How do leaf hydraulics limit stomatal conductance at high water vapour pressure deficits?

A reduction in leaf stomatal conductance (g) with increasing leaf-to-air difference in water vapour pressure (D) is nearly ubiquitous. Ecological comparisons of sensitivity have led to the hypothesis that the reduction in g with increasing D serves to maintain leaf water potentials above those that would cause loss of hydraulic conductance. A reduction in leaf water potential is commonly hypothesized to cause stomatal closure at high D. The importance of these particular hydraulic factors was tested by exposing Abutilon theophrasti, Glycine max, Gossypium hirsutum and Xanthium strumarium to D high enough to reduce g and then decreasing ambient carbon dioxide concentration ([CO2]), and observing the resulting changes in g, transpiration rate and leaf water potential, and their reversibility. Reducing the [CO2] at high D increased g and transpiration rate and lowered leaf water potential. The abnormally high transpiration rates did not result in reductions in hydraulic conductance. Results indicate that low water potential effects on g at high D could be overcome by low [CO2], and that even lower leaf water potentials did not cause a reduction in hydraulic conductance in these well-watered plants. Reduced g at high D in these species resulted primarily from increased stomatal sensitivity to [CO2] at high D, and this increased sensitivity may mediate stomatal responses to leaf hydraulics at high D.     

Elevated CO2 induces physiological, biochemical and structural changes in leaves of Arabidopsis thaliana

Leaves of Arabidopsis thaliana grown under elevated or ambient CO2 (700 or 370 micromol mol(-1), respectively) were examined for physiological, biochemical and structural changes. Stomatal characters, carbohydrate and mineral nutrient concentrations, leaf ultrastructure and plant hormone content were investigated using atomic absorption spectrophotometry, transmission electron microscopy and enzyme-linked immunosorbent assay (ELISA). Elevated CO2 reduced the stomatal density and stomatal index of leaves, and also reduced stomatal conductance and transpiration rate. Elevated CO2 increased chloroplast number, width and profile area, and starch grain size and number, but reduced the number of grana thylakoid membranes. Under elevated CO2, the concentrations of carbohydrates and plant hormones, with the exception of abscisic acid, increased whereas mineral nutrient concentrations declined. These results suggest that the changes in chloroplast ultrastructure may primarily be a consequence of increased starch accumulation. Accelerated A. thaliana growth and development in elevated CO2 could in part be attributed to increased foliar concentrations of plant hormones. The reductions in mineral nutrient concentrations may be a result of dilution by increased concentrations of carbohydrates and also of decreases in stomatal conductance and transpiration rate.     

Determination of the site of CO₂ sensing in poplar: is the area-based N content and anatomy of new leaves determined by their immediate CO₂ environment or by the CO₂ environment of mature leaves?

Exposure to an elevated CO(2) concentration ([CO(2)]) generally decreases leaf N content per unit area (N(area)) and stomatal density, and increases leaf thickness. Mature leaves can 'sense' elevated [CO(2)] and this regulates stomatal development of expanding leaves (systemic regulation). It is unclear if systemic regulation is involved in determination of leaf thickness and N(area)-traits that are significantly correlated with photosynthetic capacity. A cuvette system was used whereby [CO(2)] around mature leaves was controlled separately from that around expanding leaves. Expanding leaves of poplar (Populus trichocarpa×P. deltoides) seedlings were exposed to elevated [CO(2)] (720 μmol mol(-1)) while the remaining mature leaves inside the cuvette were under ambient [CO(2)] of 360 μmol mol(-1). Reverse treatments were performed. Exposure of newly developing leaves to elevated [CO(2)] increased their thickness, but when mature leaves were exposed to elevated [CO(2)] the increase in thickness of new leaves was less pronounced. The largest response to [CO(2)] was reflected in the palisade tissue thickness (as opposed to the spongy tissue) of new leaves. The N(area) of new leaves was unaffected by the local [CO(2)] where the new leaves developed, but decreased following the exposure of mature leaves to elevated [CO(2)]. The volume fraction of mesophyll cells compared with total leaf and the mesophyll cell density changed in a manner similar to the response of N(area). These results suggest that N(area) is controlled independently of the leaf thickness, and suggest that N(area) is under systemic regulation by [CO(2)] signals from mature leaves that control mesophyll cell division.     

胡杨叶片气孔导度特征及其对环境因子的响应

依据2005年对极端干旱区荒漠河岸林胡杨的观测资料,对胡杨气孔运动进行了分析研究以揭示胡杨的水分利用特征与抗旱机理。结果表明:(1)胡杨叶片气孔导度日变化呈现为周期波动曲线,其波动周期为2 h,傍晚(20:00)波动消失;净光合速率和蒸腾速率与气孔导度的波动相对应而呈现同步周期波动。(2)胡杨的阳生叶气孔导度高于阴生叶,且不同季节气孔导度值不同,阳生叶气孔导度的季节变幅大于阴生叶。(3)胡杨气孔导度与气温、相对湿度和叶水势有显著相关关系,当CO2浓度较小时,胡杨气孔导度随CO2浓度的增加而增加,当CO2浓度达到一定值后气孔导度不再增加,反而随CO2浓度的增加大幅度降低。(4)胡杨适应极端干旱区生境的气孔调节机制为反馈式反应,即由于叶水势降低导致气孔导度减小,从而减少蒸腾耗水,达到节约用水、适应干旱的目的,表明胡杨的水分利用效率随气孔限制值的增大而减小,二者呈显著负相关。     

Simulation of the Physiological Responses of C3 Plant Leaves to Environmental Factors by a Model Which Combines Stomatal Conductance,Photosynthesis and Transpiration

Transpiration element is included in the integrated stomatal conductance-photosynthesis model by considering gaseous transfer processes, so the present model is capable to simulate the influence of boundary layer conductance. Leuning in his revised Ball' s model replaced relative humidity with VPDs(the vapor pressure deficit from stomatal pore to leaf surface) and thereby made the relation with transpiration more straightforward, and made it possible for the regulation of transpiration and the influence of boundary layer conductance to be integrated into the combined model. If the differences in water vapor and CO2 concentration between leaf and ambient air are considered, VPDs, the evaporative demand, is influenced by stomatal and boundary layer conductance. The physiological responses of photosynthesis, transpiration, and stomatal function, and the changes of intercellular CO2 and water use efficiency to environmental factors, such as wind speed, photon flux density, leaf temperature and ambient CO2, are analyzed. It is shown that ff the boundary layer conductance drops to a level comparable with stomatal conductance, the results of simulation by the model presented here differ significantly from those by the previous model, and, in some cases, are more realistic than the latter.     

光合作用-蒸腾作用-气孔导度的耦合模型及C_3植物叶片对环境因子的生理响应(英文)

以叶片的气体传输过程为基础,将蒸腾作用包括在以往光合作用气孔导度的耦合模型中,建立了光合作用蒸腾作用气孔导度的耦合模型。该模型可以模拟边界层导度对生理过程的影响。模拟了Ｃ３植物叶片对环境因子,如光照、温度、湿度、边界层导度和ＣＯ２浓度等的生理响应（光合作用、蒸腾作用、气孔导度）以及Ｃｉ和水分利用效率的变化。在环境因子变化于较大范围的情况下,模拟结果符合许多实验结论。     

Leaf area and photosynthesis of newly emerged trifoliolate leaves are regulated by mature leaves in soybean

Leaf anatomy and the stomatal development of developing leaves of plants have been shown to be regulated by the same light environment as that of mature leaves, but no report has yet been written on whether such a long-distance signal from mature leaves regulates the total leaf area of newly emerged leaves. To explore this question, we created an investigation in which we collected data on the leaf area, leaf mass per area (LMA), leaf anatomy, cell size, cell number, gas exchange and soluble sugar content of leaves from three soybean varieties grown under full sunlight (NS), shaded mature leaves (MS) or whole plants grown in shade (WS). Our results show that MS or WS cause a marked decline both in leaf area and LMA in newly developing leaves. Leaf anatomy also showed characteristics of shade leaves with decreased leaf thickness, palisade tissue thickness, sponge tissue thickness, cell size and cell numbers. In addition, in the MS and WS treatments, newly developed leaves exhibited lower net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (E), but higher carbon dioxide (CO ₂ ) concentration in the intercellular space (Ci) than plants grown in full sunlight. Moreover, soluble sugar content was significantly decreased in newly developed leaves in MS and WS treatments. These results clearly indicate that (1) leaf area, leaf anatomical structure, and photosynthetic function of newly developing leaves are regulated by a systemic irradiance signal from mature leaves; (2) decreased cell size and cell number are the major cause of smaller and thinner leaves in shade; and (3) sugars could possibly act as candidate signal substances to regulate leaf area systemically.     

Structural and hydraulic correlates of heterophylly in Ginkgo biloba

This study investigates the functional significance of heterophylly in Ginkgo biloba, where leaves borne on short shoots are ontogenetically distinct from those on long shoots. Short shoots are compact, with minimal internodal elongation; their leaves are supplied with water through mature branches. Long shoots extend the canopy and have significant internodal elongation; their expanding leaves receive water from a shoot that is itself maturing. Morphology, stomatal traits, hydraulic architecture, Huber values, water transport efficiency, in situ gas exchange and laboratory-based steady-state hydraulic conductance were examined for each leaf type. Both structure and physiology differed markedly between the two leaf types. Short-shoot leaves were thinner and had higher vein density, lower stomatal pore index, smaller bundle sheath extensions and lower hydraulic conductance than long-shoot leaves. Long shoots had lower xylem area:leaf area ratios than short shoots during leaf expansion, but this ratio was reversed at shoot maturity. Long-shoot leaves had higher rates of photosynthesis, stomatal conductance and transpiration than short-shoot leaves. We propose that structural differences between the two G. biloba leaf types reflect greater hydraulic limitation of long-shoot leaves during expansion. In turn, differences in physiological performance of short- and long-shoot leaves correspond to their distinct ontogeny and architecture.     

水稻叶片光合作用对开放式空气CO2浓度增高(FACE)的响应与适应

利用便携式光合气体分析系统 (LI 6 4 0 0 ) ,比较测定了高CO2 浓度 (FACE ,free airCO2 enrich ment)和普通空气CO2 浓度下生长的水稻叶片的净光合速率、水分利用率、表观量子效率和RuBP羧化效率等光合参数 .在各自生长CO2 浓度 (380vs 5 80 μmol·mol-1)下测定时 ,高CO2 浓度 (5 80 μmol·mol-1)下生长的水稻叶片的净光合速率、碳同化的表观量子效率和水分利用率明显高于普通空气 (380 μmol·mol-1)下生长的水稻叶片 .但是 ,随着FACE处理时间的延长 ,高CO2 浓度对净光合速率的促进作用逐渐减小 .在相同CO2 浓度下测定时 ,FACE条件下生长的水稻叶片净光合速率和羧化效率明显比普通空气下生长的对照低 .尽管高CO2 浓度下生长的水稻叶片的气孔导度明显低于普通空气中生长的水稻叶片 ,但两者胞间CO2 浓度差异不显著 ,因此高CO2 浓度下生长的水稻叶片光合下调似乎不是由气孔导度降低造成的 .     

Juvenile Rhus glabra leaves have higher temperatures and lower gas exchange rates than mature leaves when compared in the field during periods of high irradiance

We sought to test the hypothesis that stomatal development determines the timing of gas exchange competency, which then influences leaf temperature through transpirationally driven leaf cooling. To test this idea, daily patterns of gas exchange and leaflet temperature were obtained from leaves of two distinctively different developmental stages of smooth sumac (Rhus glabra) grown in its native habitat. Juvenile and mature leaves were also sampled for ultrastructural studies of stomatal development. When plants were sampled in May-June, the hypothesis was supported: juvenile leaflets were (for part of the day) from 1.4 to 6.0 degrees C warmer than mature leaflets and as much as 2.0 degrees C above ambient air temperature with lower stomatal conductance and photosynthetic rates than mature leaflets. When measurements were taken from July to October, no significant differences were observed, although mature leaflet gas exchange rates declined to the levels of the juvenile leaves. The gas exchange data were supported by the observations that juvenile leaves had approximately half the number of functional stomata on a leaf surface area basis as did mature leaves. It was concluded that leaf temperature and stage of leaf development in sumac are strongly linked with the higher surface temperatures observed in juvenile leaflets in the early spring possibly being involved in promoting photosynthesis and leaf expansion when air temperatures are cooler.     

Photosynthetic activity and water relations of sprouts ofPasania edulis

In order to investigate the factors causing fast growth of sprouts ofPasania edulis, photosynthetic activity and water relation characteristics of lower (mature) leaves and upper (expanding) leaves of the sprouts were compared with those of seedlings and adult trees ofP. edulis. Apparent quantum yield was generally low. Maximum photosynthetic rate was highest in the lower leaves of sprouts. Stomatal frequency was higher in sprout leaves than in seedling leaves. Osmotic potential at the water saturation point and water potential at the turgor loss point, in leaves, were higher in sprouts than in seedlings and adult trees. Symplasmic water content per unit leaf area was higher in sprouts than in seedlings. These water relation parameters in leaves indicated that sprout leaves are superior in maintaining cell turgor against water loss, but are not tolerant to water stress. In field measurements, sprout leaves showed higher stomatal conductance and transpiration rates. These results indicated that sprout leaves fully realized their high potential productivity even under field conditions. The leaf specific conductance, from the soil to the leaf, was higher in sprouts than in seedlings. Large and deep root systems of the original stumps of the sprouts may be attributed to the high leaf specific conductance.     

Impact of elevated CO(2) and O(3) on gas exchange parameters and epidermal characteristics in potato (Solanum tuberosum L.)

Potato plants (Solanum tuberosum L. cv. Bintje) were grown in open-top chambers (OTCs) under three CO(2) levels (ambient and 24 h d(-1) seasonal mean concentrations of 550 and 680 micromol mol(-1)) and two O(3) levels (ambient and a seasonal mean 8 h d(-1) concentration of 50 nmol mol(-1)). The objectives were to determine the effects of season-long exposure to these key climate change gases on gas exchange, leaf thickness and epidermal characteristics. The experimental design also provided an ideal opportunity to examine within-leaf variation in epidermal characteristics at the whole-leaf level. Stomatal and epidermal cell density and stomatal index were measured at specific locations on the youngest fully expanded leaf (centre of lamina, mid-way between tip and base) and representative whole leaves from each treatment. Effects on leaf conductance, assimilation rate and instantaneous transpiration efficiency were determined by infrared gas analysis, while anatomical characteristics were examined using a combination of leaf impressions and thin sections. Exposure to elevated CO(2) or O(3) generally increased leaf thickness, leaf area, stomatal density, and assimilation rate, but reduced leaf conductance. The irregular stomatal distribution within leaves resulted from a combination of uneven differentiation and expansion of the epidermal cells. The results are discussed with reference to sampling protocols and the need to account for within-leaf variation when examining the impact of climate change or other environmental factors on epidermal characteristics.     

Stomatal control of transpiration from a developing sugarcane canopy

Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.     

提高CO2浓度对两种亚热带树苗叶片水分状况的影响

鼎湖山季风常绿阔叶林的主要优势乔木树种黧蒴和荷木的幼苗,盆栽于自然光照和人工调节CO₂浓度为500μl·L^-1或空气CO₂(340μl·L^-1)的气罩中3个月.在各自生长条件下测定,高CO₂下生长的黧蒴和荷木叶片平均气孔导度分别降低13%和20%,蒸腾速率下降20%和18%,水分利用效率提高1倍以上,不同CO₂浓度下的植物叶片气孔导度和蒸腾速率日进程曲线也有明显差异.处理后将幼苗置于自然条件下观测其后效应,第7d时处理间的气孔导度和蒸腾速率皆无明显差异     

Specification of adaxial and abaxial stomata, epidermal structure and photosynthesis to CO2 enrichment in maize leaves

Acclimation to CO2 enrichment was studied in maize plants grown to maturity in either 350 or 700 microl l-1 CO2. Plants grown with CO2 enrichment were significantly taller than those grown at 350 microl l-1 CO2 but they had the same number of leaves. High CO2 concentration led to a marked decrease in whole leaf chlorophyll and protein. The ratio of stomata on the adaxial and abaxial leaf surfaces was similar in all growth conditions, but the stomatal index was considerably increased in plants grown at 700 microl l-1 CO2. Doubling the atmospheric CO2 content altered epidermal cell size leading to fewer, much larger cells on both leaf surfaces. The photosynthesis and transpiration rates were always higher on the abaxial surface than the adaxial surface. CO2 uptake rates increased as atmospheric CO2 was increased up to the growth concentrations on both leaf surfaces. Above these values, CO2 uptake on the abaxial surface was either stable or increased as CO2 concentration increased. In marked contrast, CO2 uptake rates on the adaxial surface were progressively inhibited at concentrations above the growth CO2 value, whether light was supplied directly to this or the abaxial surface. These results show that maize leaves adjust their stomatal densities through changes in epidermal cell numbers rather than stomatal numbers. Moreover, the CO2-response curve of photosynthesis on the adaxial surface is specifically determined by growth CO2 abundance and tracks transpiration. Conversely, photosynthesis on the abaxial surface is largely independent of CO2 concentration and rather independent of stomatal function.     

高羊茅叶片表皮蜡质含量与其抗旱性的关系

以14个高羊茅品种为试验材料,在田间试验中对干旱高温胁迫下的叶片表皮蜡质含量、净光合速率、蒸腾速率、气孔导度、胞间CO2浓度等生理指标测定分析。结果表明,干热胁迫下高羊茅品种间的叶片表皮蜡质含量和水分利用效率均存在极显著差异(P<0.01);叶片蜡质含量与综合抗旱性和水分利用效率的等级相关系数分别为0.78(P<0.01)和0.68(P<0.01);蜡质含量越高的品种,其叶片气孔导度和胞间CO2浓度越低,水分利用效率越高,但所有品种的水分利用效率绝对值都较低。研究发现,在干热胁迫时,高羊茅叶片表皮蜡质可通过对气孔导度的调节来减少气孔蒸腾,提高水分利用效率,最终提高其抗旱性;表皮蜡质含量可以作为高羊茅品种抗旱性鉴定的一个新指标。     

Changes in leaf physiology caused by Calacarus heveae (Acari, Eriophyidae) on rubber tree

The influence of Calacarus heveae Feres on physiological processes was evaluated in two rubber tree clones. Experiments were conducted in a greenhouse with 5-month-old potted seedlings of RRIM 600 and GT 1 clones, that were either infested with C. heveae or not (non-infested control). The level of photosynthetic pigments, net photosynthetic rate, stomatal conductance, transpiration rate, changes in relative humidity between leaf surface and ambient air (Δw) and intercellular CO(2) concentration (Ci CO(2)) were evaluated. Infested plants showed significant reductions in the rate of transpiration, the rate of photosynthesis, stomatal conductance and Δw. RRIM 600 seedlings showed more pronounced physiological damage than GT 1 seedlings, indicating a lower physiological tolerance of the former clone to the mite. However, carotenoid levels were reduced only in GT 1 seedlings. Photosynthesis was probably reduced due to a decrease in stomatal opening, as indicated by reductions in transpiration rate and stomatal conductance and by the absence of differences in chlorophyll levels between treatments. Our results indicate that populations of C. heveae reduce the productivity of rubber trees. Thus, farmers must to be aware to control this mite pest in rubber tree plantations.     

Leaf age effects on photosynthetic rate, transpiration rate and nitrogen content in a tropical dry forest

This study examines the effect of leaf age on photosynthesis, transpiration and nitrogen concentration in four deciduous (DC) and two evergreen (EG) species coexisting in a tropical dry forest of Venezuela. Leaf age was characterized on the basis of leaf chorophyll, nitrogen content, and construction and maintenance costs. The mean leaf area-based nitrogen concentration (N) in EG was about twice that in DC species. A leaf age effect was observed in both DC and EG species, with largest N concentration in mature leaves. Fractional leaf N allocation to chlorophyll was higher in the DC than in the EG species. Differences in the construction costs of leaf mass between the youngest and the oldest leaves averaged from 2.14 to 1.55 g glucose g⁻¹ dry weight. Although variation in area-based leaf maintenance and construction costs between DC and EG species existed, they were, nevertheless, positively correlated. Individual data sets, for each species, indicated that leaf N and maximum rate of photosynthesis (A_max) were linearly related. Nitrogen use efficiency (NUE) and water use efficiency (WUE) tended to be higher in mature leaves than in expanding and old leaves. Moreover, DC species always had higher NUE than EG species. Intercellular to ambient pressures of CO₂ (P_i/P_a) were related to WUE in a negative manner. Higher P_i/P_a values were observed in expanding and old leaves. Leaf age effect on photosynthesis was, therefore, due to greater decline of carbon fixation capacity by mesophyll tissue relative to the decline in stomatal conductance in youngest and oldest leaves.