Efficiency and regulation of water transport in some woody and herbaceous species

The efficiency with which plants transport water is related to the water potential differences required to drive water fluxes from the soil to the leaf. A comparative study of two woody and three herbaceous species (Citrus sinensis L. cv. Koethen, Pyrus kawakami L., Helianthus annuus L. cv. Mammoth Russian, Capsicum frutescens L. cv. Yolo Wonder, and Sesamum indicum L. cv. Glauca) indicated contrasts in water transport efficiency. Depression of leaf water potential in response to transpiration increases was found in the woody species; the herbaceous species, however, had more efficient water transport systems and presented no measurable response of leaf water potential to transpiration changes. Different maximum transpiration rates under the same climatic conditions were observed with different species and may be accounted for by stomatal response to humidity gradients between leaf and air. Leaf diffusion resistance in sesame increased markedly as the humidity gradient was increased, while leaf resistance of sunflower responded less to humidity. Stomata appeared to respond directly to the humidity gradient because changes in leaf water potential were not detected when leaf resistance increased or decreased.     

Selective oviposition by a leaf miner in response to temporal variation in abscission

Summary Responses to humidity of net photosynthesis and leaf conductance of single attached leaves were examined in populations of herbs from wet soil sites in Beltsville, Maryland and Davis, California, USA. Plants were grown in controlled environments under three conditions which differed in the magnitude of the day-night temperature difference and in daytime air saturation deficit. No population differences in response were found in Abutilon theophrasti. In Amaranthus hybridus stomatal conductance and net photosynthesis were more reduced by increasing leaf to air water vapor pressure difference (VPD) in the population from Beltsville, but only for the growth condition with a constant 25°C temperature. In Chenopodium album, stomatal conductance was more sensitive to VPD in the population from Davis, but only for the growth condition with 28/22°C day/night temperatures. Population differences in the sensitivity to VPD of leaf conductance were associated with differences in leaf area to root weight ratio. The relative reduction of net photosynthesis as VPD increased was greater than, equal to, or less than the relative decrease in substomatal carbon dioxide partial pressure. The pattern depended on species, and on growth condition. From these results one can not conclude that environmental humidity has been a strong selective force in determining sensitivity to humidity of stomatal conductance.     

Stomatal response of engelmann spruce to humidity, light, and water stress

Stomatal response of Engelmann spruce (Picea engelmannii Engelm.) to environmental conditions was studied in the natural subalpine environment and under controlled laboratory conditions. Stomata of naturally occurring trees responded to the difference in absolute humidity from leaf to air. When foliage was exposed to full sunlight, stomatal conductance decreased as the absolute humidity difference increased. In the shade, where photosynthetically active radiation was 10% of that in full sunlight, stomatal closure at large absolute humidity differences was much more complete. No effect of soil or air temperatures on stomatal aperture was observed in the field, nor were differences among three contrasting sites detected. Under growth chamber conditions, stomata responded to photosynthetically active radiation, but conductances were influenced by leaf-to-air differences in absolute humidity. Leaf water potentials below - 15 bars resulted in lower conductances over a range of humidity and light conditions. Because net photosynthesis under shaded conditions in the natural environment must be very low, stomatal closure could result in considerable savings in water while having a minimum effect on net photosynthesis.     

Development of a photosynthesis model with an emphasis on ecological applications

Summary A physiologically based steady-state model of whole leaf photosynthesis (WHOLEPHOT) is used to describe net photosynthesis daily time courses in Prunus armeniaca. Net photosynthesis rates are calculated in response to incident light intensity, leaf temperature, air carbon dioxide concentration, and leaf diffusion resistance measured at five minute intervals. The steady-state calculations closely approximate the observed net photosynthesis rates for a broad range of weather conditions and leaf stomatal behavior.     

The diurnal time course of net photosynthesis of soybean leaves: Analysis with a physiologically based steady-state photosynthesis model

Summary A physiologically based steady-state model of whole leaf photosynthesis (WHOLEPHOT) is used to analyze observed net photosynthesis daily time courses of soybean, Glycine max (L.) Merr., leaves. Observations during two time periods of the 1978 growing season are analyzed and compared. After adjustment of the model for soybean, net photosynthesis rates are calculated with the model in response to measured incident light intensity, leaf temperature, air carbon dioxide concentration, and leaf diffusion resistance. The steady-state calculations closely approximate observed net photosynthesis. Results of the comparison reveal a decrease in photosynthetic capacity in leaves sampled during the second time period, which is associated with decreasing ability of leaves to respond to light intensity and internal air space carbon dioxide concentration, increasing mesophyll resistance, and increasing stomatal resistance.     

Responses of stomata of senescing and nonsenescing leaves of Nicotians glauca to changes in intercellular concentrations of leaf carbon dioxide

Abstract. Gas exchange measurements were performed to test the hypothesis that failure of stomata to open in senescing leaves of Nicotiana glauca is caused by elevated concentrations of carbon dioxide in the intercellular spaces of leaf mesophyll tissue (c_i). Senescing leaves selected for experiments were completely chlorotic and lacked positive rates of photosynthesis. When stomata in detached epidermis from senescing leaves were illuminated in CO2-free air, they opened to similar apertures as those in detached epidermis from nonsenescing leaves. To compare the effects of changes in c_i on stomatal responses of the two leaf types, leaf 'flags' of either nonsenescing or senescing leaves were illuminated at a photosynthetic photon flux density of 500 μmol m⁻² s⁻¹ in a gas exchange cuvette. Leaf temperatures were maintained at 23.5 ± 0.5°C, and vapour pressure differences between leaves and the air were maintained between 0.70 and 0.75kPa. C_i was adjusted by changing external concentrations of carbon dioxide in air circulating through the cuvette. Conductances and photosynthetic rates of nonsenescing leaves changed in response to changes in c_i, but neither the conductances nor the photosynthetic rates of senescing leaves were affected significantly by changes in q. We conclude that guard cells of senescing leaves of Nicotiana glauca do not lose the capacity to respond to changes in carbon dioxide concentration and that increases in c_i resulting from declining rates of mesophyll photosynthesis are not the sole cause of maintenance of stomatal closure during leaf senescence. The data suggest that factors external to guard cells may prevent them from responding to changes in carbon dioxide concentrations in intact senescing leaves.     

Does transpiration control stomatal responses to water vapour pressure deficit?

Three types of observations were used to test the hypothesis that the response of stomatal conductance to a change in vapour pressure deficit is controlled by whole-leaf transpiration rate or by feedback from leaf water potential. Varying the leaf water potential of a measured leaf by controlling the transpiration rate of other leaves on the plant did not affect the response of stomatal conductance to vapour pressure deficit in Glycine max. In three species, stomatal sensitivity to vapour pressure deficit was eliminated when measurements were made at near-zero carbon dioxide concentrations, despite the much higher transpiration rates of leaves at low carbon dioxide. In Abutilon theophrasti, increasing vapour pressure deficit sometimes resulted in both decreased stomatal conductance and a lower transpiration rate even though the response of assimilation rate to the calculated substomatal carbon dioxide concentration indicated that there was no ‘patchy’ stomatal closure at high vapour pressure deficit in this case. These results are not consistent with stomatal closure at high vapour pressure deficit caused by increased whole-leaf transpiration rate or by lower leaf water potential. The lack of response of conductance to vapour pressure deficit in carbon dioxide-free air suggests that abscisic acid may mediate the response.     

Some effects of air temperature and humidity on crop and leaf photosynthesis, transpiration and resistance to gas transfer

Measurements of carbon dioxide exchange and transpiration were made, at various air temperatures, on wheat and barley using a field enclosure system. From these were derived the stomatal and mesophyll resistances to carbon dioxide transfer. Optimum temperatures for net CO₂ uptake were about 24°C for wheat and barley. Above these optima, as temperature increased so net CO₂ uptake rates decreased, because of increasing stomatal and mesophyll resistances; transpiration rates decreased in wheat but were constant in barley. In laboratory growth cabinets, wheat plants were subjected to different regimes of temperature and humidity. Optimum temperature for net CO₂ uptake of individual leaves was 25°C. At constant humidity, a decline in net uptake rates above 25°C was associated with large increases in mesophyll resistance. At a constant 25°C, as the vapour pressure deficit (v.p.d. was increased above 1 k Pa (10 mb) v.p.d. the net uptake declined, with an increase in mesophyll resistance and a small increase in stomatal resistance. When the v.p.d. exceeded 1 k Pa at a temperature of 30°C, conditions that are experienced by field plants, then there were large increases in both mesophyll and stomatal resistances and the net uptake rates declined. Photo-respiration, as measured by CO₂ uptake in oxygen-free air, was independent of temperature, but both dark respiration and CO₂ compensation concentration increased with temperature.     

Leaf hydraulic conductivity and stomatal responses to humidity in amphistomatous leaves

The response of stomata to changes in humidity for a single surface of an amphistomatous leaf was investigated in Xanthium strumarium and Vicia faba using gas exchange and direct observation of stomatal apertures. The stomatal response to humidity for a given surface was found to be the same whether or not the humidity for the opposite surface was changed concurrently. Stomata on the surface for which humidity was constant showed no response to changes in humidity for the opposite surface. Despite large changes in epidermal turgor on the surface for which humidity was changed, there was no change in epidermal turgor for the surface with constant humidity. Measurements of transpiration and epidermal turgor as functions of the mole fraction gradient of water between leaf and air were used to calculate a value for leaf hydraulic resistance. The results suggest that in these species, the mechanism for the stomatal response to humidity resides in the epidermis or the mesophyll very close to the epidermis, and that most of the hydraulic resistance of the leaf occurs between the xylem and the evaporating sites.     

Low humidity effects on photosynthesis in single leaves of C4 plants

Summary It was hypothesized that since sub-stomatal carbon dioxide concentrations are often saturating to photosynthesis at ambient external concentrations in C₄ plants at high light, photosynthesis might be insensitive to partial stomatal closure caused by large leaf-air water vapor pressure difference. The response of stomatal conductance and photosynthesis at high irradiance to vapor pressure difference was determined under uniform conditions in C₄ plants grown under controlled conditions, and outdoors. In several cases, photosynthesis was less sensitive to stomatal closure than it would have been if photosynthesis had a linear response to sub-stomatal carbon dioxide concentration. No change in photosynthesis at up to 25 mbar vapor pressure difference was demonstrated in the C₄ species Portulaca oleracea and Amaranthus hypochondriacus, despite reductions in stomatal conductance of 32 and 17%, respectively. Sensitivity of photosynthesis to leaf-air vapor pressure difference was found to depend on the species and on the growth conditions.     

田间大豆光合午休原因的综合研究

本研究利用主成分分析构造同归模型,利用典型相关分析等方法,对自变量的重要程度进行排序,确定了光合午休的主要原因是气孔因素限制所致.光抑光合也是一个值得注意的因素,但不是主要因素.空气相对湿度、叶片与大气间水汽压差是诱发气孔限制的重要因素.当空气相对湿度较为适中时,叶温的影响较小,在空气相对湿度变坏的情况下,叶温才对光合午休有重要影响.     

Sensitivity of infrared water vapor analyzers to oxygen concentration and errors in stomatal conductance

Use of infrared analyzers to measure water vapor concentrations in photosynthesis systems is becoming common. It is known that sensitivity of infrared carbon dioxide and water vapor analyzers is affected by the oxygen concentration in the background gas, particularly for absolute analyzers, but the potential for large errors in estimates of stomatal conductance due to effects of oxygen concentration on the sensitivity of infrared water vapor analyzers is not widely recognized. This work tested three types of infrared water vapor analyzers for changes in sensitivity of infrared water vapor analyzers depending on the oxygen content of the background gas. It was found that changing from either 0 or 2% to 21% oxygen in nitrogen decreased the sensitivity to water vapor for all three types of infrared water vapor analyzers by about 4%. The change in sensitivity was linear with oxygen mole fraction. The resulting error in calculated stomatal conductance would depend strongly on the leaf to air vapor pressure difference and leaf temperature, and also on whether leaf temperature was directly measured or calculated from energy balance. Examples of measurements of gas exchange on soybean leaves under glasshouse conditions indicated that changing from 21% to 2% oxygen produced an artifactual apparent increase in stomatal conductance which averaged about 30%. Similar errors occurred for `conductances' of wet filter paper. Such errors could affect inferences about the carbon dioxide dependence of the sensitivity of photosynthesis to oxygen. This revised version was published online in June 2006 with corrections to the Cover Date.     

The effect of air pollutants on physiological processes in plants

Abstract. Important physiological processes, photosynthesis, respiration, carbon allocation and stomatal function are known to be affected by air pollutants. A wide range in sensitivity of photosynthesis both within and between species is evident from the literature for the pollutants sulphur dioxide, ozone, nitrogen oxide and hydrogen fluoride. Some of this variation is clearly due to genetic factors, but much is in response to differences in environmental conditions both prior to and during fumigation. Exposure of plants to mixtures of pollutants generally reduced the threshold at which effects were first detected and increased the level of inhibitory responses. In the majority of studies on stomatal responses to air pollutants, opening occurs at low concentrations, below the threshold for effects on photosynthesis, and closure occurs at injurious concentrations; this latter response often following the inhibition of photosynthesis. Effects on carbon allocation have been reported in response to air pollutants. Changes usually favour leaf development over root growth, which can compensate for a decline in net assimilation rate up to a certain point but may limit water uptake from soils with low moisture content. Future research into physiological effects of air pollutants should incorporate an integrated approach in which both key physiological parameters and growth parameters are measured together with estimates of the effective dose of pollutant. In this way, the underlying mechanisms to changes in growth and development will be more fully understood.     

空气湿度与土壤水分胁迫对紫花苜蓿叶表皮蜡质特性的影响

选用2个抗旱性不同的紫花苜蓿品种,敖汉(强抗旱)和三得利(弱抗旱),设置空气湿度(45%-55%和75%-85%)和土壤水分胁迫(75%和35%田间持水量)处理,分析紫花苜蓿叶表皮蜡质含量、组分及晶体结构、气体交换参数、水势及脯氨酸含量的变化规律。结果表明,单独土壤水分胁迫时,紫花苜蓿叶表皮蜡质晶体结构及蜡质总量无显著变化;敖汉蜡质组分中烷类、酯类含量增加,醇类含量下降;三得利醇类含量下降,烷类、酯类含量变化不显著。低空气湿度胁迫时,两品种蜡质总量无显著变化,烷类和酯类含量显著增加,醇类含量显著下降,叶表皮片状蜡质晶体结构熔融呈弥漫性,扩大了对叶表面积的覆盖,其蒸腾速率显著低于正常湿度。复合胁迫处理时,叶表皮片状蜡质晶体结构继续呈弥漫性,烷类、酯类、未知蜡质组分含量均高于单独胁迫处理,醇类含量最低,而蜡质总量除三得利显著高于对照外,其余均无显著差异。紫花苜蓿叶表皮蜡质各组分含量(除醇类)及蜡质总量与光合速率呈显著负相关,与蒸腾速率无显著相关关系。蜡质总量与叶水势呈显著正相关。总体上,敖汉蜡质总量显著高于三得利,蜡质组分中烷类物质的增加有助于提高植株的抗旱性。在复合胁迫下,强抗旱品种主要通过气孔因素控制水分散失,而弱抗旱品种通过气孔和非气孔因素共同控制植物水分散失。     

Studies in Stomatal Behaviour: The Effects of Moving, Normal and Carbon Dioxide-Free Air on the Stomata of Some Plants

A series of experiments were conducted to investigate the effects of moving, normal and carbon dioxide-free air on the stomata within porometer cups attached to the illuminated leaves of Datura stramonium, Pelargonium zonale, Duranta plumieri and Clerodendron inerme. A marked opening stomatal response was obtained in Datura and Pelargonium on changing to carbon dioxide-free air. In general no such response was established in Duranta and Clerodendron. Indications of even an opposite closing effect were shown in the case of Duranta. The present results seem to disprove the reduction of carbon dioxide by photosynthesis as the major factor controlling the wide stomatal opening within a porometer cup. The possibility of a humidity effect is also suggested.     

Stomatal responses to changes in air humidity are not necessarily linked to nocturnal CO2 uptake in the CAM plant Plectranthus marrubioides Benth. (Lamiaceae)

Plants of the crassulacean acid metabolism (CAM) species Plectranthus marrubioides (Lamiaceae) were subjected to short- and long-term changes in air humidity in controlled-environment experiments. Stomata of well-watered individuals of this all-cell leaf-succulent taxon responded directly, quickly and reversibly to variations of the water vapour gradient between leaf and air (Δw). Mean night-time leaf conductance to water vapour decreased curvilinearly with increasing Δw but linearly with lowered relative air humidity. Stomatal response was generally independent of the prevailing temperature and was not linked to CO₂ uptake rates. Therefore, net night-time carbon gain, nocturnal malic acid accumulation and, thus, relative carbon recycling were not influenced by changes in air humidity in the temperature range tested. Mean nocturnal molar water use efficiency, however, decreased with decreasing air humidity because of the increased transpirational water loss. If watering was repeatedly withheld for several days during the experiments, employing a temperature regime of 35/30°C day and night, stomatal conductance became low enough to inhibit CO₂ uptake, but only at the highest Δw. The results suggest that drought stress was necessary to increase responsiveness of plants to the point where CAM was also inhibited by decreases in air humidity.     

The Measurement of Stomatal Responses to Stimuli in Leaves and Leaf Discs

A comparison has been made of stomatal responses in intact leaves,leaf discs supplied with water via their cut edges and leafdiscs floating on water. Xanthium pennsylvanicum leaf discswatered via their cut edges appeared to be more turgid thanintact leaves; this considerably slowed down the rate of stomatalopening but it slightly increased the final steady-state stomatalopening. When the water potential of such leaf discs was loweredby pre-treatment with mannitol solutions rates of stomatal openingincreased whereas maximum steady-state openings decreased. In tobacco leaf discs floating on water the stomata in contactwith water were wider open than those in contact with normalair and they did not respond to treatment with carbon dioxide-freeair. The rate of photosynthesis was severely reduced in tobaccoleaf discs floating with the lower epidermis on water, mostprobably owing to the slow rate of diffusion of carbon dioxidein water. By floating such discs on osmotica the degree of stomatalopening was increased, however, a response to treatment withcarbon dioxide-free air was still not measurable. It is postulatedthat, on account of the relative unavailability of carbon dioxidefrom the water, the carbon dioxide concentration in the substomatalcavities of the lower surface is abnormally low, irrespectiveof whether ordinary air or carbon dioxide-free air is availableto the upper surface. A comparison between porometer readings and measurements ofsiliconerubber impressions of stomatal pores taken from insidethe porometer cup confirmed that the silicone-rubber impressionmethod of assessing stomatal responses to stimuli has severelimitations, especially at small stomatal apertures.     

In Situ Measurement of Epidermal Cell Turgor, Leaf Water Potential, and Gas Exchange in Tradescantia virginiana L

A combined system has been developed in which epidermal cell turgor, leaf water potential, and gas exchange were determined for transpiring leaves of Tradescantia virginiana L. Uniform and stable values of turgor were observed in epidermal cells (stomatal complex cells were not studied) under stable environmental conditions for both upper and lower epidermises. The changes in epidermal cell turgor that were associated with changes in leaf transpiration were larger than the changes in leaf water potential, indicating the presence of transpirationally induced within-leaf water potential gradients. Estimates of 3 to 5 millimoles per square meter per second per megapascal were obtained for the value of within-leaf hydraulic conductivity. Step changes in atmospheric humidity caused rapid changes in epidermal cell turgor with little or no initial change in stomatal conductance, indicating little direct relation between stomatal humidity response and epidermal water status. The significance of within-leaf water potential gradients to measurements of plant water potential and to current hypotheses regarding stomatal response to humidity is discussed.     

Interaction of ozone pollution and light effects on photosynthesis in a forest canopy experiment

Ozone pollution may reduce net carbon gain in forests, yet data from mature trees are rare and the effects of irradiance on the response of photosynthesis to ozone remain untested. We used an open-air system to expose 10 branches within the upper canopy of an 18-m-tall stand of sugar maple (Acer saccharum Marsh.) to twice-ambient concentrations of ozone (95nmol mol^?1, 0900 to 1700, 1 h mean) relative to 10 paired, untreated controls (45nmol mol^?1) over 3 months. The branch pairs were selected along a gradient from relatively high irradiance (PPFD 14.5 mol m^?2 d^?1) to deep shade (0.7mol m^?2 d^?1). Ozone reduced light-saturated rates of net photosynthesis (A_sat) and increased dark respiration by as much as 56 and 40%, respectively. Compared to sun leaves, shade leaves exhibited greater proportional reductions in A_sat and had lower chlorophyll concentrations, quantum efficiencies, and leaf absorptances when treated with ozone relative to controls. With increasing ozone dose over time, A_sat became uncoupled from stomatal conductance as ratios of internal to external concentrations of carbon dioxide increased, reducing water-use efficiency. Ozone reduced net photosynthesis and impaired stomatal function, with these effects depending on the irradiance environment of the canopy leaves. Increased ozone sensitivity of shade leaves compared to sun leaves has consequences for net carbon gain in canopies.     

冬小麦叶片气孔导度模型水分响应函数的参数化

植物气孔导度模型的水分响应函数用来模拟水分胁迫对气孔导度的影响过程, 是模拟缺水环境下植物与大气间水、碳交换过程的关键算法。水分响应函数包括空气湿度响应函数和土壤湿度(或植物水势)响应函数, 该研究基于田间实验观测, 分析了冬小麦(Triticum aestivum)叶片气孔导度对不同空气饱和差和不同土壤体积含水量或叶水势的响应规律。一个土壤水分梯度的田间处理在中国科学院禹城综合试验站实施, 不同水分胁迫下的冬小麦叶片气体交换过程和气孔导度以及其他的温湿度数据被观测, 同时观测了土壤含水量和叶水势。实验数据表明, 冬小麦叶片气孔导度对空气饱和差的响应呈现双曲线规律, 变化趋势显示大约1 kPa空气饱和差是一个有用的阈值, 在小于1 kPa时, 冬小麦气孔导度对空气饱和差变化反应敏感, 而大于1 kPa后则反应缓慢; 分析土壤体积含水量与中午叶片气孔导度的关系发现, 中午叶片气孔导度随土壤含水量增加大致呈现线性增加趋势, 但在平均土壤体积含水量大于大约25%以后, 气孔导度不再明显增加, 而是维持在较高导度值上下波动; 冬小麦中午叶片水势与相应的气孔导度之间, 随着叶水势的增加, 气孔导度呈现增加趋势。根据冬小麦气孔导度对空气湿度、土壤湿度和叶水势的响应规律, 研究分别采用双曲线和幂指数形式拟合了水汽响应函数, 用三段线性方程拟合了土壤湿度响应函数和植物水势响应函数, 得到的参数可以为模型模拟冬小麦的各类水、热、碳交换过程采用。