C3和C4植物光合途径的适应性变化和进化

 高等植物大多为C₃植物, C₄植物和景天酸代谢(Crassulacean acid metabolism, CAM)植物是由C₃植物进化而来的。C₄途径的多源进化表明, 光合途径由C₃途径向C₄途径的转变相对简单。该文分析研究了植物光合途径的进化前景, 指出C₄植物是从C₃植物进化而来的高光效种类, 且地质时期以来降低的大气CO₂浓度和升高的大气温度以及干旱和盐渍化是C₄途径进化的外部动力。C₃植物的C₄途径的发现说明植物的光合途径并非是一成不变的, C₃和C₄植物的光合特征具有极大的可塑性, 某些环境的变化会引起植物光合途径在C₃和C₄途径之间转变。C₃植物具有的C₄途径是环境调控的产物, 是对逆境的适应性进化结果, 因而光合途径的转变也适用于干旱地区植被的适应性生存机理研究。该文还利用国外最新的C₄光合进化模型介绍了植物在进化C₄途径中所经历的7个重要时期(从分子基础到形态基础、结构基础, 再到物质代谢水平、光合酶活水平, 直到C3和C4途径协调运转时期, 最后达到形态与功能最优化阶段), 并结合全球气候变化的特点对国内外相关领域的研究进行了分析, 总结了植物光合途径的适应性转变和进化的研究成果, 为今后的相关工作提出建议。     

C3和C4植物的氮素利用机制

提高植物的氮素利用效率(NUE)不仅有利于保障全球粮食安全, 也是实现农业可持续发展的重要途径。近半个世纪以来, 植物氮素利用机理研究已取得重要进展, 但NUE的调控机制仍不明确, NUE的提高仍然十分有限。高等植物集光合碳素同化和氮素同化于一体, 只有碳氮代谢相互协调, 才能维持植物体内的碳氮平衡, 保证植物正常生长发育。由于C₃和C₄植物的光合氮素利用率(PNUE)存在差异, 对氮素的利用效率也会存在差异。为了更有效地提高作物的NUE, 须更全面地了解C₃和C₄植物对氮素吸收、转运、同化和信号转导等关键因子的功能和调控机制。此外, 面对大气CO₂浓度增高和全球气候变暖条件下的植物碳氮同化及其机理的研究也不容忽视。该文综述了C₃和C₄植物氮素利用关键因素的差异及其调控机制, 并对提高C₃禾本科作物氮素利用效率的遗传改良途径进行了展望。     

植物近期光合碳分配及转化

碳 (C) 既是生命基础元素,又是与大气环境及全球气候变化密切相关的基本成分。近期 (系指数星期的时间尺度,约相当于植物某一生长期) 光合C是“大气-植物-土壤”系统C循环的重要组成部分;定量近期光合C在植物组织、土壤和呼吸损失的分配,对于理解全球C循环是必不可少的。植物近期光合C分结构性C和代谢C;由于代谢C周转快和结构性C尚未及时形成有机质等,近期光合C容易被忽视。应用同位素技术,可从多方面研究植物近期光合C分配及转化特征。这些方面主要包括根际区光合C代谢、近期光合C对CO₂ 和CH₄排放的贡献、C₃植物与C₄ 植物对全球C循环模式的作用、人类活动对近期光合C分配的影响等。我国鲜见植物近期光合C分配及转化的研究,开展此方面的工作,对我国从生命代谢角度研究C循环具有积极的推动作用。     

植物高光效基因工程育种

C₄植物所具有的C₄光合途径赋予其较高的光合作用效率,而一些主要的农作物如水稻、小麦、大豆等均为C₃作物,光合效率低下。随着生物技术的发展,通过基因工程手段利用C₄光合特性来改善C₃植物的光合效率进而提高其生物产量逐渐成为植物高光效育种的一个研究热点。综述了目前这一领域的研究进展及存在问题,预测了这一领域的发展前景。     

不同光合类型牧草对干旱-复水的光合生理响应及生长适应策略

基于干旱频率增加、强度增大这一全球降水变化背景, 探究干旱-复水条件下不同功能群(C₃和C₄)植物的光合生理响应及生长适应策略有助于预测降水格局变化条件下草地的植被组成和生态系统功能。该研究采用盆栽实验, 以松嫩草地生长的一年生C₃ (4种)和C₄ (3种)牧草为实验材料, 设置了对照、中度干旱和重度干旱3个水分处理水平, 在干旱末期及复水期对植物进行气体交换、生物量和比叶质量的测量。在干旱条件下, 各物种净光合速率和气孔导度均呈下降趋势, 水分利用效率呈上升趋势。干旱对不同植物光合指标的影响存在功能群差异, 随干旱程度的增加C₄植物逐渐丧失光合优势, 重度干旱对C₄植物净光合速率的影响较C₃植物更加明显。由于干旱条件下C₃植物光合固碳主要受气孔限制而C₄植物主要受代谢限制, 因此复水后C₄植物净光合速率恢复速度较C₃植物慢。干旱条件下, 各物种的生物量降低, 根冠比和比叶质量升高, 干旱对C₃植物各生长指标的影响均大于C₄植物; 复水处理后, C₃植物生物量随干旱强度增加呈下降趋势, 而C₄植物的生物量与对照相比无显著差异。     

开放式空气CO2浓度升高与作物/杂草的竞争关系

CO₂浓度升高对植物的光合作用、呼吸作用和水分利用等生理过程产生直接影响,进而影响植物的生长和繁殖.CO₂浓度升高对于具有C₃光合途径的植物较具C₄光合途径的植物更为有益.由于许多重要的杂草是C₄植物,而许多重要的作物是C₃植物,CO₂浓度升高对杂草/作物的相互关系将有重要影响.本文就全球CO₂浓度升高和气候变化对杂草/作物之间竞争关系影响进行综述,同时针对目前研究现状和可持续农业的需要,提出CO₂浓度升高条件下杂草/作物之间竞争关系及未来农田杂草治理方面理论与实践中有待解决的问题.     

东北草原区C3、C4植物的生态分布及其适应盐碱环境的生理特性

用5种实验方法对东北草原区233种植物光合类型进行鉴定,并对其相对分布随纬度变化关系及其与土壤含盐量和PH值的关系进行分析.在此基础上对几种典型C₃、C₄牧草适应于盐碱环境的生理特点进行深入研究结果表明,在所鉴定的233种植物中,C₃植物有144种,隶属于28科94属,C₄植物有89种;隶属于17科55属,在高纬度地区C₃植物表现出更高的生长优势,在纬度较低和盐碱化区域,C₄植物分布具相对优势.尤其在盐碱化程度较重的地区,C₄植物成为明显的优势种,分布上的差别决定于它们对环境适应机制上的差异C₃植物对盐碱环境适应机制主要通过积累脯氨酸等有机溶质进行渗透调节,而C₄植物主要通过液泡中离子区域化积累作用进行调节,并且与C₃植物相比对盐碱环境具更强的适应能力.     

光响应曲线的指数改进模型与常用模型比较

光响应曲线的参数是研究植物生理状态的重要指标, 常用的光响应曲线模型无法准确地计算出光饱和点和最大净光合速率。该文利用光响应曲线新模型——指数改进模型、直角双曲线模型、直角双曲线修正模型、非直角双曲线模型和指数模型, 拟合高粱(Sorghum bicolor)、苋(Amaranthus tricolor)、大麦(Hordeum vulgare)和半夏(Pinellia ternata)的光响应曲线, 并随机选取部分数据进行检验, 得到了各模型计算出的主要生理参数, 并对这些数据进行了比较分析, 讨论了各模型之间的优缺点和准确性, 描述了C₃、C₄植物光响应的适宜性。结果表明, 基于C₃植物得到的指数改进模型和直角双曲线修正模型能较准确地计算出C₃、C₄植物饱和光强和最大净光合速率, 并在描述光响应曲线时比另外3个模型具有更高的精确性和适宜性。实验结果可为光响应曲线模型在C₃和C₄光合途径植物中的应用提供参考。     

Effects of plant intraspecific variation on the prediction of C3/C4 vegetation ratio from carbon isotope composition of topsoil organic matter across grasslands

植物种内变异对草地表层有机质碳同位素组成预测C₃/C₄植被比的影响植物群落中C₃和C₄植物的比例和组成对诸多生态系统过程具有重要影响。解析C₃和C₄植物碳同位素的环境驱动过程与调控因子,对于从土壤碳同位素的角度来预测C₃/C₄植被比和组成具有重要意义。本研究旨在评估草原植物碳同位素特征的种内变异将如何影响C₃和C₄植物的碳同位素组成以及C₃/C₄植被比的预测。沿中国北方草原的自然干旱梯度选择26个植物群落,通过分析植物和土壤的碳同位素组成,采用混合模型来预测C₄植物对土壤有机碳的相对贡献。本研究对比分析了如下3种情境：(1)考虑C₃和C₄植物碳同位素的种内和种间效应;(2)仅考虑碳同位素的种间变异;(3)忽略碳同位素的种内和种间变异。研究结果表明,植物碳同位素组成沿中国北方草原自然干旱梯度的变化具有物种特异性。C₃和C₄植物的碳同位素组成与干旱指数之间呈显著负相关关系,但C₃植物比C₄植物对环境的干旱变化更为敏感。植物碳同位素特征的种内变异在驱动C₃植物功能群碳同位素沿干旱梯度的分布格局中发挥着重要作用。如果忽略植物碳同位素特征的种内变异将会显著高估C₄植物的相对贡献。本研究结果表明,草原植物碳同位素特征的种内变异对于准确预测C₃/C₄植被组成具有重要意义。     

大气NO2影响植物生长与代谢的研究进展

二氧化氮(NO₂)是大气氮氧化物之一,是大气气溶胶颗粒形成的主要成分,降低大气NO₂浓度可减轻空气中的雾霾.大气NO₂通过干沉降和湿沉降两种方式降落到植物叶片.植物吸收NO₂后主要通过两种代谢途径来降低空气中NO₂浓度: 一是主要在细胞质和叶绿体中利用还原酶的氮代谢途径,二是在质外体和细胞质中的歧化反应.植物吸收NO₂干扰了植物正常的生长和生理代谢,包括: 植物营养和生殖生长,植物体内硝酸还原酶(NaR)活性、亚硝酸还原酶(NiR)活性、氮素吸收、光合等生理代谢过程.对目前国内外有关大气NO₂影响植物生长与代谢的研究进展进行了综述,并对植物吸收NO₂的生理及分子机制的未来研究方向进行了展望.     

Climatic warming changes plant photosynthesis and its temperature dependence in a temperate steppe of northern China

Warming responses of photosynthesis and its temperature dependence in two C₃ grass (Agropyron cristatum, Stipa krylovii), one C₄ grass (Pennisetum centrasiaticum), and two C₃ forb (Artemisia capillaris, Potentilla acaulis) species in a temperate steppe of northern China were investigated in a field experiment. Experimental warming with infrared heater significantly increased daily mean assimilation rate (A) in P. centrasiaticum and A. capillaris by 30 and 43%, respectively, but had no effects on other three species. Seasonal mean A was 13, 15, and 19% higher in the warmed than control plants for P. centrasiaticum, A. capillaries, and S. krylovii, respectively. The mean assimilation rate in A. cristatum and P. acaulis was not impacted by experimental warming. All the five species showed photosynthetic acclimation to temperature. The optimum temperature for photosynthesis (T_opt) and the assimilation rate at T_opt in the five species increased by 0.33–0.78 °C and 4–27%, respectively, under experimental warming. Elevated temperature tended to increase the maximum rate of ribulose-1,5-bisphosphate (RuBP) carboxylation (V_cmax) and the RuBP regeneration capacity (J_max) in the C₃ plants and carboxylation efficiency and the CO₂-saturated photosynthetic rate in the C₄ plant at higher leaf temperature, as well as the optimum temperatures for the four parameters. Our results indicated that photosynthetic responses to warming were species-specific and that most of the species in the temperate steppe of northern China could acclimate to a warmer environment. The changes in the temperature dependence of V_cmax and J_max, as well as the balance of these two processes altered the temperature dependence of photosynthesis under climatic warming.     

Photosynthetic carbon metabolism in Panicum milioides, a C3-C4 intermediate species: Evidence for a limited C4 dicarboxylic acid pathway of photosynthesis

Panicum milioides, a naturally occurring species with C₄-like Kranz leaf anatomy, is intermediate between C₃ and C₄ plants with respect to photorespiration and the associated oxygen inhibition of photosynthesis. This paper presents direct evidence for a limited degree of C₄ photosynthesis in this C₃-C₄ intermediate species based on:

1. (a) the appearance of 24% of the total ¹⁴C fixed following 4 s photosynthesis in ¹⁴CO₂-air by excised leaves in malate and aspartate and the complete transfer of label from the C₄ acids to Calvin cycle intermediates within a 15 s chase in ¹²CO₂-air;

2. (b) pyruvate- or alanine-enhanced light-dependent CO₂ fixation and pyruvate stimulation of oxaloacetate- or 3-phosphoglycerate-dependent O₂ evolution by illuminated mesophyll protoplasts, but not bundle sheath strands; and

3. (c) NAD-malic enzyme-dependent decarboxylation of C₄ acids at the C-4 carboxyl position, C₄ acid-dependent O₂ evolution, and ¹⁴CO₂ donation from [4-¹⁴C]C₄ acids to Calvin cycle intermediates during photosynthesis by bundle sheath strands, but not mesophyll protoplasts.

However, P. milioides differs from C₄ plants in that the activity of the C₄ cycle enzymes is only 15 to 30% of a C₄ Panicum species and the Calvin cycle and phosphoenolpyruvate carboxylase are present in both cell types. From these and related studies (Rathnam, C.K.M. and Chollet, R. (1979) Arch. Biochem. Biophys. 193, 346–354; (1978) Biochem. Biophys. Res. Commun. 85, 801–808) we conclude that reduced photorespiration in P. milioides is due to a limited degree of NAD-malic enzyme-type C₄ photosynthesis permitting an increase in pCO₂ at the site of bundle sheath, but not mesophyll, ribulosebisphosphate carboxylase-oxygenase.     

Integration and expression of Sorghum C4 phosphoenolpyruvate carboxylase and chloroplastic NADP-malate dehydrogenase separately or together in C3 potato plants

We have integrated two cDNAs expressing Sorghum photosynthetic phosphoenolpyruvate carboxylase (C₄-PEPC) and NADP-malate dehydrogenase (cpMDH), two key enzymes involved in the primary carbon fixation pathway of NADP-malic enzyme-type C₄ plants, separately or together into a C₃ plant (potato). Analysis of the transgenic plants showed a 1.5-fold increase in PEPC and cpMDH activities compared to untransformed plants. Immunolocalization confirmed an increase at the protein level of these two enzymes in the transgenic plants and indicated that the Sorghum cpMDH was specifically addressed to the chloroplasts of potato mesophyll cells. However, integration of either or both of the cDNAs into the potato genome did not appear to significantly modify either tuber starch grain content or the rate of photosynthetic O₂ production compared to control untransformed plants. The low level of transgene expression probably explains the lack of influence on carbon metabolism and photosynthetic rates. This general observation suggests that some complex mechanism may regulate the level of production of foreign C₄ metabolism enzymes in C₃ plants.     

The evolutionary relationship between stomatal mechanism, crassulacean acid metabolism and C4 photosynthesis

Abstract. All of the features of crassulacean acid metabolism (CAM) and most characteristics of C₄ photosynthesis are exhibited by stomatal guard cells. It is proposed that CAM and possibly also C₄ photosynthesis result from the expression in photosynthetic cells of genetic information which is expressed only in guard cells of C₃ plants.     

Seasonal differences in photosynthesis between the C3 and C4 subspecies of Alloteropsis semialata are offset by frost and drought

The regional abundance of C₄ grasses is strongly controlled by temperature, however, the role of precipitation is less clear. Progress in elucidating the direct effects of photosynthetic pathway on these climate relationships is hindered by the significant genetic divergence between major C₃ and C₄ grass lineages. We addressed this problem by examining seasonal climate responses of photosynthesis in Alloteropsis semialata , a unique grass species with both C₃ and C₄ subspecies. Experimental manipulation of rainfall in a common garden in South Africa tested the hypotheses that: (1) photosynthesis is greater in the C₄ than C₃ subspecies under high summer temperatures, but this pattern is reversed at low winter temperatures; and (2) the photosynthetic advantage of C₄ plants is enhanced during drought events. Measurements of leaf gas exchange over 2 years showed a significant photosynthetic advantage for the C₄ subspecies under irrigated conditions from spring through autumn. However, the C₄ leaves were killed by winter frost, while photosynthesis continued in the C₃ plants. Unexpectedly, the C₄ subspecies also lost its photosynthetic advantage during natural drought events, despite greater water-use efficiency under irrigated conditions. This study highlights previously unrecognized roles for climatic extremes in determining the ecological success of C₃ and C₄ grasses.     

Photosynthetic traits in C3 and C4 grassland species in mesocosm and field environments

The North American tallgrass prairie is composed of a diverse mix of C₃ and C₄ plant species that are subject to multiple resource limitations. C₄ grasses dominate this ecosystem, purportedly due to greater photosynthetic capacity and resource-use efficiency associated with C₄ photosynthesis. We tested the hypothesis that intrinsic physiological differences between C₃ and C₄ species are consistent with C₄ grass dominance by comparing leaf gas exchange and chlorophyll fluorescence variables for seven C₄ and C₃ herbaceous species (legumes and non-legumes) in two different settings: experimental mesocosms and natural grassland sites. In the mesocosms, C₄ grasses had higher photosynthetic rates, water potentials and water-use efficiency than the C₃ species. These differences were absent in the field, where photosynthetic rates declined similarly among non-leguminous species. Thus, intrinsic photosynthetic advantages for C₄ species measured in resource-rich mesocosms could not explain the dominance of C₄ species in the field. Instead, C₄ dominance in this ecosystem may depend more on the ability of the grasses to grow rapidly when resources are plentiful and to tolerate multiple limitations when resources are scarce.     

华南地区3个棕榈藤种光合途径的判别

C3植物uT以通过转入C4植物基因而具备C4植物光合特性,从而提高产量.有鉴于此,本文通过测定我国华南地区分布的黄藤(Daemonorops margarttae(Hance)Becc.)、单叶省藤(Calamus simplicifolius C.F.Wei)和白藤(C.tetradactylus Hance)等3个棕榈藤种苗木和成年植株叶片的叶绿素含量、气孔密度、磷酸烯醇式丙酮酸羧化酶(phosphoenolpyruvate carboxylase,PEPC)、丙酮酸磷酸二激酶(pyruvate phosphate dikinase,PPDK)和稳定碳同位素比值等指标以判别3个藤种的光合途径,为棕榈藤转入C4植物基因工作提供理论依据.结果表明,3种藤种苗木和成年植株的叶绿素含量和气孔密度比常见C3植物和C4植物高,但叶绿素a/b值、叶片上下表面气孔比值、PEPC酶活性、PPDK酶活性和稳定碳同位素比值等指标均较低,与常见C3植物的对应指标相当或略低,而远小于常见C4植物,因此认为黄藤、单叶省藤和白藤等3个藤种是C3植物.     

转玉米PEPC基因水稻中有限的C4光合微循环及其生理作用

用转PEPC基因水稻(Oryza sativa L. subsp.japonica Kitaake)和原种水稻Kitaake为材料,研究了不同基因型水稻叶片中的C4光合微循环及其功能.通过测定与光合C4途径有关的关键酶,如磷酸烯醇式丙酮酸羧化酶(PEPC)、NADP -苹果酸酶(NADP -ME)、NADP -苹果酸脱氢酶(NADP -MDH)和丙酮酸磷酸双激酶(PPDK),说明原种水稻叶片中具有完整的C4光合酶体系;用外源OAA或MA饲喂叶切片或叶绿体后明显增加光合速率,证明原种水稻中具有一个有限的光合C4微循环.将玉米的PEPC基因导入原种水稻后,可大幅度提高光合C4微循环的速率.测定不同基因型的CO2交换速率,看出水稻中C4光合微循环的增强有提高净光合速率(Pn)和降低光呼吸速率/净光合速率(Pr/Pn)比值的作用.叶绿素荧光特性分析表明,C4光合微循环的增强伴随着PSⅡ电子传递效率(Fv/Fm)和光化学猝灭(qP)的增加以及非光化学猝灭(qN)的降低;这些结果为通过基因工程手段提高作物光合效率的遗传育种提供了科学根据.