杨树——林木基因组学研究的模式物种

林木植物是大多数生态系统中的主要生命形式, 在所有陆地生态系统中具有最大的生物量和生物生产力, 因此它具有重要的经济及生态价值。林木植物特有的多年生习性使其生物学研究相对困难, 因此需要寻找一种适用于遗传学及分子生物学方法做精细分析的模式植物。杨属(Populus)植物作为林业研究的模式植物, 具有优良的实验特性: 容易进行种间杂交和无性繁殖; 生长迅速, 并已建立完善的遗传转化系统; 基因组相对较小, 约450～550 Mbp; 易于进行遗传研究; 适应性强, 生长速度快, 丰产性强。自2002年起, 美国能源部与多家研究机构正式启动了杨属植物基因组计划, 目前已接近完成。本文综述了模式植物杨树及其基因组学的研究进展。     

多年生植物模式物种基因组研究的历史及进展

木本植物有许多不同于一年生草本植物的生物学特性,生物学家提出将木本植物 作为研究多年生植物的模式体系。杨属Populus树种由于研究基础较好且基因组较小,目前已 被广泛地接受作为多年生植物基因组研究的模式物种。随着杨属树种全基因组序列的测定, 杨属树种在多年生植物的功能基因组研究及一些基础科学问题的研究中将发挥重要作用。本 文综述了杨属树种基因组研究的历史、进展及将来的研究热点,旨在为我国多年生植物基因 组研究提供参考和借鉴。本文主要论述了以下几个方面的内容:（1）对杨属树种开展的细胞 遗传学研究;（2）在分子水平上对杨属树种进行的基因组研究,内容包括遗传作图、基因组 测序、物理图谱构建、基因芯片及连锁不平衡分析;（3）杨属树种基因组信息在探讨一些基 础科学问题中的潜在应用。     

杨属遗传多样性研究进展

 生物多样性的研究和保护是目前全世界普遍关注的问题,其中研究较多的是遗传多样性、物种多样性及生态系统多样性,遗传多样性作为生物多样性的重要组成部分,是生态系统多样性和物种多样性的基础方面。杨树是世界各国普遍种植的木本植物,研究其遗传多样性具有重要的理论和实践意义,国内外都开展了广泛研究。本文综述了杨属(Populus)植物的派、种及无性系等在表型、染色体、蛋白质及DNA水平的遗传多样性的研究进展。其表型多样性主要体现在不同水平上的种群大小、形态特征、物候期、材性以及对生物或非生物逆境的抗性等方面;在杨属植物中已发现有天然的三倍体及三倍体/非整倍体的杂交种存在;蛋白质的多态性主要集中于同工酶的研究,用于不同杨属植物的遗传差异,无性系或品种的特征、分化和遗传研究;DNA多态性是研究最多的,RFLP、RAPD、AFLP和SSR等分子标记已广泛用于杨属的遗传多样性研究中,根据不同的研究目的所选择的DNA（nDNA, cpDNA或mtDNA）也不同。并根据国内的研究现状,提出了几点建议。     

刨花楠不同相对生长速率下林木叶片碳氮磷的适应特征

分别对9年生与13年生刨花楠林木叶片氮磷养分之间关系及林木生物量相对生长速率与叶片碳氮磷化学计量比关系进行分析,探讨不同相对生长速率下的林木叶片N、P养分适应特征,并检验相对生长速率假说理论对刨花楠树种的适应性。结果表明:两种年龄刨花楠林木生物量相对生长速率、叶片C、N、P含量及其计量比值均存在显著差异;同一年龄的林木叶片N、P之间存在显著相关性,二者具有协同相关性;9年生林木叶片P含量及C∶P、N∶P与生物量相对生长速率呈二次曲线相关,而13年生林木叶片N、P含量及C∶N、C∶P、N∶P则与生物量相对生长速率均呈线性相关。研究表明,在能满足植物生长所需养分供给的土壤环境中,叶片N、P含量与林木相对生长速率间呈线性正相关,但当土壤中养分供应满足不了植物高速生长时,植物则会对有限的养分资源进行适应性调整。     

模式植物二穗短柄草研究进展与展望

二穗短柄草(Brachypodium distachyon)是一种温带禾本科植物,其植株矮小,自花授粉,生活周期短,生长条件简单,基因组小,易于进行遗传转化,与小麦、柳枝稷同属禾本科早熟禾亚科,是研究小麦、大麦等经济作物以及柳枝稷等能源植物的比较适合的模式植物。最近,二穗短柄草基因组测序及注释正式完成,有必要对其研究进展进行全面的总结。综述了二穗短柄草的基因组特征、基因表达模式、遗传转化等方面的最新研究进展,并对今后的研究方向作了展望,以促进对禾谷类经济作物和能源植物的深入研究。     

杨树抗冻性的研究现状

杨树是世界上广泛栽培的重要造林树种之一,我国是世界上杨树资源丰富的国家。杨树具有速生丰产、实用性强、无性繁殖能力强,且基因组较小等特点,现已成为研究林木生理和基因工程研究的模式植物。本文概述了国内外有关杨树抗冻性的生理生化性质及其指标、天然杂种的选择以及人工杂交选育种等方面研究现状,并展望了杨树基因工程在杨树抗冻性遗传改良中的应用前景。此外,对今后如何开展杨树抗冻改良育种的研究提出了一些设想。     

林木基因组学研究进展

林木基因组学研究进展迅速。结构基因组学方面,已构建了近40个主要造林树种的遗传连锁图谱,在不同树种中定位了30余个重要的数量性状位点,在部分树种中开展了基因组比较和综合图谱构建研究,杨树的全基因组测序已经完成,桉树的全基因组测序正在进行。功能基因组学方面,已分析了主要造林树种多种组织的转录组EST序列,对林木次生生长与木材形成、开花和抗寒性的形成等过程开展了功能基因组学研究。另外,探讨了林木基因组学研究的发展趋势,以期为我国林木基因组学研究提供有益的参考。     

杨树抗冻性的研究现状

杨树是世界上广泛栽培的重要造林树种之一,我国是世界上杨树资源丰富的国家,杨树具有速生丰产、实用性强、无性繁殖能力强,且基因组较小等特点,现已成为研究林木生理和基因工程研究的模式植物。本文概述了国内外有关杨树抗冻性的生理生化性质及其指标、天然杂种的选择以及人工杂交选育种等方面研究现状,并展望了杨树基因工程在杨树抗冻性遗传改良中的应用前景,此外,对信后如何开展杨树抗冻改良育种的研究提出了一些设想。     

豆科模式植物——蒺藜苜蓿

蒺藜苜蓿（Medicago truncatula）是国际上广泛用于研究仅属于豆科植物或者与之有关的某些生物过程的豆科模式植物,这些生物过程无法采用模式植物拟南芥进行研究。蒺藜苜蓿的基因组较小,二倍体,遗传学简单,遗传转化相对容易,再生时间较短。2003年,国际间合作的蒺藜苜蓿基因组测序计划已经启动。文章概述了蒺藜苜蓿基因组测序、生物信息数据库、遗传转化系统以及功能基因组研究工具的研究进展。     

单核苷酸多态性在林木中的研究进展

摘要: 单核苷酸多态性(Single nucleotide polymorphisms, SNPs)是许多生物体最丰富的遗传变异形式。林木是重要的植物类群和陆地植物生态系统的重要组成部分, SNP作为新的分子标记已应用于松、杨、黄杉、桉和云杉等属的多个树种的遗传育种学研究, 获得了包括核苷酸多样性、连锁不平衡及群体结构等相关的遗传信息, 这些研究主要建立在对候选基因序列进行测序分析的基础上。基于SNP的关联遗传学分析或连锁不平衡(Linkage disequilibrium, LD)作图, 已成为研究林木复杂数量性状的理想工具, 对桉树和火炬松的关联遗传学研究发现, 多个基因内的SNP位点与不同的木材性状相关联。利用SNP标记对林木遗传参数的估算从不同程度上揭示了林木群体进化规律及其生态学意义。SNP标记在林木中应用的不断深入, 必将极大地推动林木遗传育种学研究的发展。     

Out of the woods: forest biotechnology enters the genomic era

Trees represent a unique life form of upmost importance for mankind, as these organisms have developed a perennial lifestyle that produces the majority of terrestrial biomass. The difference between trees and annual plants is one of the main arguments behind the effort to sequence the entire genome of the poplar tree. This initiative is being backed up with a full-scale functional genomics effort on trees that will set a completely new agenda for forest research.     

Populus: arabidopsis for forestry. Do we need a model tree?

Trees are used to produce a variety of wood-based products including timber, pulp and paper. More recently, their use as a source of renewable energy has also been highlighted, as has their value for carbon mitigation within the Kyoto Protocol. Relative to food crops, the domestication of trees has only just begun; the long generation time and complex nature of juvenile and mature growth forms are contributory factors. To accelerate domestication, and to understand further some of the unique processes that occur in woody plants such as dormancy and secondary wood formation, a 'model' tree is needed. Here it is argued that Populus is rapidly becoming accepted as the 'model' woody plant and that such a 'model' tree is necessary to complement the genetic resource being developed in arabidopsis. The genus Populus (poplars, cottonwoods and aspens) contains approx. 30 species of woody plant, all found in the Northern hemisphere and exhibiting some of the fastest growth rates observed in temperate trees. Populus is fulfilling the 'model' role for a number of reasons. First, and most important, is the very recent commitment to sequence the Populus genome, a project initiated in February 2002. This will be the first woody plant to be sequenced. Other reasons include the relatively small genome size (450-550 Mbp) of Populus, the large number of molecular genetic maps and the ease of genetic transformation. Populus may also be propagated vegetatively, making mapping populations immortal and facilitating the production of large amounts of clonal material for experimentation. Hybridization occurs routinely and, in these respects, Populus has many similarities to arabidopsis. However, Populus also differs from arabidopsis in many respects, including being dioecious, which makes selfing and back-cross manipulations impossible. The long time-to-flower is also a limitation, whilst physiological and biochemical experiments are more readily conducted in Populus compared with the small-statured arabidopsis. Recent advances in the development of large expressed sequence tagged collections, microarray analysis and the free distribution of mapping pedigrees for quantitative trait loci analysis secure Populus as the ideal subject for further exploitation by a wide range of scientists including breeders, physiologists, biochemists and molecular biologists. In addition, and in contrast to other model plants, the genus Populus also has genuine commercial value as a tree for timber, plywood, pulp and paper.     

What genes make a tree a tree?

Woody growth is evolutionarily ancient, yet has been gained and lost multiple times in plant evolution and is readily enhanced or minimized in eudicot speciation. New molecular genetic and genomic studies in Populus and Arabidopsis that are defining the genes responsible for cambium function and woody growth suggest that the genes regulating woody growth are not unique to woody plants. Surprisingly, key genetic mechanisms originally characterized as regulating the meristematic cells of the shoot apical meristem are also expressed in the vascular cambium during woody growth. This has important implications for the development of Populus as a model species and illustrates why forest trees constitute a contrived group of plants that have more in common with herbaceous relatives than we foresters like to admit.     

林木基因克隆研究进展

林木种质资源丰富, 种质间遗传差异大, 控制林木重要性状的基因克隆及转化对培育优良林木新品种具有很强的实用价值, 但许多具有潜在应用价值的林木基因未得到充分发掘和有效分离。近年来, 随着各种不同林木cDNA文库的建立, 大规模随机EST测序技术的运用以及克隆技术的不断完善, 特别是毛果杨(Populus trichocarpa)基因组测序计划的完成, 大量与林木重要性状相关的基因被分离和鉴定。这些重要基因的获得为利用转基因技术培育高产、优质、抗逆、抗病虫害的林木新品种奠定了一定的基础。该文综述了20多年来国内外林木基因克隆的研究进展, 对基因克隆及其应用过程中亟待解决的问题进行了讨论, 并对其发展趋势进行展望。     

Insertional mutagenesis in <Emphasis Type="Italic">Populus</Emphasis>: relevance and feasibility

The recent sequencing of the first tree genome, that of the black cottonwood (Populus trichocarpa), opens a new chapter in tree functional genomics. While the completion of the genome is a milestone, mobilizing this significant resource for better understanding the growth and development of woody perennials will be an even greater undertaking in the years to come. In other model organisms, a critical tool for high-throughput analysis of gene function has been the generation of large mutagenized populations. Some mutagenesis technologies and approaches cannot be applied to trees because of their typically outcrossing breeding systems, high heterozygosity, large body size, and delayed flowering. In contrast, gene-tagging approaches that use insertional mutagenesis to create dominant phenotypes are ideally suited for trees and, especially, Populus. Both activation tagging and enhancer trap programs have been successful in identifying new genes important to tree development. The generation of genome-wide insertional mutant populations, which provide direct functional links between genes and phenotypes, should help to integrate in silico analyses of gene and protein expression, association studies of natural genetic polymorphism, and phenotypic analyses of adaptation and development.