植物光合结构与非光合结构的功能平衡:来自三种亚热带乔木树种的实验证据

来自三种亚热带乔木树种的实验证据 曾波 （1.西南师范大学生命科学学院,重庆400715;2.Department of Plant Ecology, Utrecht Unixersity, P.O.Box 80084,3508TB Utrecht,The Netherlands) 摘要： 植物的地上部分和地下部分存在功能性平衡现已十分清楚,但植物的地上部分是否在其光合结构（叶组织）和非光合结构（枝和茎）之间也存在功能性平衡尚不明晰。本文提出两个研究假设并检验之：1）植物地上部分在其光合与非光合结构之间存在功能性平衡;2）此功能性平衡的维持依赖于对光合和非光合结构生物量分配的调节。为验证此假设,采用枝叶修剪的方式（连续两年修剪,四个修剪强度：0,20%,50%,70%）对3种亚热带乔木树种榕(Ficus microcarpa）、黄桷树（Ficus uirens）和樟（Cinnamomum camphora）进行了研究。结果表明,修剪使所有树种地上部分的光合与非光合结构生物量比率（P/NP）立即下降,下降程度随修剪强度的升高而增大。但不论是首次修剪还是第二次修剪,修剪处理一年后,修剪株地上部分的光合与非光合结构生物量比率升高,且此生物量比率不低于非修剪株的光合与非光合结构生物量比率。此研究结果证实了植物地上部分光合与非光合结构间存在功能性平衡的假设。与假设一致,植株的生物量分配在修剪后发生了改变：修剪株加大了对光合结构（叶组织）的生物量分配（大量的新生产地上部分生物量被分配到光合结构）,同时却减少了对非光合结构（枝和茎）的分配。此分配格局的改变保证了光合与非光合结构功能性平衡的恢复与维持。可以认为,通过改变生物量分配格局以维持光合 与非光合结构功能平衡是植物抵御外来干扰和e 或损伤的一种有效策略。 关键词： 生物量分配;功能平衡;非光合结构;光合结构;乔木     

植物光合结构与非光合结构的功能平衡：来自三种亚热带乔木树种的实验证据

植物的地上部分和地下部分存在功能性平衡现已十分清楚,但植物的地上部分是否在其光合结构（叶组织）和非光合结构（枝和茎）之间也存在功能性平衡尚不明晰,本文提出两个研究假设并检验之：1）植物地上部分在其光合与非光合结构之间存在功能性平衡;2）此功能性平衡的维持依赖于对光合和非光合结构生物量分配的调节,为验证此假设,采用枝叶修剪的方式（连续两年修剪,四个修剪强度：0,20%,50%,70%）对3种亚热带乔木树种榕（Ficus microcarpa),黄桷树（Ficus virens)和樟（Cinnamomum camphora)进行了研究。结果表明,修剪使所有树种地上部分的光合与非光合结构生物量比率（P/NP）立即下降,下降程度随修剪强度的程式高而增大,但不论是首次修剪还是第二次修剪,修剪处理一年后,修剪株地上部分的光合与非光合结构生物量比率升高,且此生物量比率不低于非修剪株的光合与非光合结构生物量比率。此研究结果证实了植物地上部分光合与非光合结构间存在功能性平衡的假设,与假设一致。植株的生物量分配在修剪后发生了改变;修剪株加大了对光合结构（叶组织）的生物量分配（大量的新生产地上部分生物量被分配到光合结构）,同时却减少了对非光合结构（枝和茎）的分配,此分析格局的改变保证了光合与非光合结构功能性平衡的恢复与维持,可以认为,通过改变生物量分配格局以维持光合与非光合结构功能平衡是植物低御外来干扰和/或损伤的一种有效策略。     

Expansion Mechanisms and Functional Divergence of the Glutathione S-Transferase Family in Sorghum and Other Higher Plants

Glutathione S-transferases (GSTs) exist in various eukaryotes and function in detoxification of xenobiotics and in response to abiotic and biotic stresses. We have carried out a genome-wide survey of this gene family in 10 plant genomes. Our data show that tandem duplication has been regarded as the major expansion mechanism and both monocot and dicot plants may have practiced different expansion and evolutionary history. Non-synonymous substitutions per site (Ka) and synonymous substitutions per site (Ks) analyses showed that N- and C-terminal functional domains of GSTs (GST_N and GST_C) seem to have evolved under a strong purifying selection (Ka/Ks < 1) under different selective pressures. Differential evolutionary rates between GST_N and GST_C and high degree of expression divergence have been regarded as the major drivers for the retention of duplicated genes and the adaptability to various stresses. Expression profiling also indicated that the gene family plays a role not only in stress-related biological processes but also in the sugar-signalling pathway. Our survey provides additional annotation of the plant GST gene family and advance the understanding of plant GSTs in lineage-specific expansion and species diversification.     

Ferredoxin and Ferredoxin-NADP Reductase from Photosynthetic and Nonphotosynthetic Tissues of Tomato

Ferredoxin and ferredoxin-NADP⁺ oxidoreductase (FNR) were purified from leaves, roots, and red and green pericarp of tomato (Lycopersicon esculentum, cv VFNT and cv Momotaro). Four different ferredoxins were identified on the basis of N-terminal amino acid sequence and charge. Ferredoxins I and II were the most prevalent forms in leaves and green pericarp, and ferredoxin III was the most prevalent in roots. Red pericarp of the VFNT cv yielded variable amounts of ferredoxins II and III plus a unique form, ferredoxin IV. Red pericarp of the Momotaro cv contained ferredoxins I, II, and IV. This represents the first demonstration of ferredoxin in a chromoplast-containing tissue. There were no major differences among the tomato ferredoxins in absorption spectrum or cytochrome c reduction activity. Two forms of FNR were present in tomato as judged by anion exchange chromatography and by sodium dodecyl sulfate-polyacrylamide gel electrophoresis. FNR II had a lower apparent relative molecular weight, a slightly altered absorption spectrum, and a lower specific activity for cytochrome c reduction than FNR I. FNR II could be a partially degraded form of FNR I. The FNRs from the different tissues of tomato plants all showed diaphorase activity, with FNR II being more active than FNR I. The presence of ferredoxin and FNR in heterotrophic tissues of tomato is consistent with the existence of a nonphotosynthetic ferredoxin/FNR redox pathway to support the function of ferredoxin-dependent enzymes.     

Genome Reduction Promotes Increase in Protein Functional Complexity in Bacteria

Obligate pathogenic and endosymbiotic bacteria typically experience gene loss due to functional redundancy, asexuality, and genetic drift. We hypothesize that reduced genomes increase their functional complexity through protein multitasking, in which many genes adopt new roles to counteract gene loss. Comparisons of interaction networks among six bacteria that have varied genome sizes (Mycoplasma pneumoniae, Treponema pallidum, Helicobacter pylori, Campylobacter jejuni, Synechocystis sp., and Mycobacterium tuberculosis) reveal that proteins in small genomes interact with proteins from a wider range of functions than do their orthologs in larger genomes. This suggests that surviving proteins form increasingly complex functional relationships to compensate for genes that are lost.     

茶树CPP转录因子家族的全基因组鉴定及分析

CPP(cysteine-rich polycomb-like protein)转录因子广泛存在于动植物中,是一类成员数目较少的转录因子家族,在调控植物生长发育和响应非生物胁迫中起重要作用。该研究通过对茶树基因组系统的鉴定,获得10个CsCPP转录因子均具有典型的CXC结构域。系统发育分析将CsCPP家族成员分为4类(A^D),且大部分成员与葡萄在进化关系上更为接近。A类和C类成员的CXC结构域分布在蛋白序列的N端,而B类和D类成员分布在C端。茶树组织表达分析表明,CsCPP转录因子在生长活跃的顶芽和嫩叶中普遍高表达,不同组织的表达水平排序为:顶芽和嫩叶>根和茎>成熟叶和果>老叶和花。启动子分析发现了CsCPP家族成员的启动子区域存在大量的ABA和干旱响应元件;干旱处理下,6个CsCPP成员的表达水平均有不同程度上调,其中4个成员在ABA处理后表达迅速上调,表明CsCPP转录因子可能在ABA介导的干旱胁迫响应中起正调控作用;低温处理下,大部分CsCPP成员的表达均有轻微下调,而CsCPP 2和CsCPP 6的表达水平在6 h达到顶峰,表达量均超过2倍。研究结果为进一步发掘茶树CPP转录因子家族的功能奠定了基础。     

田间大豆叶片成长过程中的光合特性及光破坏防御机制

田间大豆叶片在成长进程中光饱和光合速率持续提高,但气孔导度的增加明显滞后.尽管叶片在成长初期就具有较高的最大光化学效率,但是仍略低于发育成熟的叶片.随着叶片的成长,光下叶片光系统Ⅱ实际效率增加;非光化学猝灭下降.幼叶叶黄素总量与叶绿素之比较高,随着叶面积的增加该比值下降,在光下,幼叶的脱环氧化程度较高.因此认为大豆叶片成长初期就能够有效地进行光化学调节;在叶片生长过程中依赖叶黄素循环的热耗散机制迅速建立起来有效抵御强光的破坏.     

Carbohydrate Metabolism in Photosynthetic and Nonphotosynthetic Tissues of Variegated Leaves of Coleus blumei Benth

Mature, variegated leaves of Coleus blumei Benth. contained stachyose and other raffinose series sugars in both green, photosynthetic and white, nonphotosynthetic tissues. However, unlike the green tissues, white tissues had no detectable level of galactinol synthase activity and a low level of sucrose phosphate synthase indicating that stachyose and possibly sucrose present in white tissues may have originated in green tissues. Uptake of exogenously supplied [¹⁴C]stachyose or [¹⁴C]sucrose into either tissue type showed conventional kinetic profiles indicating combined operation of linear first-order and saturable systems. Autoradiographs of white discs showed no detectable minor vein labelling with [¹⁴C]stachyose, but some degree of vein labeling with [¹⁴C]sucrose. Autoradiographs of green discs showed substantial vein loading with either sugar. In both tissues, p-chloromercuribenzenesulfonic acid had no effect on the linear component of sucrose or stachyose uptake but inhibited the saturable component. Both tissues contained high levels of invertase, sucrose synthase and α-galactosidase and extensively metabolized exogenously supplied ¹⁴C-sugars. In green tissues, label from exogenous sugars was recovered as raffinose-series sugars. In white tissues, exogenous sugars were hydrolysed and converted to amino acids and organic acids. The results indicate that variegated Coleus leaves may be useful for studies on both phloem loading and phloem unloading processes in stachyose-transporting species.     

Functional Activity of the Photosynthetic Apparatus in Tobacco and Arabidopsis Plants Exposed to Chilling Temperatures

Russian Journal of Plant Physiology - Photosynthetic activities of a chilling-sensitive species, Nicotiana tabacum L. (cv. Samsun), and a chilling-resistant plant, Arabidopsis thaliana Heynh. (L)...     

Functional and Structural Divergence of an Unusual LTR Retrotransposon Family in Plants

Retrotransposons with long terminal repeats (LTRs) more than 3 kb are not frequent in most eukaryotic genomes. Rice LTR retrotransposon, Retrosat2, has LTRs greater than 3.2 kb and two open reading frames (ORF): ORF1 encodes enzymes for retrotransposition whereas no function can be assigned to ORF0 as it is not found in any other organism. A variety of experimental and in silico approaches were used to determine the origin of Retrosat2 and putative function of ORF0. Our data show that not only is Retrosat2 highly abundant in the Oryza genus, it may yet be active in rice. Homologs of Retrosat2 were identified in maize, sorghum, Arabidopsis and other plant genomes suggesting that the Retrosat2 family is of ancient origin. Several putatively cis-acting elements, some multicopy, that regulate retrotransposon replication or responsiveness to environmental factors were found in the LTRs of Retrosat2. Unlike the ORF1, the ORF0 sequences from Retrosat2 and homologs are divergent at the sequence level, 3D-structures and predicted biological functions. In contrast to other retrotransposon families, Retrosat2 and its homologs are dispersed throughout genomes and not concentrated in the specific chromosomal regions, such as centromeres. The genomic distribution of Retrosat2 homologs varies across species which likely reflects the differing evolutionary trajectories of this retrotransposon family across diverse species.     

植物GH3基因家族的功能研究概况

植物GH3基因是一种典型的植物生长素原初反应基因,此类基因与植物的生长发育密切相关。GH3基因在植物生长素信号途径、光信号途径以及植物的防卫反应中起着重要作用。植物GH3蛋白具有植物生长素氨基酸化合成酶活性,这有助于维持植物生长素的动态平衡。该文介绍拟南芥等植物中GH3基因的生物学功能研究概况和最新进展,为植物GH3基因家族的进一步研究提供参考。     

植物GH3 基因家族的功能研究概况

植物GH3基因是一种典型的植物生长素原初反应基因, 此类基因与植物的生长发育密切相关。GH3基因在植物生长素信号途径、光信号途径以及植物的防卫反应中起着重要作用。植物GH3蛋白具有植物生长素氨基酸化合成酶活性, 这有助 于维持植物生长素的动态平衡。该文介绍拟南芥等植物中GH3基因的生物学功能研究概况和最新进展, 为植物GH3基因家族的进一步研究提供参考。     

A Lack of Parasitic Reduction in the Obligate Parasitic Green Alga Helicosporidium

The evolution of an obligate parasitic lifestyle is often associated with genomic reduction, in particular with the loss of functions associated with increasing host-dependence. This is evident in many parasites, but perhaps the most extreme transitions are from free-living autotrophic algae to obligate parasites. The best-known examples of this are the apicomplexans such as Plasmodium, which evolved from algae with red secondary plastids. However, an analogous transition also took place independently in the Helicosporidia, where an obligate parasite of animals with an intracellular infection mechanism evolved from algae with green primary plastids. We characterised the nuclear genome of Helicosporidium to compare its transition to parasitism with that of apicomplexans. The Helicosporidium genome is small and compact, even by comparison with the relatively small genomes of the closely related green algae Chlorella and Coccomyxa, but at the functional level we find almost no evidence for reduction. Nearly all ancestral metabolic functions are retained, with the single major exception of photosynthesis, and even here reduction is not complete. The great majority of genes for light-harvesting complexes, photosystems, and pigment biosynthesis have been lost, but those for other photosynthesis-related functions, such as Calvin cycle, are retained. Rather than loss of whole function categories, the predominant reductive force in the Helicosporidium genome is a contraction of gene family complexity, but even here most losses affect families associated with genome maintenance and expression, not functions associated with host-dependence. Other gene families appear to have expanded in response to parasitism, in particular chitinases, including those predicted to digest the chitinous barriers of the insect host or remodel the cell wall of Helicosporidium. Overall, the Helicosporidium genome presents a fascinating picture of the early stages of a transition from free-living autotroph to parasitic heterotroph where host-independence has been unexpectedly preserved.     

Molecular Properties and Functional Divergence of the Dehydroascorbate Reductase Gene Family in Lower and Higher Plants

Dehydroascorbate reductase (DHAR), which reduces oxidized ascorbate, is important for maintaining an appropriate ascorbate redox state in plant cells. To date, genome-wide molecular characterization of DHARs has only been conducted in bryophytes (Physcomitrella patens) and eudicots (e.g. Arabidopsis thaliana). In this study, to gain a general understanding of the molecular properties and functional divergence of the DHARs in land plants, we further conducted a comprehensive analysis of DHARs from the lycophyte Selaginella moellendorffii, gymnosperm Picea abies and monocot Zea mays. DHARs were present as a small gene family in all of the land plants we examined, with gene numbers ranging from two to four. All the plants contained cytosolic and chloroplastic DHARs, indicating dehydroascorbate (DHA) can be directly reduced in the cytoplasm and chloroplast by DHARs in all the plants. A novel vacuolar DHAR was found in Z. mays, indicating DHA may also be reduced in the vacuole by DHARs in Z. mays. The DHARs within each species showed extensive functional divergence in their gene structures, subcellular localizations, and enzymatic characteristics. This study provides new insights into the molecular characteristics and functional divergence of DHARs in land plants.     

The Functional State of the Photosynthetic Apparatus of Potato Plants upon Destruction of the Tubulin Cytoskeleton

Biophysics - We have investigated the effect of colchicine, a disrupting agent of the tubulin cytoskeleton, on chlorophyll fluorescence parameters, the Hill reaction, noncyclic...     

Isolation, Identification, and Activity of Mycoherbicidal Pathogens from Juvenile Broomrape Plants

Although there are reports of isolation of mycoherbicidal pathogens attacking the widespread broomrapes (Orobanche spp.) that parasitize legumes and vegetables, none is in use or available. This is despite there being no good method of controlling broomrapes in most crops other than by preplant fumigation with methyl bromide. Two highly parasitic fungi, Fusarium arthrosporioides strain E4a (CNCM I-164) and F. oxysporum strain E1d (CNCM I-1622), were isolated from nearly 100 organisms found on diseased, juvenile, emerging Orobanche flower stalks. A near-axenic polyethylene envelope system for culturing broomrape on tomato roots was used to ascertain pathogenicity of these strains. Both organisms fulfilled Koch's postulates for being primary pathogens. Their DNAs were analyzed and fingerprinted by restriction fragment length polymorphism and random amplified polymorphic DNA, showing that they are indeed different from each other and from many other Fusarium spp. and other formae speciales of F. oxysporum including a strain that attacks O. cumana on sunflowers. Both strains infect O. aegyptiaca, O. cernua, and O. ramosa, but not O. cumana. They did not infect any of the vegetable and legume crops tested and thus seem specific to Orobanche. Tomato plant roots dipped into a fungal spore and mycelial suspension and planted in broomrape-infested soil were protected for 6 weeks, as were tomato transplants in pot experiments. About 90% control was also achieved by posttransplant soil drench with fungal suspensions in pot experiments. These pathogens may be effective as seed, transplant, or soil-drench treatments of high-value vegetable and other crops.