新的C4及CAM光合途径植物

以稳定性碳同位素比(δ¹³C)鉴别禾本科、莎草科、苋科和萝摩科共46种植物的光合作用途径,发现了36种新的C₄植物(δ¹³C-10.43到-13.66‰)和1种CAM植物(δ¹³C-15.24‰)。根据Hattersley区分C₄植物三种亚类型的不同δ¹³C值,提出在36种C₄中有8种具δ₁₃C值-10.4到-10.9‰者是NADP-ME型,6种具δ¹³C值-13‰左右的是NAD-ME型,其余种类可能是NADP-ME或PCK型。     

近70年来黄土高原典型植物δ13C值变化研究

对黄土高原地区 4种典型C3 植物狼牙刺 (Sophoraviciifolia) 、辽东栎 (Quercusliaotungensis) 、虎榛子 (Os tryopsisdavidiana) 和酸枣 (Zizyphusjujubavar.spinosa) 样品稳定性碳同位素组分 (δ¹³ C) 进行分析, 研究了从 2 0世纪 30年代至今近 70年中不同年代植物δ¹³ C值的变化。结果表明, 在近 70年中, 4种植物δ¹³ C值变化范围为- 2 5.0 5‰~ - 2 9.75‰, 平均值为 - 2 7.0 4‰。 4种植物叶片δ¹³ C值均呈下降趋势, 表明随气候环境变化, 近 70年4种植物的水分利用效率 (WUE) 均呈降低趋势。但不同植物叶片δ¹³ C值下降幅度不同 :狼牙刺和辽东栎叶片δ¹³ C值下降非常明显, 虎榛子叶片δ¹³ C值下降也较明显, 而酸枣叶片δ¹³ C值下降不明显。 4种植物δ¹³ C值的降低率分别为 14.6 5 %、14.4 6 %、11.99%和 2.4 4 %, 说明不同植物对气候环境因子的敏感性不同, 具有不同的适应环境变化的策略, 酸枣是 4种植物中耐旱能力较强, WUE较高的物种。     

控温条件下C3、C4草本植物碳同位素组成对温度的响应

采取人工控制实验,探讨了6种C3、C4草本植物在昼/夜温度指标为20/12℃!36/28℃的范围内植物碳同位素组成(δ13C)及其对温度变化的响应,并结合植物比叶面积(SLA)、胞间CO2浓度(ci)与环境CO2浓度(ca)的比值、碳同化率(净光合速率Pn/胞间CO2浓度ci)等光合生长指标对植物δ13C的影响进行了分析。结果表明:所有C3、C4植物样品的δ13C值分别变化在-28.3‰!-32.1‰和-14.4‰!-17.6‰之间;在C3植物中,油菜δ13C值分布范围最集中,位于-31.1‰!-32.1‰之间;C4植物中,谷子δ13C值分布范围最窄。在控制的温度范围内,3种C3植物的平均δ13C值随温度升高而显著变低,而C4植物δ13C平均值与温度呈先增大后减小的抛物型关系,但线性回归结果未达到显著水平(P0.05)。单个植物种的δ13C值对温度的响应不同,茄子、高粱的δ13C值与温度呈线性负相关,其它4种植物与温度均呈二次抛物线关系,这可能与不同植物种具有不同的光合最适温度以及植物δ13C分馏对温度变化的敏感程度不同有关。     

黄土高原植被演替对土壤碳库及δ~(13)C的影响

采集宁夏云雾山草原植被演替阶段的土壤和植物样品,测定土壤容重(BD)、土壤含水量(WC)、土壤有机碳(SOC)含量、以及土壤和植物优势种稳定碳同位素组成(δ13C),并比较了它们之间的差异,分析其反映的环境信息。结果表明:在表层0~10 cm深度,灌木地SOC含量为71.7 g·kg-1,显著高于草地的54.6 g·kg-1;灌木地0~20 cm深度SOC含量占1 m深SOC含量的58.1%,而草地在0~40 cm深度的SOC含量占1 m深的34.1%;短期植被演替(约10 a)对土壤剖面0~20 cm层的δ13C值有显著影响,其中对0~10 cm层δ13C影响最大;草地演替为灌木地后δ13C偏负1.4‰,30 cm以下土壤δ13C值对短期植被变化不敏感;通过δ13C在C3植被的短期演替过程中的响应关系发现,δ13C值作为土壤碳库更替和碳循环的研究工具有很好的辨识力;灌木的δ13C值从叶子到根系存在明显差异,逐渐偏正,变化为2.2‰,草地为1.9‰;通过对0~40 cm深度植物根系δ13C值的测定发现,灌木地由表层的-28.15‰变为-26.11‰,偏正了2.04‰,草地根系δ13C值从表层的-27.08‰变为-27.57‰,偏负了0.49‰。     

内蒙古草原常见植物叶片δ~(13)C和δ~(15)对环境因子的响应

在中国东北样带沿线的内蒙古草原地区采集了一些常见植物的叶片样品,并测定其δ~(13)C和δ~(15)N值,分析了其统计学特征以及对环境因子(年平均降雨量和温度)的响应模式。发现东北样带草原区同时存在C_3和C_4两种不同光合途径的植物,但是C_3植物占主导地位,C_4植物数量有限。C_3植物叶片δ~(13)C随着年平均降雨量和年平均温度的升高而显著降低,反映了此区域C_3植物δ~(13)C受控于降水量和温度。C_4植物的叶片δ~(13)C值随着降雨量的增多而有轻微升高的趋势,但是C_4植物的叶片δ~(13)C值对年平均温度的响应不敏感。不论对C_3植物还是C_4植物而言,叶片δ~(15)N都随降雨量增加而显著降低,即干旱区的植物叶片δ~(15)N大于湿润地区,这说明降水是影响植物叶片δ~(15)N的一个重要因素。然而两者叶片δ~(15)N对温度的响应不敏感。     

中国暖温带落叶阔叶林中某些树种的13C自然丰度:δ13C值及其生态学意义

利用质谱分析仪对暖温带地区落叶阔叶林优势种稳定碳同位素的分析发现不同树种叶片的稳定碳同位素比率差别较大 ,大多数优势种叶片δ13C值在 -2 4.75 1‰± 0 .85 4‰～ -2 8.1 1 3‰± 1 .5 1 9‰之间。叶片的δ13C值可以分为 3个等级 , 级 ,叶片的 δ13C≥ -2 5 .5‰ , 级 ,叶片的 δ13C值在 -2 5 .5‰～ -2 7.5‰之间 , 级 ,叶片的 δ13C≤ -2 7.5‰ ,由于δ13C值在一定程度上能够反映植物的生理生态特性 ,这表明所研究的植物在生理生态特性方面也可以分为 3个类型。同时 ,由于植物的不同器官具有不同的生理生态特性 ,导致器官对 13C具有不同的分馏特性 ,也导致器官之间的δ13C值产生差异 ,分析结果显示树干、根和小枝的δ13C值一般要较叶片的δ13C值高 ,但不同树种又各具特点。生境的差异是影响稳定碳同位素比率的另一个重要原因 ,良好生境条件下生长的植物的δ13C值一般较生长在干旱瘠薄生境下的低。     

不同生活型绿化植物叶片碳同位素组成的季节特征

通过测定北京地区不同生活型绿化植物叶片的碳同位素组成(δ13C值),从植物种和生活型两个方面研究植物水分利用效率的自然可变性。结果表明:所测定的75种植物(隶属于35科65属)的叶片的δ13C值变幅,春季为-30.7‰--23.4‰,夏季为-31.5‰--25.1‰,秋季为-31.4‰--23.9‰;落叶灌木种间差异不显著(p=0.114),而常绿乔木(p=0.005)、落叶乔木(p0.001)、常绿灌木(p=0.022)、草本植物(p0.001)和藤本植物(p=0.001)的种间差异显著或极显著;同一生活型植物叶片的δ13C季节差异显著,春季叶片的δ13C值显著大于夏秋两季(常绿乔木除外),不同生活型植物叶片的δ13C值在春、夏、秋3个季节差异都达到了极显著水平(春季p=0.001、夏季p0.001、秋季p0.001),且叶片的δ13C值表现出乔木树种灌木树种藤本植物草本植物、常绿植物落叶植物的规律。因此,植物种和生活型均会引起植物叶片δ13C值的变化,但δ13C受生活型变化的影响较大,表明不同生活型植物的水分利用效率具有明显差异。     

樟子松、油松、蒙古栎水分利用效率种间变化及其对环境因子的响应差异

以碳同位素值(δ~(13)C)作为反映植物水分利用效率的指标,对辽宁地区3种典型林型—樟子松人工林(Pinus sylvestris var.mongolica)、油松人工林(Pinus tabulaeformis)和蒙古栎天然林(Quercus mongolica Fischer ex Ledebour)进行调查并分别对樟子松、油松和蒙古栎进行样品采集,探讨其水分利用效率之间的差异特征及其对年均降水量、年均大气温度和海拔高度变化的响应。樟子松、油松和蒙古栎δ~(13)C值变化范围分别为-30.37‰~-25.10‰、-30.32‰~-24.07‰和-29.85‰~-23.51‰,且随经度的增加显著降低;统计分析也表明,樟子松、油松和蒙古栎δ~(13)C值在辽西显著高于辽东,说明3种树种水分利用效率在辽西较高。进一步分析发现,3种树种叶片δ~(13)C值受年均降水量和海拔的影响较大,均随年均降水量增加显著降低,随海拔的升高显著增大,而受温度的影响较小;其中,又以樟子松的水分利用效率受海拔的影响较大(P0.01);与蒙古栎相比,樟子松和油松受降水量的影响更加明显。综上所述,3种树种叶片δ~(13)C值以及由其衍生来的水分利用效率之间种间差异较小,而受环境因子及地理学指标降雨、海拔和经度的影响较大,这为今后基于δ~(13)C值的水分利用效率进一步科学研究奠定了基础。     

荒漠植物梭梭稳定碳同位素组成与环境因子的关系

以广泛分布于新疆荒漠地区的建群种植物——梭梭(Haloxylon ammodendron)为研究对象,通过对23个样地101份梭梭同化枝样品δ~(13)C值的测定,分析了梭梭稳定碳同位素组成的变化特征及其与环境因子(海拔、日照时数、潜在蒸散量、年平均降水量和年平均温度)的关系,并讨论了不同生境下梭梭同化枝δ~(13)C值的变化特征。研究结果显示:(1)梭梭同化枝δ~(13)C平均值为-14.15‰,其在95%置信区间的变化范围为-13.14‰—-15.38‰,表明梭梭是C_4光合途径的植物。(2)梭梭同化枝δ~(13)C值与年平均降水量和年平均温度呈显著负相关关系,而与日照时数、潜在蒸散量和海拔呈显著正相关关系。我们推测梭梭同化枝δ~(13)C值对各环境因子响应趋势的不同,可能是由气孔限制因素造成的,它是梭梭适应干旱荒漠环境的一种策略。(3)在不同生境下,梭梭同化枝的碳同位素组成存在显著差异。当梭梭群落中的主要伴生种为白刺、红砂时,其δ~(13)C值最高,当主要伴生种为沙拐枣和假木贼时,其δ~(13)C值最低。在灰漠土与灰棕漠土样地中的梭梭δ~(13)C值高于棕钙土、风沙土、石质土样地;盆地中梭梭同化枝δ~(13)C值低于平原、山地、丘陵地形条件下的样地。以上结果表明:梭梭水分利用效率在不同环境梯度和生境中,存在着显著不同,表现出显著的适应策略差异。     

沙棘属植物叶片碳稳定同位素含量与气候的关系

本试验以131个沙棘属植物种群为研究对象,通过测定其叶片碳稳定同位素(δ¹³C)值,分析了碳稳定同位素特征与环境因子之间的关系。结果表明: 沙棘属植物叶片的δ¹³C值介于-24.65‰～-29.11‰,平均值为-26.97‰,属于C3植物,叶片δ¹³C值变异系数为种内大于种间,表明环境因子是影响沙棘属植物叶片δ¹³C含量变化的主导因素。沙棘属植物叶片的δ¹³C值与经纬度的变化无显著相关,与海拔呈显著负相关。通过建立回归方程: δ¹³C(‰)=0.118VAP-0.007GST-0.000028RDA-20.721(R²=0.212,P<0.0001),说明影响沙棘属叶片δ¹³C值最主要的因素是水蒸气压(VAP)、生长季温度(GST)和太阳辐射(RDA)。研究结果可为沙棘属植物对全球气候变化的响应提供理论依据。     

C4 photosynthesis in a single C3 cell is theoretically inefficient but may ameliorate internal CO2 diffusion limitations of C3 leaves

Attempts are being made to introduce C₄ photosynthetic characteristics into C₃ crop plants by genetic manipulation. This research has focused on engineering single‐celled C₄‐type CO₂ concentrating mechanisms into C₃ plants such as rice. Herein the pros and cons of such approaches are discussed with a focus on CO₂ diffusion, utilizing a mathematical model of single‐cell C₄ photosynthesis. It is shown that a high bundle sheath resistance to CO₂ diffusion is an essential feature of energy‐efficient C₄ photosynthesis. The large chloroplast surface area appressed to the intercellular airspace in C₃ leaves generates low internal resistance to CO₂ diffusion, thereby limiting the energy efficiency of a single‐cell C₄ concentrating mechanism, which relies on concentrating CO₂ within chloroplasts of C₃ leaves. Nevertheless the model demonstrates that the drop in CO₂ partial pressure, pCO₂, that exists between intercellular airspace and chloroplasts in C₃ leaves at high photosynthetic rates, can be reversed under high irradiance when energy is not limiting. The model shows that this is particularly effective at lower intercellular pCO₂. Such a system may therefore be of benefit in water‐limited conditions when stomata are closed and low intercellular pCO₂ increases photorespiration.     

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。     

Effects of Relative Humidity on Carbon Isotope Fractionation in Plants

Four C₃ plants and a C₄ plant were grown from seeds at four levels (30, 45, 60, and 75 %) of relative humidity. All plants were subjected to a 16 h day, at 500 μE/m².s^?1 photon flux density. Mature leaves were analyzed for their carbon isotopic composition. Isotope fractionation decreased by up to 3 ‰ with decreasing relative humidity in all C₃ plants, while the opposite trend was observed in the C₄ plant. The observed shifts in both C₃ and C₄ plants are attributed to decreased stomatal conductance at low relative humidity, resulting in a smaller P_i.     

判别分析方法在鉴别C_3 、C_4植物中的应用——以中国东北样带 (NECT)的研究为例

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便携式叶面积仪在野外所测定的植物生理生态参数,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。用马氏距离和后验概率判别准则进行回判,准确率达到98.48%。利用此判别模型可以根据任一植物的光合速率、蒸腾速率、气孔导度和叶温与大气温度之差4项指标判别该植物的光合碳同化功能型     

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.     

四种荒漠植物的光合响应

研究了胡杨（Populus euphratica Oliv）、疏叶骆驼刺（Alhagi sparsifolia B.Keller et Shap)、多枝柽柳（Tamarix ramosissima Lbd)和头状沙拐枣（Calligonum caput-medusae Schrenk)等四种塔克拉玛干荒漠植物的光合响应曲线,结果表明：（1）四种植物中,疏叶骆驼刺光呼吸（Rp)和CO2补偿点（Г）最高,净光合速率（An）以及水分利用效率（WUE）最低;而C4植物头状沙拐枣无论Rp还是Г都最低,它的An、光补偿点（LCP）及光饱和点（LSP）都显著高于其它3种植物（P≤0．05）,而且因为非常低的蒸腾速率（E）,沙拐枣的WUE也较高。（2）4种植物都为喜光植物,但仍然表现出一定的差异。根据它们光曲线中LCP以及暗呼吸速率（Rd),4种喜光植物的喜光顺序为：头状沙拐枣,疏叶骆驼枣刺,多枝柽柳,胡杨。（3）柽柳光合响应曲线与日变化中的An,gs等值差异较大,可能是该植物的温度生态位较其它三种植物范围较窄,设置的25℃不在其最适温度范围内,或者是在这个温度下气孔对光强变化的不敏感。（4）综合前人的结果表明,在特定的环境条件,发育阶段以及经过一定的诱导处理,胡杨可以因诱导而表现出一些C4植物特征。     

Longitudinal variation in food sources and their use by aquatic fauna along a subtropical river in Taiwan

1. River food webs rely on two major food sources: autochthonous primary production within the river and allochthonous organic matter transferred to the river. We characterised the consumer communities and assessed the food sources of dominant consumers along a subtropical mountainous river (the Lanyang River of north‐eastern Taiwan) at the catchment scale from the headwater to the estuary using natural abundances of stable carbon and nitrogen isotopes. 2. The downstream transport of fine particulate organic matter (FPOM) was two orders of magnitude greater than that of coarse particulate organic matter (CPOM). Transport of both materials increased from the headwater and reached a maximum in the midstream reach. CPOM composition exhibited a gradual shift from leaves and branches in the headwater, an area characterised by high canopy cover, to algae in the midstream reaches and marsh plants in the downstream reaches. 3. Consumer communities can be classified into two regional categories: the upland category in the headwater and upstream and midstream reaches and the lowland category comprised of samples from the downstream reach and estuary. The upland category revealed a clear and gradual seasonal shift in community composition, but a seasonal shift was not apparent for the lowland category. Nutrient concentrations and water temperature were the main factors explaining longitudinal and seasonal variations. 4. The use of sources of organic matter by dominant consumers along the Lanyang River was primarily determined by their availability. Riparian C₃ plants were the major food sources in the headwater, upstream reach and estuary, but the contribution of periphyton increased in the upper midstream reach where the river flows through an agricultural area. In the lower midstream and downstream reaches, the contribution of riparian C₄ plants became dominant. 5. The trophic transfer of organic materials in the Lanyang River may be influenced by the fast current velocity and by sewage nutrient loading in the river, both of which have important implications for predicting how the functioning of subtropical river food webs will respond to human‐related changes in land use.     

Root exudation (net efflux of amino acids) may increase rhizodeposition under elevated CO2

Increases in atmospheric CO₂ concentration ([CO₂]) can lead to global climate change and theoretically could enhance carbon (C) deposition in soil, but data on this complex issue are contradictory. One approach for clarifying the diverse forces influencing plant‐derived C in the rhizosphere involves defining how elevated [CO₂] alters the fundamental process of C transfer from plant roots to the soil. We examine here how a step increase in [CO₂] affects the innate influx and efflux components of root exudation in axenic plants, as one foundation for understanding how climate change may affect rhizodeposition. Increasing [CO₂] from 425 to 850 μmol mol^?1 during short‐term trials enhanced shoot and root dry weight (P<0.01) of annual rye grass (Lolium multiflorum Lam.) and medic (Medicago truncatula L.) but had no effect on growth of maize (Zea mays L.). Root amino‐acid flux in the same plants changed only in maize, which increased the efflux rate (nmol g root fresh weight^?1 h^?1) of six amino acids (arginine, alanine, proline, tyrosine, lysine and leucine) significantly (P<0.05) under elevated [CO₂]. None of the three plant species altered the steady‐state concentration of 16 amino acids released into a hydroponic solution with changing [CO₂], apparently because amino‐acid influx rates, measured at 2.5 μm , consistently exceeded efflux rates. Indeed, plants recovered amino acids at rates 94–374% higher than they were lost from roots regardless of [CO₂]. These results indicate that, in theory, any effect of [CO₂] doubling on amino‐acid efflux can be offset by innately higher rates of influx. In practice, however, higher rates of amino‐acid cycling (i.e., efflux+influx) for each root segment (in C₄ maize) or from more root tissue (in the two C₃ species) should increase root exudation by plants exposed to elevated [CO₂] as additional amino acids would be adsorbed to soil particles or be taken up by soil microorganisms.     

Application of Discriminant Analysis in Distinguishing Plant Photosynthetic Types-A Case Study in Northeast China Transect (NECT) Area

Discriminant analysis is an important method in multivariate statistic analysis to distinguish whatever type an individual should belong to. Based on the field actual photosynthetic data obtained from the research platform--Northeast China Transect (NECT), the concept and principle of discriminant analysis were used to distinguish the different plant photosynthetic types. A number of indices related to plant photosynthetic rate measured by a LCA4 photosynthesis system were selected to build the discriminant model. In this case study, 15 plant species from C4 plant functional groups and 51 from C3 plant functional groups were selected to build a discriminant model. The rate of accuracy, of returned classification using methods of squared Mahalanobis distances from group centroids and posterior probabilities, reached to 98.48 %. With the help of this model, any plants' photosynthetic types could be distinguished simply by using their four related parameters, viz., photosynthetic rate, transpiration, stomatal conductance and the temperature difference between leaf surface and atmosphere.