Major diffusion leaks of clamp‐on leaf cuvettes still unaccounted: how erroneous are the estimates of Farquhar et al. model parameters?

Estimates of leaf gas-exchange characteristics using standard clamp-on leaf chambers are prone to errors because of diffusion leaks. While some consideration has been given to CO(2) diffusion leaks, potential water vapour diffusion leaks through chamber gaskets have been neglected. We estimated diffusion leaks of two clamp-on Li-Cor LI-6400 (Li-Cor, Inc., Lincoln, NE, USA) leaf chambers with polymer foam gaskets and enclosing either 2 or 6 cm(2) leaf area, and conducted a sensitivity analysis of the diffusion leak effects on Farquhar et al. photosynthesis model parameters - the maximum carboxylase activity of ribulose 1 x 5-bisphosphate carboxylase/oxygenase (Rubisco) (V(cmax)), capacity for photosynthetic electron transport (J(max)) and non-photorespiratory respiration rate in light (R(d)). In addition, net assimilation rate (A(n)) versus intercellular CO(2) (C(i)) responses were measured in leaves of Mediterranean evergreen species Quercus ilex L. enclosing the whole leaf chamber in a polyvinyl fluoride bag flushed with the exhaust air of leaf chamber, thereby effectively reducing the CO(2) and water vapour gradients between ambient air and leaf chamber. For the empty chambers, average diffusion leak for CO(2), K(CO2), (molar flow rate corresponding to unit CO(2) mole fraction difference) was ca. 0.40 micromol s(-1). K(CO2) increased ca. 50% if a dead leaf was clamped between the leaf chamber. Average diffusion leak for H(2)O was ca. 5- to 10-fold larger than the diffusion leak for CO(2). Sensitivity analyses demonstrated that the consequence of a CO(2) diffusion leak was apparent enhancement of A(n) at high CO(2) mole fraction and reduction at lower CO(2) mole fraction, and overall compression of C(i) range. As the result of these modifications, Farquhar et al. model parameters were overestimated. The degree of overestimation increased in the order of V(cmax) < J(max) < R(d), and was larger for smaller chambers and for leaves with lower photosynthetic capacity, leading to overestimation of all three parameters by 70-290% for 2 cm(2), and by 10-60% for 6 cm(2) chamber. Significant diffusion corrections (5-36%) were even required for leaves with high photosynthetic capacity measured in largest chamber. Water vapour diffusion leaks further enhanced the overestimation of model parameters. For small chambers and low photosynthetic capacities, apparent C(i) was simulated to decrease with increasing A(n) because of simultaneous CO(2) and H(2)O diffusion leaks. Measurements in low photosynthetic capacity Quercus ilex leaves enclosed in 2 cm(2) leaf chamber exhibited negative apparent C(i) values at highest A(n). For the same leaves measured with the entire leaf chamber enclosed in the polyvinyl fluoride bag, C(i) and A(n) increased monotonically. While the measurements without the bag could be corrected for diffusion leaks, the required correction in A(n) and transpiration rates was 100-500%, and there was large uncertainty in Farquhar et al. model parameters derived from 'corrected'A(n)/C(i) response curves because of uncertainties in true diffusion leaks. These data demonstrate that both CO(2) and water vapour diffusion leaks need consideration in measurements with clamp-on leaf cuvettes. As plants in natural environments are often characterized by low photosynthetic capacities, cuvette designs need to be improved for reliable measurements in such species.     

Low stomatal and internal conductance to CO2 versus Rubisco deactivation as determinants of the photosynthetic decline of ageing evergreen leaves

A novel A-Ci curve (net CO2 assimilation rate of a leaf -An- as a function of its intercellular CO2 concentration -Ci) analysis method (Plant, Cell & Environment 27, 137-153, 2004) was used to estimate the CO2 transfer conductance (gi) and the maximal carboxylation (Vcmax) and electron transport (Jmax) potentials of ageing, non-senescing Pseudotsuga menziesii leaves in relation to their nitrogen (N) content and protein and pigment composition. Both gi and the stomatal conductance (gsc) of leaves were closely coupled to Vcmax, Jmax and An with all variables decreasing with increasing leaf age. Consequently, both Ci and Cc (chloroplastic CO2 concentration) remained largely conserved through successive growing seasons. The N content of leaves, as well as the amount of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and other sodium dodecyl sulfate-soluble proteins, increased during the first three growing seasons, then stabilized or decreased only slightly afterwards. Thus, the age-related photosynthetic nitrogen use efficiency (PNUE) decline of leaves was not a consequence of decreased allocation of N towards Rubisco and other proteins involved in bioenergetics and light harvesting. Rather, loss of photosynthetic capacity was the result of the decreased activation state of Rubisco and proportional down-regulation of electron transport towards the photosynthetic carbon reduction (PCR) and photorespiratory (PCO) cycles in response to a reduction of CO2 supply to the chloroplasts' stroma. This study emphasizes the regulatory potential and homeostaticity of Cc- rather than photosynthetic metabolites or Ci- in relation to the commonly observed correlation between photosynthesis and gsc.     

干旱区胡杨光合作用对高温和CO2浓度的响应

采用LI-6400便携式光合作用测定仪实测的塔里木河下游胡杨(Populus euphratica oliv)光合作用参数,探讨了不同地下水埋深下的胡杨光合作用对CO2浓度增加和温度升高的响应.结果表明:(1)CO2浓度升高减小了胡杨气孔导度,促进了光合速率、胞间CO2浓度和水分利用效率的增加,但不同地下水埋深下,胡杨光合作用参数对CO2浓度升高的响应不同,干旱环境(地下水埋深较深)下的响应程度大于水分适宜(地下水埋深浅)环境下的响应;(2) 高温引起胡杨气孔发生不完全关闭,导致了光合作用的光抑制发生,从而降低了胡杨光合速率,但降低程度受水分条件的影响,地下水埋深较深环境下的影响程度大于地下水埋深浅的;(3)地下水埋深是控制干旱区胡杨光合作用对CO2浓度和温度升高的根本因素,6m是胡杨生长正常的临界地下水埋深,地下水埋深>6m,胡杨即遭到水分胁迫,地下水埋深>7m,胡杨即受到了较严重的水分胁迫.     

Long term water stress inactivates Rubisco in subterranean clover

In long-term field experiments, during consecutive years, microswards of subterranean clover were irrigated to minimise water deficits or subjected to progressively increasing drought over 30 days. Both leaf water potential and relative water content steadily decreased during the experiments. Plants affected by drought grew more slowly and photosynthesis was decreased. Photosynthetic rate (A) and Rubisco were analysed in relation to midday water potentials and relative water contents. The difference in A between draughted and irrigated plants increased progressively, in part as a result of decreased stomatal conductance and CO₂ concentration within leaf (Ci). However, A-Ci curves suggest that the photosynthetic capacity in plants experiencing long-term stress was reduced by 50% when compared with irrigated plants. Drought decreased both the initial and the total Rubisco activity per unit area in a similar way but did not reduce the amount of Rubisco protein per unit leaf area. Thus, the specific activity of Rubisco, rather than its activation state, decreased suggesting that under water stress the active sites were blocked by inhibitors.     

杭州湾湿地不同演替阶段优势物种光合生理生态特性

采用LI-6400光合作用系统测定了杭州湾滨海湿地不同演替阶段6种优势植物-包括早期植物海三棱藨草(Scirpus mariqueter)和糙叶苔草(Carex scabrifolia)、中期植物芦苇(Phragmites communis)和柽柳(Tamarixchinensis)、后期植物白茅(Imperata cylindrica)和旱柳(Salix matsudana)的光合作用光响应曲线(LRC)和CO2响应曲线(A/Ci),并拟合得出多个光合生理指标.结果表明,6种优势植物LRC净光合速率(Pn)大小顺序为海三棱藨草>糙叶苔草>芦苇>柽柳>白茅>旱柳,且早期植物显著大于后期植物(P<0.05);光补偿点(LCP)、光饱和点(LSP)、最大净光合速率(Amax)、和暗呼吸速率(Rd)变化与Pn相同,也表现出演替早期植物>中期植物>后期植物;而表观量子效率(AQY)则表现出相反趋势.由A/Ci曲线可以发现,演替早期优势植物较后期植物具有更低的CO2羧化效率(CCE)和相对较高的CO2补偿点(CCP).可见群落演替与各阶段优势植物的光合生理特征密切相关.     

甜瓜幼苗叶片光合变化特性

为探讨甜瓜光响应变化特性与环境因子的关系,选择光响应曲线适宜测定的时段,以甜瓜幼苗为试材,将1 d分为3个时段:10:00-12:00、12:00-15:00和15:00-17:00,每个叶位叶片测定1 d,并采用直角双曲线修正模型拟合光响应曲线,研究不同时段下甜瓜叶片光响应曲线、光响应参数的变化趋势和不同叶位叶片光响应参数特性。结果表明:当环境中光合有效辐射增强,叶面温度(Tl)升高,空气相对湿度(RH)降低;当环境中光合有效辐射减弱,Tl降低,RH升高。10:00-12:00光响应曲线和12:00-15:00的第1-4叶光响应曲线呈双曲线,在强光下趋向饱和状况,12:00-15:00的第5叶光合速率和15:00-17:00光合速率在强光下出现明显的光抑制现象。1 d的不同时段均表现为10:00-12:00最大净光合速率(P_max)和光饱和点(LSP)最高,12:00-17:00降低;12:00-15:00光补偿点(LCP)和暗呼吸速率(R_d)较高,其它两个时段较低,10:00-17:00光补偿点量子效率(φ_c)、气孔导度(G_s)和胞间CO₂浓度(C_i)总体呈降低趋势,气孔限制值(L_s)升高。10:00-15:00相同时段测得的不同叶位叶片光响应参数,以第4-5叶光合性能较好,15:00-17:00以第3叶P_max最高,第5叶次之;10:00-17:00 G_s和C_i以第5叶较低,第1叶较高,L_s以第5叶较高,第1叶较低。RH为影响P_max的主要决策因子,测定时段、叶面饱和蒸汽压亏缺(Vpdl)和Tl为主要限制因子。10:00-12:00适宜进行光响应曲线测定,气孔限制为不同时段光合作用不同的主要因素,非气孔限制为影响不同叶位叶片光合作用的主要因素。     

田间冬小麦叶片光合午休过程中的非气孔限制

1　引　　言作物的光合午休是近几年作物栽培生理研究的热点之一 ,研究大都从生态、生理的各个方面阐述光合午休的成因与机理 ,如土壤水分降低 ,ＣＯ2 浓度下降 ,ＶＰＤ过高 ,气孔限制与非气孔限制 ,羧化效率降低等[4 ,6,10 ] .其中有关作物光合午休过程中的非气孔限制问题 ,多数研究仅根据Ｆａｒｑｕｈａｒ判据[2 ] 判断发生了非气孔限制即止 ,至于非气孔限制的具体内容 ,多见于讨论 ,缺乏直接的实验证据 .本文对大田生态条件下 ,冬小麦生育后期旗叶光合午休过程中的非气孔限制问题进行了较深入的研究 .2　材料与方法2 1　供试材料选用冬…     

叶肉细胞导度研究进展

叶肉细胞导度指叶片叶肉细胞内部的CO2扩散能力,在植物生理生态及全球气候变化和陆地生态系统相互关系的研究中具有重要作用。系统介绍了叶肉细胞导度的发现、发展过程及其研究进展、几种目前国际上常用的叶肉细胞导度测度方法的原理、计算过程;强调了叶肉细胞导度作为光合作用扩散过程一部分的重要意义,明确了叶肉细胞导度的定义及分布范围。并探讨了不同方法的优缺点及注意事项。总结分析了叶肉细胞导度对不同环境因子(温度、水分及环境中CO2和O3浓度等)的响应,从不同角度对叶肉细胞导度的生态学意义进行了简单的概括。对叶肉细胞导度的未来研究进行了展望。     

Rapid variations of mesophyll conductance in response to changes in CO2 concentration around leaves

The effects of short-term (minutes) variations of CO2 concentration on mesophyll conductance to CO2 (gm) were evaluated in six different C3 species by simultaneous measurements of gas exchange, chlorophyll fluorescence, online carbon isotope discrimination and a novel curve-fitting method. Depending on the species, gm varied from five- to ninefold, along the range of sub-stomatal CO2 concentrations typically used in photosynthesis CO2-response curves (AN)-Ci curves; where AN is the net photosynthetic flux and Ci is the CO2 concentrations in the sub-stomatal cavity), that is, 50 to 1500 micromol CO2 mol(-1) air. Although the pattern was species-dependent, gm strongly declined at high Ci, where photosynthesis was not limited by CO2, but by regeneration of ribulose-1,5-bisphosphate or triose phosphate utilization. Moreover, these changes on gm were found to be totally independent of the velocity and direction of the Ci changes. The response of gm to Ci resembled that of stomatal conductance (gs), but kinetic experiments suggested that the response of gm was actually faster than that of gs. Transgenic tobacco plants differing in the amounts of aquaporin NtAQP1 showed different slopes of the gm-Ci response, suggesting a possible role for aquaporins in mediating CO2 responsiveness of gm. The importance of these findings is discussed in terms of their effects on parameterization of AN-Ci curves.     

4个观赏竹种的光合特性及其影响因子分析

本研究以白纹阴阳竹( Hibanobambusa tranquillans f. shiroshima H. Okamura )、鼓节竹( Bambusa tul doides‘Swolleninternode')、花秆早竹( Phyllostachys violascens f. viridisulcata P. X. Zhang et W. X. Huang)和美丽箬竹(Indocalamus decorus Q. H. Dai)4个观赏竹种为供试材料,分析了各竹种的光响应和CO2响应曲线及参数、光合和气体交换参数以及相关环境因子的日变化规律,并通过相关性分析、逐步多元回归分析和通径分析探讨了影响4个竹种叶片净光合速率( Pn)的主要因子。测定结果表明：随光合有效辐射强度( PAR)或胞间CO2浓度( Ci)的提高,各竹种叶片的Pn值均逐渐增大,但增幅有一定差异;各竹种的光响应和CO2响应参数均有较大差异,其中,白纹阴阳竹叶片的表观量子产量( AQY)最低,CO2饱和点( CSP)和CO2补偿点( CCP)也较低,其他参数均最高;而花秆早竹叶片的AQY和CCP值均最高,而其他参数总体上均最低。4个竹种的光合及气体交换参数日变化曲线均有一定差异;美丽箬竹叶片Pn值日变化曲线呈“双峰型”;白纹阴阳竹、鼓节竹和花秆早竹叶片的Pn值日变化曲线均呈“单峰型”,峰值分别出现在14：00、10：00和12：00;总体上看,4个竹种叶片的气孔导度( Gs)、Ci、蒸腾速率( Tr)和水分利用效率( WUE)总体上具有相似的日变化趋势。分析结果表明：按照总影响效应由高至低进行排序,对白纹阴阳竹叶片Pn值的主要影响因子为PAR、Tr、大气CO2浓度( Ca)、Ci,对鼓节竹叶片Pn值的主要影响因子为PAR、Tr、Ca、Gs,对花秆早竹叶片Pn值的主要影响因子为PAR、Tr、Gs、气温( Ta)、Ci,对美丽箬竹叶片Pn值的主要影响因子为PAR、空气相对湿度( RH)、Ta、Ca、Tr、Gs。综合分析结果显示：4个竹种中,白纹阴阳竹光合能力最强,但对弱光的适应和利用能力较弱,CO2同化能力强且适应范围广;鼓节竹和美丽箬竹对光和CO2的利用能力和适应范围均居中;花秆早竹光合能力和CO2同化能力最差,但对弱光的适应和利用能力较强,CO2适应范围较窄。     

曼陀罗光合特性研究

采用LI-6400型便携式光合测定仪对潜在能源植物曼陀罗(Datura stramonium L.)在6月份至9月份的光合特性指标(净光合速率、光合有效辐射强度和气孔导度)的日变化以及净光合速率和叶绿素含量的月平均值变化进行了研究,并采用非直角双曲线对曼陀罗叶片净光合速率的光响应曲线和C02响应曲线进行了拟合.结果表明:实验期间(6月份至9月份),光合有效辐射强度和曼陀罗叶片净光合速率的日变化均呈"单峰型",峰值出现在11:00左右;7月份的净光合速率峰值最高,为19.01μmol·m-2·s-1;气孔导度日变化呈"双峰型",最大峰值和次峰值分别出现在12:00和14:00.不同月份的净光合速率月平均值有极显著差异(P＜0.01),其中7月份的净光合速率月平均值最大,为9.41 μmol·m-2·s-1;不同月份叶绿素的月平均含量变化与净光合速率月平均值的变化相似,7月份叶绿素的月平均含量也最高,但叶绿素a、b及总叶绿素月平均含量有一定差异.曼陀罗叶片净光合速率的光响应曲线和CO2响应曲线相似,在光合有效辐射强度和CO2浓度较低的条件下呈线性升高,至饱和点后缓慢升高并趋于稳定;曼陀罗叶片的光补偿点为22.42 μmol·m-2·s-1,光饱和点为689.26 μmol·m-2·s-1;CO2补偿点为74.06μmol·mol-1,CO2饱和点为1 331.97μmol·mol-1.研究结果说明,曼陀罗为具有一定耐阴能力的阳性植物,但CO2同化能力较弱.     

冬小麦对有限水分高效利用的生理机制

通过对不同土壤供水条件下的孕穗开花期的冬小麦叶片CO₂/H₂O气体交换参数的系统测定,研究了光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、细胞间隙CO₂浓度(Ci)、叶温(Tl)与水分利用效率(WUE)间的关系。结果表明,WUE并不随Pn的增长直线增长,而是呈现出二次曲线的变化趋势;只有当蒸腾达到一定程度时,Tr才对WUE产生影响,而Tr过大时WUE则有下降的趋势;WUE与Ci呈负相关,随Ci的增加WUE呈递减趋势;叶温升高对光合和蒸腾都有促进作用,当超过了某种限度则表现为抑制作用,表明在一定温度范围内,Tl升高对水分利用不利;随Gs的增大,WUE增大到一定程度则不再增加,甚至出现一种回落趋势.     

当归叶片光合参数日变化及其与环境因子的关系

在当归根茎膨大期,利用CI-310便携式光合仪,田间活体测定了当归的净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、胞间CO2浓度(Ci)等光合生理生态因子,以及光合有效辐射(PAR)、田间CO2浓度(Ca)、相对湿度(RH)、大气温度(Ta)、叶温(TL)等环境因子的日变化。结果表明,当归Pn日变化曲线为"双峰"型,最高峰出现在10:00左右,次高峰出现在16:00左右,午间有明显的"午休"现象,气孔因素是导致其"午休"的主要原因。Tr、Gs和Ci的日变化曲线均为"双峰"型,最高峰出现在10:00,次高峰出现在17:00。当归叶片净光合速率随环境因子日变化最优逐步多元回归方程为yPn=14.3108 0.0076xPAR-0.5271xTL(R2=0.6601,P<0.05),TL和PAR是对光合速率直接影响最大的生态因子,而Ca、RH和Ta主要是通过TL而间接影响光合速率的变化。     

Physiological effects of kaolin applications in well-irrigated and water-stressed walnut and almond trees

BACKGROUND AND AIMS: Kaolin applications have been used to mitigate the negative effects of water and heat stress on plant physiology and productivity with variable results, ranging from increased to decreased yields and photosynthetic rates. The mechanisms of action of kaolin applications are not clear: although the increased albedo reduces leaf temperature and the consequent heat stress, it also reduces the light available for photosynthesis, possibly offsetting benefits of lower temperature. The objective of this study was to investigate which of these effects are prevalent and under which conditions. METHODS: A 6% kaolin suspension was applied on well-irrigated and water-stressed walnut (Juglans regia) and almond (Prunus dulcis) trees. Water status (i.e. stem water potential, psi(s)), gas exchange (i.e. light-saturated CO2 assimilation rate, Amax; stomatal conductance, g(s)), leaf temperature (T(l)) and physiological relationships in treated and control trees were then measured and compared. KEY RESULTS: In both species, kaolin did not affect the daily course of psi(s) whereas it reduced Amax by 1-4 micromol CO2 m(-2) s(-1) throughout the day in all combinations of species and irrigation treatments. Kaolin did not reduce g(s) in any situation. Consequently, intercellular CO2 concentration (C(i)) was always greater in treated trees than in controls, suggesting that the reduction of Amax with kaolin was not due to stomatal limitations. Kaolin reduced leaf temperature (T(l)) by about 1-3 degrees C and leaf-to-air vapour pressure difference (VPD(l)) by about 0.1-0.7 kPa. Amax was lower at all values of g(s), T(l) and VPD(l) in kaolin-treated trees. Kaolin affected the photosynthetic response to the photosynthetically active radiation (PAR) in almond leaves: kaolin-coated leaves had similar dark respiration rates and light-saturated photosynthesis, but a higher light compensation point and lower apparent quantum yield, while the photosynthetic light-response curve saturated at higher PAR. When these parameters were used to model the photosynthetic response curve to PAR, it was estimated that the kaolin film allowed 63% of the incident PAR to reach the leaf. CONCLUSIONS: The main effect of kaolin application was the reduction, albeit minor, of photosynthesis, which appeared to be related to the shading of the leaves. The reduction in T(l) and VPD(l) with kaolin did not suffice to mitigate the adverse effects of heat and water stress on Amax.     

Diffusion and Quasi-diffusion Resistances in Relation to the Carboxylation Kinetics of Maize Leaves

A “quasi-diffusion resistance”, rq, is defined to accommodate the role which the thermochemical and photochemical phenomena of photosynthesis play in the control of CO2 fixation to the terminology and approach of the diffusion resistance analogue for the CO2 exchange of leaves. The relationship of rq to Rabinowitch's classical rectangular hyperbolic model of photosynthesis rate as a function of CO2 concen-tration at the carboxylating surface is discussed. Examination of Kmapp for phos-phopyruvate carboxylase (the predominant carboxylase in maize) suggests, as a reasonable hypothesis for light-saturated maize leaves, that rq may be essentially independent of ambient CO2-concentration up to at least 300 μ1/l. A corollary of the hypothesis is that an increase of diffusion resistance, rather than of rq, may account for the observed curvature of the response curves of light-saturated maize leaf photosynthesis to ambient CO2-concentration. An experiment carried out on fieldgrown maize plants, using a well controlled leaf chamber as a nitrous oxide diffusion porometer, gave evidence which strongly supported the hypothesis.     

Far-red photons have equivalent efficiency to traditional photosynthetic photons: Implications for redefining photosynthetically active radiation

Far-red photons (701–750 nm) are abundant in sunlight but are considered inactive for photosynthesis and are thus excluded from the definition of photosynthetically active radiation (PAR; 400–700 nm). Several recent studies have shown that far-red photons synergistically interact with shorter wavelength photons to increase leaf photochemical efficiency. The value of far-red photons in canopy photosynthesis has not been studied. Here, we report the effects of far-red photons on single leaf and canopy photosynthesis in 14 diverse crop species. Adding far-red photons (up to 40%) to a background of shorter wavelength photons caused an increase in canopy photosynthesis equal to adding 400–700 nm photons. Far-red alone minimally increased photosynthesis. This indicates that far-red photons are equally efficient at driving canopy photosynthesis when acting synergistically with traditionally defined photosynthetic photons. Measurements made using LEDs with peak wavelength of 711, 723, or 746 nm showed that the magnitude of the effect was less at longer wavelengths. The consistent response among diverse species indicates that the mechanism is common in higher plants. These results suggest that far-red photons (701–750 nm) should be included in the definition of PAR.     

Functional analysis of corn husk photosynthesis

The husk surrounding the ear of corn/maize (Zea mays) has widely spaced veins with a number of interveinal mesophyll (M) cells and has been described as operating a partial C(3) photosynthetic pathway, in contrast to its leaves, which use the C(4) photosynthetic pathway. Here, we characterized photosynthesis in maize husk and leaf by measuring combined gas exchange and carbon isotope discrimination, the oxygen dependence of the CO(2) compensation point, and photosynthetic enzyme activity and localization together with anatomy. The CO(2) assimilation rate in the husk was less than that in the leaves and did not saturate at high CO(2), indicating CO(2) diffusion limitations. However, maximal photosynthetic rates were similar between the leaf and husk when expressed on a chlorophyll basis. The CO(2) compensation points of the husk were high compared with the leaf but did not vary with oxygen concentration. This and the low carbon isotope discrimination measured concurrently with gas exchange in the husk and leaf suggested C(4)-like photosynthesis in the husk. However, both Rubisco activity and the ratio of phosphoenolpyruvate carboxylase to Rubisco activity were reduced in the husk. Immunolocalization studies showed that phosphoenolpyruvate carboxylase is specifically localized in the layer of M cells surrounding the bundle sheath cells, while Rubisco and glycine decarboxylase were enriched in bundle sheath cells but also present in M cells. We conclude that maize husk operates C(4) photosynthesis dispersed around the widely spaced veins (analogous to leaves) in a diffusion-limited manner due to low M surface area exposed to intercellular air space, with the functional role of Rubisco and glycine decarboxylase in distant M yet to be explained.     

狗尾草蒸腾特性与水分利用效率对模拟光辐射增强和CO2浓度升高的响应

 在人工控制光照强度和CO2浓度条件下,测量了禾本科C4植物狗尾草（Setaria viridis）的光合速率（Pn）,蒸腾速率（Tr）,胞间CO2浓度（Ci）,气孔导度（Gs）和叶面饱和水汽压亏缺（Vpdl）对不同模拟光辐射（SPR）强度与CO2浓度的响应。结果表明：Pn, Tr 及Gs均随SPR的升高而增大,增幅趋缓,最终趋于动态平衡。SPR增强的起始阶段,水分利用率（WUE）逐渐增大,在SPR为1200 μmol·m-2·s-1时达到最大值,然后逐渐降低。Ci与Vpdl则随SPR的增强而减小,SPR高于600 μmol·m-2·s-1之后,两者均达到平衡状态。CO2浓度从300增至600 μmol·mol-1的过程中,狗尾草Pn逐渐增大,从600增至1 000 μmol·mol-1过程中,其Pn逐渐降低。Ci、Vpdl和WUE随CO2浓度的升高而增大,Gs和Tr则随CO2浓度的升高而减小。即禾本科一年生C4植物的光合作用对CO2浓度升高响应不敏感,水分蒸腾消耗的减少和WUE的提高对CO2浓度升高的响应极显著。可见,CO2浓度升高对C4植物光合作用的直接促进作用有限,但是却能从提高现有水分利用效率途径促进植物的第一性生产。     

Effects of HgCl(2) on CO(2) dependence of leaf photosynthesis: evidence indicating involvement of aquaporins in CO(2) diffusion across the plasma membrane

Experiments were conducted to examine whether mercury-sensitive aquaporins facilitate photosynthetic CO(2) diffusion across the plasma membrane of leaf mesophyll cells. Discs without abaxial epidermes from Vicia faba leaflets were treated with HgCl(2), an inhibitor of aquaporins. Hydraulic conductivity of the plasma membrane of these discs, measured as the weight loss of the discs in the 1 M sorbitol solution, was inhibited by sub-mM concentrations of HgCl(2) by 70 to 80%. Photosynthetic CO(2) fixation was also inhibited by the HgCl(2) treatment in a similar concentration range. When 0.3 mM HgCl(2) solution was fed to the V. faba leaflets with intact epidermes via the transpiration stream, the rate of photosynthesis on leaf area basis (A) measured at photosynthetically active photon flux density of 700 micromol m(-2) s(-1) and at leaf temperature of 25 degrees C, decreased by about 20 to 30% at any CO(2) concentration in the intercellular spaces (C(i)). However, when CO(2) concentration in the chloroplast stroma (C(c)) was calculated from fluorescence and gas exchange data and A was plotted against C(c), A at low C(c) concentrations did not differ before and after the treatment. The conductance for CO(2) diffusion from the intercellular spaces to the chloroplast stroma (g(i)) decreased to 40 and 30% of the control value, when the leaflets were fed with 0.3 mM and 1.2 mM HgCl(2), respectively. Similar results were obtained with leaves of Phaseolus vulgaris. Although effects of HgCl(2) were not specific, the present results showed that HgCl(2) consistently lowered g(i). It is, thus, probable that the photosynthetic CO(2) uptake across the plasma membrane of the mesophyll cells is facilitated by mercury-sensitive aquaporins.     

Effects of carbohydrate accumulation on photosynthesis differ between sink and source leaves of Phaseolus vulgaris L

Accumulation of non-structural carbohydrate in leaves represses photosynthesis. However, the extent of repression should be different between sink leaves (sugar consumers) and source leaves (sugar exporters). We investigated the effects of carbohydrate accumulation on photosynthesis in the primary leaves of bean (Phaseolus vulgaris L.) during leaf expansion. To increase the carbohydrate content of the leaves, we supplied 20 mM sucrose solution to the roots for 5 d (sugar treatment). Plants supplied only with water and nutrients were used as controls. The carbohydrate contents, which are the sum of glucose, sucrose and starch, of the sugar-treated leaves were 1.5-3 times of those of the control leaves at all developmental stages. In the young sink leaves, the photosynthetic rate at saturating light and at an ambient CO2 concentration (A360) did not differ between the sugar-treated and control leaves. The A360 of sugar-treated source leaves gradually decreased relative to the control source leaves with leaf expansion. The initial slope of the A-Ci (CO2 concentration in the intercellular space) curve, and the Rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase) content per leaf area showed trends similar to that of A360. Differences in Amax between the treatments were slightly smaller than those in A360. These results indicate that the effect of carbohydrate accumulation on photosynthesis is significant in the source leaves, but not in the young sink leaves, and that the decrease in Rubisco content was the main cause of the carbohydrate repression of photosynthesis.