生理干燥过程中水分传输和散失机制探讨

木材中的水分会严重影响木材加工和使用性能,必须通过干燥使木材含水率控制在适宜范围内。常规蒸汽干燥耗能大、干燥缺陷多,热泵除湿干燥、太阳能干燥等新型节能干燥技术工业化应用尚不理想,因此,本文从木材水分来源和树木水分生理特性出发,探讨基于蒸腾作用降低木材水分的活立木生理干燥理论和技术,并从水分与植物生理的角度阐述活立木生理干燥的理论基础。通过分析树叶水分蒸发研究进展,总结叶内水分可能的3种蒸发位点,即暴露在内部气体空间的所有叶肉细胞和表皮细胞、气孔下腔室周围大部分区域的叶肉细胞和表皮细胞以及气孔下腔室周围其他区域的叶肉细胞和表皮细胞。通过分析植物体内水分传输机制研究进展和现状,总结植物叶内水分传输的3种可能途径,即通过胞间连丝的共质体传输途径、通过水孔蛋白的跨细胞传输途径以及通过未栓化细胞壁的质外体传输途径。阐明被广泛用于解释木质部水分长距离运输的内聚力-张力学说,分析其目前存在的争论及一些新提出的学说,如补偿压学说、多驱动力学说或水门学说,并分析木质部水分运输过程中时常发生的空穴和栓塞现象及其可能的恢复机制。在此基础上,提出今后研究活立木生理干燥过程中水分传输和散失机制的几个重点和方向:一是探讨生理干燥过程中处于水分胁迫状态下树叶叶孔蒸腾和角质层蒸腾之间的关系;二是探讨生理干燥过程中处于严重水分胁迫状态下树木叶内水分传输途径和蒸发位点;三是探讨纹孔等微观构造在木质部水分长距离传输中的作用以及在空穴和栓塞产生和恢复过程中的作用;四是探讨生理干燥过程中木质部内空穴和栓塞的产生和恢复机制及其对水分长距离传输的作用和影响。     

气候变化对树木木质部生长影响的研究进展

【目的】树木生长受多种气候因子的影响,与年轮宽度相比,树木木质部细胞的生长变化记录了更为详尽的气候信息,可以丰富当前从传统树轮气候学或树轮生态学研究中所得到的认识。本研究以期为利用树轮木质部细胞特征研究全球变化提供参考。【方法】综述了树木木质部形成过程、研究木质部细胞生长变化对气候因子响应的常用方法以及气候变化(温度、降水、光照强度、光周期和CO2浓度等)对形成层活动及木质部细胞生长变化的影响。【结果】目前,在细胞水平上研究树木木质部生长变化对气候因子响应过程的主要方法为：钉针法和微芯取样法,钉针法主要适用于研究形成层对损伤的响应或是记录热带树木木质部生长的增量,而微芯取样法可以清楚地观察到生长季各阶段形成层细胞数量的变化及木质部形成过程;树木木质部的生长过程一般从形成层恢复活动开始,经过细胞的伸展、次生细胞壁的加厚和木质化后,形成成熟的次生木质部细胞,其中的每一个阶段都会受到气候因子变化的影响;树木木质部细胞生长的变化与温度、降水、光照强度、光周期和 CO2浓度等气候因子有着复杂的相关关系;温度主要影响树木生长季前期形成层的活动,改变形成层恢复活动的时间、细胞的分裂速率及活动周期,但并不会改变木质部细胞分化的序列和季节生长模式;降水主要影响形成层细胞分裂的速率、在生长季的持续时间以及管胞的直径等,降水不足会降低树木细胞分裂的速率,缩短细胞分化的时间,导致生长季的持续时间减少,同时抑制细胞伸展,管腔直径较小;光照强度主要影响树木的光合速率来间接影响形成层细胞的活动;光周期主要控制木质部细胞的最大生长速率和生长结束期;有关 CO2浓度升高对树木木质部细胞的影响,研究结果并不一致,一些研究认为,CO2浓度升高影响细胞的分裂速率和细胞的扩张而不是次生细胞壁的加厚,从而造成早材生长轮宽度明显增加,但也有研究表明,CO2浓度升高并没有引起早材细胞结构较大的变化,相反晚材生长环的宽度却明显增加。【结论】随着研究理论和技术手段的不断成熟,极端灾害下树木木质部细胞的生长特征、不同树种木质部细胞的生长对气候变化的响应以及多种气候因子对树木木质部生长的综合影响将成为今后研究的主要方向。     

木麻黄树干细胞组织结构与对星天牛抗虫性的关系

本文报道了４个品系木麻黄木材组织显微解剖结果，分析了木材导管、导管间区木化细胞和射线等物理特征，探讨了木麻黄抗虫机制。高抗品系Ｃ４４木材导管数量少，单位面积（１ｍｍ２）仅有２０．２５个导管，且占木材面积少，仅为９．７２％；导管间区木化细胞较明显，壁厚与腔径比值最大，为５６．５９；单位面积射线的束数、长度、宽度和射线面积／总面积等指标均为最低。说明该品系木麻黄木质部水分含量低、材质坚硬致密、木质部提供养分少，因而不利于星天牛幼虫的生存发育。     

在成熟的次生木质部细胞中发现细胞核

细胞质及细胞核是生活的植物细胞的重要组成部分是无可非议的。然而,在成熟的木材细胞中,即在次生壁完全增厚并木质化的次生木质部内尚存在完整的细胞核及细胞质则实为罕见。除作者近年的报道外,国内外从未见报道。一般认为,立木木质部的大部分(首先是心材部分),以及树木伐倒后去皮的木质部,都是由原生质已经死亡的死细胞组成,这种细胞只包括细胞壁和细     

答读者问

问:树木为什么会出现连理现象? 答:树木枝干发生连结现象,称为连理枝。要了解这种现象的形成,首先需要知道树木茎的构造和生长。树木的茎干是由树皮、韧皮部、形成层、木质部和髓心构成的。茎在横断面上的生长加粗,表现为年轮的形式。形成层每年向内产生木质部细胞,向外产生韧皮部细胞。木质部的增加,直接构成木材的本身;韧皮部的增加,继续产生新的树皮。年复一年,树木就逐渐加     

木材干燥过程中声发射信号分析北大核心

对木材在干燥过程中产生的声发射信号进行采集和特征分析,结果表明:木材干燥过程中主要采集到两种不同类型的声发射信号:一种信号来自木材内部自由水的蒸发,与木质部导管内水分空穴化过程的声发射信号一致;另一种信号与木材干燥过快时自身形变开裂相关。作为一种木材干燥质量的控制方法,测量和分析声发射信号,可为木材干燥温度和湿度的控制提供依据,实现防止木材干裂、提高木材干燥质量的目的。     

木材干燥过程中声发射信号分析

对木材在干燥过程中产生的声发射信号进行采集和特征分析,结果表明:木材干燥过程中主要采集到两种不同类型的声发射信号:一种信号来自木材内部自由水的蒸发,与木质部导管内水分空穴化过程的声发射信号一致;另一种信号与木材干燥过快时自身形变开裂相关.作为一种木材干燥质量的控制方法,测量和分析声发射信号,可为木材干燥温度和湿度的控制提供依据,实现防止木材干裂、提高木材干燥质量的目的.     

木质部细胞分化的研究简述

本文主要从木质部细胞分化常用的实验系统、木质部细胞分化的诱导、木质部细胞的编程性死亡以及次生壁的构建4个方面阐述了木质部细胞分化的研究。     

核桃芽接愈合的组织学机制

【目的】通过对核桃芽接愈合过程的组织学观察,研究不同嫁接时期砧穗在愈伤组织形成、增殖和输导组织连接过程中的组织学变化机制,旨在为核桃和其他木本植物嫁接提供理论指导。【方法】以12年生核桃品种‘香玲’半木质化新梢中部芽片为接穗,2年生本地核桃实生苗为砧木,采用方块芽接法分别于砧木萌芽后40~47天和100~107天2个时间进行芽接试验,采用本课题组建立的非均质化材料切片方法对芽接愈合过程进行组织学观察。【结果】核桃萌芽后40~47天的砧穗愈合方式与前人报道一致,即形成层细胞在嫁接体愈合过程中起主导作用,愈合过程中主要组织学变化包括:隔离层的产生(3天)→砧穗形成层细胞分化产生愈伤组织(4天)→愈伤组织连接(6天)→砧穗原有形成层的修复与连接(12天),完成愈合需要12~15天,成活率达95.6%。而在萌芽后100~107天的砧穗愈合过程中,发现了与上述常规愈合不同的嫁接愈合新方式。此过程中木射线细胞起主导作用,愈合过程依次历经隔离层的产生(3天)→木射线细胞分化产生愈伤组织(4天)→砧穗愈伤组织连接(9天)→导管分化形成(15天)→新的形成层产生以及木射线的连接(25天),需要25~30天才能完成愈合,成活率为61.7%;在木射线细胞主导的砧穗愈合过程中砧木一侧的隔离层要先于接穗发生溶解,在砧穗愈伤组织完成连接时消失,而接穗表面隔离层在愈伤组织分化时依然存在,直至新的形成层产生后才完全溶解消失。【结论】核桃芽接过程中,除了常见的由形成层细胞主导的砧穗愈合方式外,还存在由木射线细胞主导的砧穗木质部的愈合新方式,木射线细胞脱分化产生愈伤组织,并进一步分化出导管、形成层和木射线等组织完成最终的砧穗连接;在此过程中愈伤组织主要来自于砧木木射线细胞且砧木隔离层要先于接穗发生溶解。     

Lignin deposition during earlywood and latewood formation in Scots pine stems

Lignin deposition at consecutive secondary wall thickening stages of early and late xylem cells during annual ring wood formation in Scots pine (Pinus sylvestris L.) stems was studied. Lignin patterns, isolated by thioglycolic acid method, consisted of alcohol-soluble (LTGA-I) and alkali-soluble (LTGA-II) fractions. The sum of two fractions, being the total lignin content, gradually increased in the course of lignification. However, the increments of lignin amount at each development stage of early and late tracheids were different. The intensity of lignin deposition increased in the course of earlywood tracheid maturation and decreased toward the end of latewood cell differentiation. The deposition of two lignin fractions in each layer of forming wood also occurred oppositely. The increment of LTGA-I descended, whereas that of LTGA-II increased from the beginning to the end of early xylem lignification. In contrast, LTGA-I increment dropped, whereas LTGA-II rose during late xylem lignification. Gel permeation chromatography showed that the lignins, formed at the beginning of lignification, were more homogeneous and had higher molecular weight compared with the lignins at the end of cell differentiation. Besides, the content of cellulose, estimated as the residue after lignin isolation, and of cell wall substances, presented as cell wall cross-section areas, at consecutive maturation stages of early and late xylem cells have been found to be different. The data show that lignin deposition occurred in different conditions and with opposite dynamics during early and late xylem formation.     

Effects of applied gibberellins and uniconazole-P on gravitropism and xylem formation in horizontally positionedFraxinus mandshurica seedlings

The present study deals with the effects of gibberellins (GA₃ GA₄) and uniconazole-P, an inhibitor of gibberellin biosynthesis, on negative gravitropism and xylem formation in the stems of horizontally positioned, 2-year-oldFraxinus mandshurica Rupr. var.japonica Maxim. seedlings. Each growth regulator (100 g) dissolved in 5 l acetone (50%) was applied to the basal node of the current shoot on May 24, 1995. The same treatment was repeated five times weekly until June 28. Five seedlings were used for each treatment. The seedlings were positioned horizontally 24h after the first application on May 25. Within 5 weeks the horizontal stem of control and GA-treated seedlings exhibited negative gravitropism. In contrast, the application of uniconazole-P inhibited negative gravitropic stem bending. The application of GAs increased the number of gelatinous fibers having thickened cell walls on the upper side of stems. The uniconazole-P application decreased xylem cell formation but did not inhibit the formation of gelatinous fibers. These results indicate that not only the differentiation of gelatinous fibers but also xylem increment is important in the negative gravitropism of horizontally positionedF. mandshurica seedlings. These results also suggest that GAs may be involved in xylem cell :formation rather than the differentiation of gelatinous fibers in this species.Part of this research was reported at the 23rd annual meeting of the Plant Growth Regulation Society of America, Calgary, Canada, July 1996     

Tracheary elements that resemble secondary xylem in calli derived from the conifers, Torreya nucifera and Cryptomeria japonica

Differentiated cells were recognized in calli derived from needles of Torreya nucifera and in calli derived from immature zygotic embryos of Cryptomeria japonica. Some differentiated cells resembled tracheary elements of primary xylem with spiral or reticulate thickening of cell walls. Other cells resembled tracheary elements with thick cell walls and bordered pits, which are features of secondary xylem. These tracheary elements were formed in cell clusters. Tracheary elements in calli of T. nucifera formed more highly developed structures, such as bordered pits and spiral thickening, than those of C. japonica. Cultured cells derived from conifers might provide a good model for studies of the differentiation of secondary xylem in vitro.     

树木木质部细胞凋亡及心材形成机理研究进展

概述有关木纤维、管状分子与薄壁细胞等木质部细胞凋亡和心材形成方面的研究成果,探讨木质部细胞凋亡与心材形成之间的信号通路及调控机理,并基于当前研究现状,提出几点建议。     

毛白杨形成层的活动周期及其POD同工酶的变化

北京地区生长的毛白杨 (PopulustomentosaCarr .)在春季芽膨大前 ,形成层带已经进入了恢复活动期。在芽展开后一周形成层进入活动期 ,细胞开始分裂 ,细胞层数也有所增加 ,同时分化出未成熟的韧皮部细胞。未成熟韧皮部细胞的数量在形成层活动初期快速增长 ,随后是一个持续而缓慢的连续过程。木质部的产生比韧皮部要晚 3周 ,未成熟的木质部细胞层数在开始变化不大 ,5月底～ 6月底 ,增长速度明显高于初期 ,随后便急剧下降。成熟木质部细胞的出现比成熟的韧皮部细胞要晚约一个月。与其他研究结果不同 ,毛白杨在4月底到 6月底这段时间 ,形成层细胞层数迅速下降 ,未成熟韧皮部细胞也略有减少 ,而未成熟和成熟的木质部细胞则大量增加。木质部在 9月底停止分化 ,较韧皮部要晚约 2 0d。 9月底形成层进入休眠期后 ,形成层细胞层数基本保持不变 ,直到翌年春天恢复活动。毛白杨形成层区域的过氧化物酶 (POD)同工酶在形成层活动期其活性低、酶带少 ,而恢复活动期和休眠期其活性高且酶带多 ,组织化学反应还表明 ,不同时期POD在不同的细胞和组织中的活性不同。本文对POD同工酶在形成层活动周期中的变化与其组织分化的关系进行了讨论。     

Ultrastructural analysis of the differentiation process of secondary xylem vessel element in Populus deltoides

Using electronic microscopy, ultrastructural changes were observed during differentiation in a secondary xylem vessel element (VE) in Populus deltoides. Results showed that morphological development of VE differentiation was successively divided into three stages. First was primary cell wall outspread (the initial stage), where the VE was highly vacuolated and the protoplasma distributed along the cell wall. Second was secondary cell wall construction (the pivotal stage), where substances accumulated before the tonoplast broke, and the VE organelle was distinct. Golgi bodies and vesicles, which were associated closely with synthesis and transportation of secondary cell wall substances, were also abundant. After the tonoplast broke, these substances accumulated faster. Simultaneously, the protoplasm was disaggregated and the agglomerated chromatin was distributed over the margin of the nucleus, showing the typical characteristics of programmed cell death (PCD). During secondary cell wall formation, no cell wall substances accumulated between the terminal cell walls of neighboring VEs. In addition, terminal cell wall substances were disaggregated in the post secondary cell wall formation. Later, when the remnant terminal cell wall was broken in the third stage, perforation occurred. Thus, for these successive stages of VE differentiation, the critical stage, when differentiation was not reversible, was at the start of secondary cell wall formation with succeeding VE differentiation similar to a typical PCD process.     

拟南芥MYB基因对次生维管系统发育的影响

根据拟南芥ATH1全基因芯片分析毛白杨维管再生过程中基因表达谱的结果,选取了在芯片中特异表达且表达量较高的MYB转录因子家族的7个基因,通过拟南芥突变体研究其对次生维管系统发育的影响及与木材形成相关的关系。通过"三引物法"的纯杂合鉴定和切片显微观察,发现其中1个突变系只有纯合突变体没有杂合体,并且茎基部维管束间纤维细胞壁和野生型相比明显增厚,细胞数量减少,细胞腔也随之减小,推测该突变体可能与次生壁的沉积有关系。     

Distribution of lignin interunit bonds in the differentiating xylem of compression and normal woods of Pinus thunbergii

The lignification process and lignin distribution at different stages of cell wall differentiation in the secondary xylem of compression and normal woods of Pinus thunbergii were investigated by thioacidolysis and subsequent desulfuration. We prepared 50-µm-thick, contiguous tangential sections of pine shoots, cut from the cambial zone through to mature xylem. In compression wood, uncondensed guaiacyl (G) and p-hydroxyphenyl (H) lignins were deposited simultaneously from early to late stages of lignification. The various types of G-G, G-H, and H-H dimers were detected in compression wood, and the ratio of G-H and H-H dimers to total dimers increased as lignification proceeded. In contrast, uncondensed and condensed H units were detected in trace amounts in normal wood. Significant differences in the relative distributions of lignin interunit linkages were not observed between compression and normal woods or between differentiating and mature xylems in either compression or normal woods.Part of this report was presented at the 10th International Symposium on Wood and Pulping Chemistry, Yokohama, June, 1999