中国东北样带羊草群落C3和C4植物功能群生物量及其对环境变化的响应

在全球动态植被模型的发展中,受限于人力、物力和财力使得在物种水平上的研究变得既不可能也无必要。植物功能群的划分是从生态学的,而不是系统发育的角度来相互比较地对待不同地区的植物,从而削减了植被变化研究中植物分类群的数量,已成为研究植被变化及生物多样性对生态系统功能作用的重要单位。植物的不同光合途径（C3、C4和CAM）从叶片组织结构到生理功能,从生态适应到地理分布均表现出对不同水、热、光环境的响应,是理想的植物功能群分类。为此,分析了中国东北样带以羊草（Leymus chinensis）为建群种或共建种的草原群落植物光合类型功能群生物量及其与群落初级生产力和环境变化的关系。结果表明：（1）C4植物生物量具有明显的变化规律,且对环境变化的响应显著,其变异性较高,更能反映样地问环境变化的差异;（2）C4与C3植物变化具有明显的互补性,并且多数C4植物常在逆境中起到更大作用,如干旱化、盐碱化和放牧干扰;（3）C4植物种类少,在所有调查样方中仅出现7种,占总出现种类的9．72％。这些特点说明C4植物可以考虑作为评估和预测我国温带草原植被及其生态系统变化的重要植物功能群。     

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。     

中国东北样带草地群落放牧干扰植物多样的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带９个草地群落放牧干扰植物多样性性变化的研究结果表明：中牧或重牧阶段Ｓhannon指数达最大值。形成中牧（重牧）＞重牧（中牧）＞轻牧＞过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率。而丰富度呈下降趋势,即轻牧（中牧）＞中牧（轻牧）＞重牧＞过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原＞典型草原＞典型草原＞荒漠草原＞碱化草甸,并且群落物种丰富度对多样性有更大贡献率。     

C4荒漠植物猪毛菜与木本猪毛菜的叶片解剖结构及光合生理特征

 为了比较C₄荒漠植物猪毛菜(Salsola collina)和木本猪毛菜(S. arbuscula)的抗旱结构和适应环境的光合作用特征, 在二者混生的群落中, 选择代表性植株, 采集叶片进行叶片解剖结构分析, 在自然条件下测定了二者叶片的气体交换参数。研究结果表明：猪毛菜叶片具表皮毛, 具有更发达的薄壁贮水组织;木本猪毛菜叶片具有更厚的角质层, 表皮下有1层下皮细胞, 其栅栏组织细胞较长, 排列更紧密。猪毛菜的净光合速率明显高于木本猪毛菜, 日平均值分别为21.5和15.7 μmol CO₂·m^–2·s^–1。猪毛菜的蒸腾速率也明显高于木本猪毛菜, 日平均值分别为14.9和10.2 mmol·m^–2·s^–1。猪毛菜和木本猪毛菜的水分利用效率的日平均值分别为1.39和1.53 μmol CO₂·mmol^–1 H₂O, 特别是在14:00时分别为1.61和2.30 μmol CO₂·mmol^–1 H₂O, 木本猪毛菜高出猪毛菜约42%。猪毛菜的光补偿点低于木本猪毛菜, 而光饱和点和光量子效率较高, 具有更低的CO₂补偿点。这表明：二者的旱生结构不同, 木本猪毛菜具有更显著的荒漠植物特征;在适于二者混生的环境下, 猪毛菜比木本猪毛菜的光合能力更强, 而木本猪毛菜的水分利用效率更高。     

3种C3木本植物绿色组织C4酶活性、色素含量及叶绿素荧光参数的比较

对3种C₃木本植物丁香（Syringa meyeri）、杨树（Populus alba）、落叶松（Larix gmelinii）叶片和树枝绿色组织相关指标分析发现：3种植物叶片Chl.a+b含量均显著高于树枝,但是树枝较叶片Chl.a/b比值差异不明显（丁香、落叶松）（p>0.05）或显著高（杨树）(p<0.05)。单位叶绿素内各酶活性显示,丁香树枝PEPC、PEPCK、PPDK、NADP-ME、NADP-MDH酶活性分别是叶片的2.4、4.1、3.2、4.5和1.9倍;杨树树枝分别是叶片的2.1、15、6.3、6.3和2.8;落叶松树枝分别是叶片的6.8、6.3、4.3、4和5.5倍,而单位鲜重内各酶活性显示出的树枝与叶片的差异均小于单位叶绿素的相应结果。种间比较发现,杨树叶片和树枝5种酶活性均显著高于丁香和落叶松。树枝绿色组织内C₄酶活性普遍高于相应叶片的事实说明,木质化绿色组织内可能存在C₄酶相关的生化过程。丁香与落叶松树枝的电子传递速率、PSⅡ效率大于叶片,杨树叶片的相应值大于树枝,相关分析表明,这种高的C₄酶活性并没有显著影响叶绿素荧光参数（p>0.05）,显示出树枝绿色组织和C₃植物叶片内C₄酶对光合影响的复杂性。     

超高产杂交稻两优培九的光合作用、光抑制和C4途径酶特性

比较研究了超高产杂交稻(Oryza sativa L.)两优培九和常规杂交稻汕优63剑叶的光合气体交换、光抑制, 以及其剑叶和稃片中的C₄酶活性. 发现两优培九和汕优63的光饱和同化速率(Asat)与汕优63相差无几, 但是, 其表观量子效率(AQY)、羧化效率(CE)和CO₂同化的量子效率(FCO₂)都远远高于汕优63. 强光条件下, 超高产杂交稻两优培九表现出比汕优63更强的抗光抑制能力, 并且具有较高的以叶黄素循环为量度的非辐射能量耗散. 此外, 两优培九的剑叶和稃片中的C₄途径酶活性均高于汕优63. 这些结果表明, 较高的光能和CO₂利用效率、较强的抗光抑制能力, 以及剑叶和稃片中C₄途径的较高表达可能是超高产杂交稻两优培九高产的重要保证.     

中国东北样带草地群落放牧干扰植物多样性的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带9个草地群落放牧干扰植物多样性变化的研究结果表明：中牧或重牧阶段Shannon指数达最大值,形成中牧（重牧）>重牧（中牧）>轻牧>过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率,而丰富度呈下降趋势,即轻牧（中牧）>中牧（轻牧）>重牧>过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原>典型草原>典型草原 >荒漠草原>碱化草原,并且群落物种丰富度对多样性有更大贡献率。     

差别分析方法在鉴别C3,C4植物中的应用——以中国东北样带（NECT） …

判别分析是多元统计分析中判断个体所属类型的一种重要方法。以中国东北样带（ＮＥＣＴ）作为研究平台,利用判别分析鉴别植物光合功能型。采用国际上先进的植物光合测定系统ＬＣＡ４便携式光合仪和ＣＩＤ－２０３便携式嘿面积仪在野外所测定的植物生理生态参,选取５１个来自Ｃ３功能群的植物种和１５个来自Ｃ４功能群的植物种构建判别模型,进行光合碳同化途径的判别。     

浓度NO对高表达转玉米C4型pepc水稻光合的促进

为了揭示C₃植物中C₄ pepc高表达带来的生理差异与其高光合效率的关系。本文以高表达的转玉米C₄ pepc光合基因水稻（PC）及原种Kitaake(WT）为材料,通过水培在孕穗期通过根吸入的方法,进行不同浓度的NO供体、NO合成抑制剂以及相关影响信号分子的试剂单独和联合过夜处理12 h,选取倒二叶研究NO对供试材料净光合速率（P_n）,气孔导度(G_s)和胞间CO₂浓度(C_i)的影响。结果表明：WT和PC在200 μmol·L^-1 SNP（Sodium nitroprusside）和1 mmol·L^-1 L-Arg(L-Arginine)处理下,P_n分别增加20.8%、10.7%,差异显著（p<0.05）;随SNP和L-Arg浓度的增加,其表现不同程度的抑制,与PC相比,WT的P_n抑制更显著（p<0.05）,而G_s和C_i的变化则相反（p<0.05）;进一步结合200 μmol·L^-1 SNP和1 mmol·L^-1 L-Arg与SA处理,结果与高浓度的NO供体处理类似;在联合6 mmol·L^-1 Ca²⁺螯合剂EGTA处理下,与PC相比,WT的P_n抑制达到极显著水平（p<0.01）,C_i的变化则相反（p<0.05）。相关分析结果表明：PC的P_n的高低与G_s的相关性小于WT,PC与WT决定系数分别为0.654 9、0.773 5;而与C_i的相关性则更大些,PC与WT决定系数分别为0.466 5、0.419 6,显示PC可能有不同的调节方式,尤其在低浓度的NO,PC可在Ca²⁺参与下调节气孔的开放,在气孔关闭的条件下,仍能维持一定的P_n。     

锡林河流域羊草草原植物种群和功能群的长期变异性及其对群落 …

分析了内蒙古锡林河流域羊草（Ｌｅｙｍｕｓ ｃｈｉｎｅｎｓｉｓ）草原群落植物种群、功能群和群落生物量的长期变异性,以及植物功能群间的补偿作用对群落稳定性的影响。结果表明：从种群水平到功能群落水平,地上生物量的年度间变异性逐渐降低,而稳定性则逐渐增加。植物功能群组成对群落稳定性具有更强烈的影响,在植物生活型功能群组成中多年生根茎禾草与多年生丛生禾草和多年生杂类草功能群,在生物量的年度变化上具有补偿作用     

中国东北样带关键种羊草水分利用效率与气孔密度

沿中国东北样带, 从长春到阿巴嘎旗, 大约每个经度设1个样地, 共10个样地, 研究了草原段关键种羊草的水分利用效率、叶片下表面气孔密度和气孔大小的变化, 以及它们与年降水量、年均温度、土壤水分和海拔的关系. 结果表明: 自东向西, 随年降水量、年均温度、土壤水分的降低和海拔的升高, 羊草水分利用效率有明显增强的趋势,而气孔密度有明显增大的趋势, 气孔大小没有明显变化规律并主要与取样时刻开张状态有关, 说明羊草气孔数量的增多有利用提高水分利用效率. 逐步回归分析结果表明, 土壤水分是决定羊草气孔密度变化的第一显著因子, 其次是年降水量, 说明水分条件是羊草气孔密度变化的主要生态因子. 从数据分布散点图、温度和放牧干扰作用的复杂性以及受土壤和地形条件特殊性影响样地的分析表明, 羊草水分利用效率和气孔密度对环境变化的响应是非常复杂的, 是包括植被利用方式与强度在内的环境因子综合作用的结果, 也是羊草长期适应各样地环境的结果.     

青海湖北岸环境梯度上植物群落的生物量与物种多样性及其相互关系

研究在土壤含水量（Ｘ１）、土壤ｐＨ（Ｘ２）、全盐量（Ｘ３）和海拔高程（Ｘ４）等４个主要环境梯度上,分析了青海湖北岸１８个草本植物的生物量、物种多样性（以物工为测度指标）变化规律,以及其基于环境梯度的相互关系,并提出了相关模型。研究结果表明,植物群落地上生物量与物种丰富度之间没有明显的相关性（Ｒ＾２＝０．１０４）,而物种丰富度（Ｙ）与环境因子之间的多元线性回归关系显著,回归模型为：Ｙ＝－８８．２９－     

鼎湖山针阔叶混交林冠层下方CO2通量及其环境响应

精确估算典型森林生态系统冠层下方CO₂通量（Fcb）对验证陆地生态系统碳平衡模型具有重要意义。采用开路涡度相关法对鼎湖山针阔叶混交林Fcb进行定位测定,根据1周年数据分析Fcb及其对环境要素的响应特征,结果表明：(1) 白天Fcb呈下降趋势表明地表植被全年具有光合能力,但总体上地表植被和土壤表现为CO2排放源;(2) Van’t Hoff方程、Arrhenius方程和Lloyd-Taylor方程均可以较好反映土壤温度(Ts)与Fcb的关系,其中仅Lloyd-Talor方程能够反映温度因子敏感性指标Q10随温度的变异性特征;(3) Lloyd-Talor方程模拟的Fcb完全由Ts控制,而连乘模型由Ts和土壤水分(Ms)控制,可以反映水热条件的综合影响,对Fcb具有更强的拟合能力;(4)在Ms较大时连乘模型对Fcb的估算高于Lloyd-Talor方程,反之在干旱时段连乘模型模拟结果低于Lloyd-Talor方程,表明当存在水分胁迫时,Ms可以成为影响Fcb的主导因子;(5) 2003年鼎湖山针阔叶混交林Fcb总量((787.4±296.8)gCm^-2a^-1)比静态箱-气相色谱法测得的土壤呼吸偏低17%。与箱式法相比,涡度相关法通量测定结果普遍存在偏低估算现象。     

Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2

To model the effect of increasing atmospheric CO2 on semi-arid grasslands, the gas exchange responses of leaves to seasonal changes in soil water, and how they are modified by CO2, must be understood for C3 and C4 species that grow in the same area. In this study, open-top chambers were used to investigate the photosynthetic and stomatal responses of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) grown at 360 (ambient CO2) and 720 micro mol mol-1 CO2 (elevated CO2) in a semi-arid shortgrass steppe. Assimilation rate (A) and stomatal conductance (gs) at the treatment CO2 concentrations and at a range of intercellular CO2 concentrations and leaf water potentials (psileaf) were measured over 4 years with variable soil water content caused by season and CO2 treatment. Carboxylation efficiency of ribulose bisphosphate carboxylase/oxygenase (Vc,max), and ribulose bisphosphate regeneration capacity (Jmax) were reduced in P. smithii grown in elevated CO2, to the degree that A was similar in elevated and ambient CO2 (when soil moisture was adequate). Photosynthetic capacity was not reduced in B. gracilis under elevated CO2, but A was nearly saturated at ambient CO2. There were no stomatal adaptations independent of photosynthetic acclimation. Although photosynthetic capacity was reduced in P. smithii growing in elevated CO2, reduced gs and transpiration improved soil water content and psileaf in the elevated CO2 chambers, thereby improving A of both species during dry periods. These results suggest that photosynthetic responses of C3 and C4 grasses in this semi-arid ecosystem will be driven primarily by the effect of elevated CO2 on plant and soil water relations.     

Leaf vascular systems in C(3) and C(4) grasses: a two-dimensional analysis

BACKGROUND AND AIMS: It is well documented that C(4) grasses have a shorter distance between longitudinal veins in the leaves than C(3) grasses. In grass leaves, however, veins with different structures and functions are differentiated: large longitudinal veins, small longitudinal veins and transverse veins. Thus, the densities of the three types of vein in leaves of C(3) and C(4) grasses were investigated from a two-dimensional perspective. METHODS: Vein densities in cleared leaves of 15 C(3) and 26 C(4) grasses representing different taxonomic groups and photosynthetic subtypes were analysed. KEY RESULTS: The C(4) grasses had denser transverse veins and denser small longitudinal veins than the C(3) grasses (1.9 and 2.1 times in interveinal distance), but there was no significant difference in large longitudinal veins. The total length of the three vein types per unit area in the C(4) grasses was 2.1 times that in the C(3) grasses. The ratio of transverse vein length to total vein length was 14.3 % in C(3) grasses and 9.9 % in C(4) grasses. The C(3) grasses generally had greater species variation in the vascular distances than the C(4) grasses. The bambusoid and panicoid C(3) grasses tended to have a denser vascular system than the festucoid C(3) grasses. There were no significant differences in the interveinal distances of the three vein types between C(4) subtypes, although the NADP-malic enzyme grasses tended to have a shorter distance between small longitudinal veins than the NAD-malic enzyme and phosphoenolpyruvate carboxykinase grasses. CONCLUSIONS: It seems that C(4) grasses have structurally a superior photosynthate translocation and water distribution system by developing denser networks of small longitudinal and transverse veins, while keeping a constant density of large longitudinal veins. The bambusoid and panicoid C(3) grasses have a vascular system that is more similar to that in C(4) grasses than to that in the festucoid C(3) grasses.     

Changes in CH4 and CO2 emissions from soils under flooded conditions after being exposed to FACE (free-air CO2 enrichment) for three years

To evaluate the variations of CO₂ and CH₄ emissions from FACE (free-air CO₂ enrichment, F) soils three years after rice-wheat rotation FACE treatment, incubation experiments in the laboratory with laboratory and elevated CO₂ concentration (1000 μl L?1) were carried out under flooded conditions at 25°C. Results show that soil organic carbon is increased by 11% after exposure to FACE treatment for three years. The results indicate that at laboratory and elevated CO₂, the cumulative CO₂ emissions from FACE soils are 35% and 22% higher than those from the ambient soils, whereas the cumulative CH₄ emissions from FACE soils are 2.6 and 2.3 times that of ambient soils. Thus, there is a larger ratio of cumulative emissions of CH₄ to CO₂ in the soil F. The elevated CO₂ concentration during the incubation stimulates the cumulative CO₂ emission significantly, but its stimulation on CH₄ emission is not statistically significant. The results indicate that the elevated atmospheric CO₂ concentration stimulates the turnover rates of soil organic matter, with a net increase in soil organic matter content, and alters the CH₄/CO₂ ratio.     

Differential sensitivity of C3 and C4 turfgrass species to increasing atmospheric vapor pressure deficit

Temperature and vapor pressure deficit (VPD) effects on turfgrass growth are almost always confounded in experiments because VPD commonly is substantially increased in elevated-temperature treatments. The objective of this study as to examine specifically the influence of VPD on transpiration response of four ‘warm-season’ (C₄) and four ‘cool-season’ (C₃) turfgrasses to increasing VPD at a stable temperature (29.3 ± 1.5 °C). Although transpiration rates were noticeably lower in C₄ grasses, transpiration rates increased linearly in response to increasing VPD across the range of 0.8–3.0 kPa. In contrast, transpiration rates of C₃ increased sharply with increasing VPD across the range of low VPDs, but became constrained at higher VPDs (>1.35 kPa). Restricted transpiration rate at elevated VPD was most evident in Agrostis palustris and Lolium perenne. Assuming restricted transpiration rates reflect a limitation on leaf CO₂ uptake, these results indicate that the commonly observed decline in growth of C₃ (and success of C₄) grasses at elevated temperature may include a sensitivity to elevated VPD.     

Photosynthetic and morphological functional types from different steppe communities in Inner Mongolia, North China

Morphological functional types and photosynthetic pathway types were identified for the forage species from steppe communities in Inner Mongolia, China, using the data of both field survey and published papers. Seven typical steppe communities were selected to investigate the morphological functional type and photosynthetic pathway type compositions and plant functional type (PFT) diversity in steppe communities at regional scale. Morphological functional types, based on plant height and leaf type combined with life span, were optimal for comparing the community differences in the region, while photosynthetic pathway types were fairly coarse for such studies. Of the seven morphological functional types in the steppe communities, perennial forbs (PEF) were the dominant type, and 60 % of species belonged to this type. Each of the high perennial grass (HPG), short perennial grass (SPG), and annual grass (ANG) types represented less than 10 % of the total, even though the grass species were dominant in the seven steppe communities. The differences of PFTs between the steppe communities were remarkable, and the PFT richness and diversity increased from the communities with moist conditions to the ones with dry environments.This revised version was published online in March 2005 with corrections to the page numbers.