Cyclic Variations in Plant Properties under Constant Environmental Conditions

Cyclic fluctuations in stomatal aperture, transpiration rate and leaf water potential under constant environmental conditions have been investigated in intact plants of cotton, pepper, and sunflower. Stomatal aperture and transpiration rate were least when leaf water potential was high and were greatest when leaf water potential was low. Lowest leaf water potential values lagged behind the occurrence of highest transpiration rates, and high overall resistance to water flow occurred in cycling plants. Both of these are considered essential for the occurrence of persistent cyclic behaviour. Hydropassive opening of stomates as the leaves wilted facilitated cycling in cotton and pepper, but not in sunflower, where hydropassive opening did not occur. The roots were identified as the site of the major resistance to water flow in the plant and further experiments directly showed the importance of this root resistance in initiating cycling by causing water stress in the leaves as the stomates opened. Root resistance varied diurnally, becoming increasingly important at night. Root resistance naturally rose to high levels in cotton. High levels were induced in pepper or sunflower by having the roots in deionized water for several days or by anoxia. Quantitative measurements of overall plant resistance were made from leaf water potential and transpiration rate data. The results are discussed and it suggested that plant resistance may indirectly be of importance in the movement of water from the plant to the air.     

Nonstomatal inhibition of photosynthesis at low water potentials in intact leaves of species from a variety of habitats

Mesophyll resistance to CO₂ uptake was calculated from gas exchange data on intact leaves of 12 species of woody plants. Plants studied were native to habitats ranging from streamsides to deserts. Gas exchange measurements were made at light saturation and constant temperature to eliminate possible effects of light and temperature on estimates of mesophyll resistance. Cuticular transpiration was measured and used in calculation of stomatal resistances from whole leaf transpiration rates. In all species examined, an increase in mesophyll resistance was observed as leaves dried. The increase in mesophyll resistance in all cases occurred at the same water potential as the initial decline in net photosynthesis, and was accompanied by an increase in stomatal resistance.     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

Whole-plant gas exchange responses of Spartina alterniflora (Poaceae) to a range of constant and transient salinities

A two-chamber-system was used to study whole-plant gas exchange responses of Spartina alterniflora to long-term and transient salinity treatments over the range of 5 to 40 ppt NaCl. Lower photosynthetic rates, leaf water vapor conductances, belowground respiration rates, and higher aboveground respiration rates in plants adapted to 40 ppt NaCl were observed. Area-specific leaf weight increased with salinity, although the salt content of leaf tissues did not. A reduced rate of gross photosynthesis and higher aboveground respiration rate in 40-ppt NaCl plants significantly lowered the net whole-plant CO₂ gain below that of 5-ppt NaCl plants, while the net CO₂ gain of 25-ppt NaCl plants was intermediate. Within 6 hr of increasing the salinity of 5- and 25-ppt NaCl plants by 20 and 15 ppt NaCl, S. alterniflora responded by reducing leaf water vapor conductance, which in turn reduced the photosynthetic rate. This response was reversed by returning the plants to their original salinity, which indicates that S. alterniflora adjusts water loss and gas exchange in response to transient salinity stress by regulating stomatal aperture. On the other hand, decreasing salinity of the growth media of plants cultured at 25 and 40 ppt NaCl had little or no effect on gas exchange characteristics. This suggests that S. alterniflora adapts to constant salinity through fixed, salinity-dependent structural modifications, such as stomatal density.     

Changes in the Water Balance and Photosynthesis of Onion, Bean and Cotton Plants under Saline Conditions

The osmotic concentration (osmotic potential) of onion leaf sap did not adjust to chloride salinity, and consequently water potential, turgor, stomatal aperture and transpiration were reduced. Although osmotic concentration of bean and cotton leaf sap did adjust to a saline root medium and turgor was no less in the salinized plants than in the controls, stomata of the salinized plants remained only partly open and transpiration was reduced. Net photosynthesis of onion plants was reduced by salinity (this effect being much enhanced in a hot dry atmosphere) but it could be rapidly raised to the level of the controls by inducing elevated leaf turgor. Stomatal closure was initially responsible for most of the ～30 % reduction in photosynthesis of salinized beans. This was due to interference with CO₂ diffusion and could be overcome by raising the CO₂ concentration in the air. At a later stage of growth, salinity affected the light reaction of bean photosynthesis, and elevation of the air CO₂ had little effect. Closure of stomata of salinized cotton plants had only a relatively small effect on net photosynthesis. Light intensity and CO₂ concentration experiments showed that salinity was reducing the photosynthesis of cotton leaves mainly by affecting the light reaction of photosynthesis. It is concluded that chloride salinity does affect the water balance and rate of photosynthesis of plants and that the nature and degree of the effects will depend upon climatic conditions and may be very different between plant species and in the same species at different periods of growth.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

两种生态型榕树的叶绿素含量、荧光特性和叶片气体交换日变化的比较研究

比较盆栽 生榕树和两栖型树的形态差异、叶片叶绿素含量、叶绿素荧光特性和气体交换的日变化。两栖型榕树具有较发达的气生根和水生不定根,叶片比陆生榕树宽,并有向中生性 倾向,陆生榕树的叶绿素含量比两栖榕树高,净光合速率略高于水培两栖型榕树,但明显高于土培两栖型榕树,蒸腾速率以水培两栖型树最高,陆生榕树次之,土培两栖型榕树最低,线性回归分析表明,三者的叶片气孔导度与净光合速率变化均呈正相关,气孔导度的变化     

SEASONAL PATTERN OF NET PHOTOSYNTHESIS OF ARTEMISIA TRIDENTATA

Gas exchange studies were carried out on Artemisia tridentata during the course of a growing season using microclimatically controlled cuvettes and infrared gas analysis. A definite seasonal pattern of net photosynthesis emerged. This pattern was influenced by the interaction of four major factors: plant water potential, leaf temperature, irradiation, and stage of phenological development. In spring and early summer, when plant water stress was minimal, photosynthesis rate was mainly correlated with leaf temperature and irradiation. During mid and late summer, increased plant water stress and phenological changes assumed at least equal importance with temperature and irradiation in limiting net photosynthesis. Indeed, plant water potential, mainly through its influence on stomatal aperture, r_s‘, was probably the single most important factor influencing assimilation rate of this species on a seasonal basis. However, variations in mesophyll resistance to CO₂ flux, r_m‘, in response to temperature, water stress, or phenological changes also were involved. Sagebrush photosynthesis under field conditions was highest in late May and early June, and declined thereafter, minimum rates occurring in August during the driest period. Optimal temperatures for net photosynthesis were higher later in the season, indicating a change in gas exchange capacity more suitable to the warmer temperatures later in the season.     

Inhibition of Corn and Sunflower Photosynthesis by Lead

Detached corn and sunflower leaves supplied with PbCl² via the transpiration stream exhibited reduced rates of photosynthesis. The difference between species in the amount of Pb taken up was in direct proportion to their respective transpiration rates. For both species the reduction in photosynthesis and the amount of Pb taken up increased with increasing treatment concentrations. A corresponding reduction occurred in the rate of transpiration suggesting that stomatal resistance may be increased by Pb contamination. The pathways of CO₂ and water vapor exchange are discussed in relation to the effects of Pb on photosynthesis and transpiration.     

SEASONAL PATTERNS OF CO2 AND WATER VAPOR EXCHANGE OF JUNCUS ROEMERIANUS SCHEELE IN A GEORGIA SALT MARSH

CO₂ and water vapor exchange studies of intact plants of black needle rush (Juncus roemerianus Scheele) were conducted in an undisturbed marsh community on Sapelo Island, Georgia. The seasonal patterns of the light and temperature responses of net photosynthesis, transpiration, leaf diffusive conductance, water-use efficiency and respiration were determined five times over the year. Internal resistances to CO₂ uptake were also evaluated. Net photosynthesis was highest in early spring, but declined only slightly through the year. A distinct and moderate temperature optimum of net photosynthesis was observed with decreasing rates above 30 C. Leaf conductances to water vapor were similar at all seasons and were high at cooler temperatures and decreased with increasing temperature. Transpiration was relatively high and constant during all seasons. The water-use efficiency of photosynthesis was high below 25 C, but decreased sharply above that temperature. Dark respiration was relatively low. Seasonal changes reflected changes in leaf density. Decreasing stomatal conductances and increasing respiration rates reduced net photosynthesis at higher temperatures. The stomatal resistance increased and internal resistances to CO₂ uptake decreased over the year, but the total resistance remained constant. The internal resistance to CO₂ uptake was consistently higher than the stomatal resistance. These seasonal response patterns show that J. roemerianus is well adapted to the seasonal changes in ambient temperature and irradiance and other microenvironmental factors in the high marsh. These physiological characteristics permit this C₃ species to maintain a high productivity in a seasonally hot and stressful environment.     

Flavescence dorée phytoplasma deregulates stomatal control of photosynthesis in Vitis vinifera

Flavescence dorée (FD) is among the major grapevine diseases causing high management costs; curative methods against FD are unavailable. In FD‐infected plants, decrease in photosynthesis is usually recorded, but deregulation in stomatal control of leaf gas exchange during FD infection and recovery is unknown. We measured the seasonal time course of gas exchange rates in two cultivars (‘Barbera’ and ‘Nebbiolo’) during the term of 1 year when grapevines experienced a water stress and another with no drought, with difference in gas exchange rates in response to FD infection and recovery as assessed by symptom observation and phytoplasma detection through PCR analysis. Chlorophyll fluorescence was also evaluated at the time of maximum symptom severity in ‘Barbera’, the cultivar showing the most severe stress response to FD infection, causing the highest damage in vineyards of north‐western Italy. In FD‐infected plants, net photosynthesis and transpiration gradually decreased during the season, more during the no drought year than during drought. During recovery, healthy (PCR negative) plants infected 2 years before, but not those infected an year before, regained the gas exchange performances to the level as measured before infection. The relationships between stomatal conductance and the residual leaf intercellular CO₂ concentration (c_i) discriminated healthy versus FD‐infected and recovered plants; at the same c_i, FD‐infected leaves had higher non‐photochemical quenching than healthy ones. We conclude that metabolic, not stomatal, leaf gas exchange limitation in FD‐infected and recovered grapevines is the basis of plant response to FD disease. In addition, we also suggest that such response is dependent upon water stress, by showing that water stress superimposes on FD infection in terms of stomatal and metabolic non‐stomatal limitations to carbon assimilation.     

Stomatal control of gas exchange in barley awns

The gas exchange of barley ears and awns was measured in the field using a gas analysis system and a diffusion porometer. Awn stomatal resistance decreased with increasing irradiance but to a smaller extent than leaf stomatal resistance. Measurements on ears immediately before and after successively removing awns showed that awn transpiration and photosynthesis were proportional to awn area and that awns accounted for 73% of transpiration by the ear. Although the maximum rates of photosynthesis of which awns were capable declined with age, awns accounted for 80–115% of the net CO₂ uptake of complete ears because the ears-less-awns could respire more CO₂ than they absorbed. Ear photosynthesis accounted for 52% of the weekly increment in ear dry weight after ear emergence, but 5 weeks later photosynthesis by the ear balanced respiration. Overall photosynthesis by the ear accounted for 35 % of its final weight. Differences in the light response curves of leaves and ears can be fully accounted for by the different relationships between stomatal resistance and irradiance of the two organs.     

Two Steps of Gas Exchange in Leaf Photosynthesis

Photosynthesis and transpiration of excised leaves of Taraxacum officinale L. and a few other species of plants were measured, using an open gas analysis system. The rates of CO₂ uptake and transpiration increased in two steps upon illumination of stomata-bearing epidermis of these leaves at a light intensity of 50 mW × cm⁻². Abscisic acid inhibited only the second step of gas exchange. Illumination of the astomatous epidermis of hypostomatous leaves caused only the first step of gas exchange. These data indicate that the first and second steps arise from cuticular and stomatal gas exchange, respectively. The rate of the cuticular photosynthesis in a Taraxacum leaf reached saturation at a light intensity of 5 mW × cm⁻², and the rates of the stomatal photosynthesis and transpiration reached saturation at a higher intensity of 35 mW × cm⁻². The cuticular photosynthesis of a Taraxacum leaf was 18% of the stomatal photosynthesis at 50 mW × cm⁻² and 270% at 5 mW × cm⁻². The other species of leaves showed the same trend. The importance of cuticular CO₂ uptake in leaf photosynthesis, especially under low light intensity was stressed from these data.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

The responses of leaf conductance, leaf water potential and rates of transpiration and net photosynthesis at different vapour pressure deficits ranging from 10 to 30 Pa kPa^-1 were followed in the sclerophyllous woody shrub Nerium oleander L. as the extractable soil water content decreased. When the vapour pressure deficit around a plant was kept constant at 25 Pa kPa^-1 as the soil water content decreased, the leaf conductance and transpiration rate showed a marked closing response to leaf water potential at-1.1 to-1.2 MPa, whereas when the vapour pressure deficit around the plant was kept constant at 10 Pa kPa^-1, leaf conductance decreased almost linearly from-0.4 to-1.1 MPa. Increasing the vapour pressure deficit from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, decreased leaf conductance at all exchangeable soil water contents. Changing the leaf water potential in a single leaf by exposing the remainder of the plant to a high rate of transpiration decreased the water potential of that leaf, but did not influence leaf conductance when the soil water content was high. As the soil water content was decreased, leaf conductances and photosynthetic rates were higher at equal levels of water potential when the decrease in potential was caused by short-term increases in transpiration than when the potential was decreased by soil drying.As the soil dried and the stomata closed, the rate of photosynthesis decreased with a decrease in the internal carbon dioxide partial pressure, but neither the net photosynthetic rate nor the internal CO₂ partial pressure were affected by low water potentials resulting from short-term increases in the rate of transpiration. Leaf conductance, transpiration rate and net photosynthetic rate showed no unique relationship to leaf water potential, but in all experiments the leaf gas exchange decreased when about one half of the extractable soil water had been utilized. We conclude that soil water status rather than leaf water status controls leaf gas exchange in N. oleander.     

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

Responses of Net Photosynthesis and Conductance to Independent Changes in the Humidity Environments of the Upper and Lower Surfaces of Leaves of Sunflower and Soybean

A dual-surface leaf chamber was used to investigate the responsesof net photosynthesis and leaf conductance to independent changesin the humidity environments of the upper and lower surfacesof leaves of sunflower and soybean. In sunflower decreasingthe humidity around the upper leaf surface while maintainingthat of the lower surface constant and high reduced both thephotosynthetic rate and the conductance of the lower surface.These reductions could not be attributed to changes in bulkleaf water potential since the transpiration rate of the wholeleaf remained constant. Similarly, the reductions were not relatedto localized water deficits in the lower epidermis or lowermesophyll since the transpiration rate of the lower surfacewas reduced. Possible mechanisms whereby the gas exchange characteristicsof the lower leaf surface of sunflower respond to the humidityenvironment of the upper surface are discussed. In contrastto sunflower, the photosynthetic rate of the lower surface ofsoybean was insensitive to the humidity environment of the uppersurface. In leaves of sunflower grown under a moderate temperature anda medium light level, simultaneous decreases of humidity atboth leaf surfaces reduced the photosynthetic rate of the wholeleaf without affecting the substomatal partial pressure of CO₂.In contrast, with leaves developed under a cool temperatureand a high light level, both the photosynthetic rate and thesubstomatal partial pressure of CO₂ were reduced. Evidently,the occurrence in sunflower of the response pattern suggestinga non-stomatal inhibition of photosynthesis by low humiditydepends upon the environment during growth. The possibilitythat this non-stomatal inhibition may be an artifact due toan error in the assumption of water vapour saturation withinthe leaf airspace is considered. Key words: Vapour pressure deficit, photosynthesis, conductance, non-stomatal inhibition, Helianthus annuus, Glycine max     

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.     

The effect of light on stomatal control of gas exchange in Douglas fir (Pseudotsuga menziesii) saplings

Summary Attached twigs of young Pseudotsuga menziesii (Mirb.) Franco plants were subjected to variations in irradaince. Stomatal responsiveness to irradiance, measured in an open type gas exchange system, varied seasonally. During the autumn and winter, stomatal conductance was relatively unresponsive to changes in irradiance, but during the summer stomatal conductance decreased in response to reduced irradiance. The summer stomatal response to irradiance was such that a nearly constant ratio of stomatal conductance to net photosynthesis was maintained as irradiance was varied. This caused intercellular CO₂ concentration (c _i) and water use efficiency (net CO₂ uptake/transpiration) to also remain relatively constant. At constant irradiance, stomatal conductance was relatively insensitive to experimentally-induced changes in c _i. This, and the observation that c _i remained relatively constant as irradiance was varied, suggest that changes in c _i played a minor role in mediating the stomatal response to light.The ecological significance of the seasonal changes in stomatal response to light is discussed.     

LIGHT,TEMPERATURE AND SALINITY EFFECTS ON GROWTH,LEAF ANATOMY AND PHOTOSYNTHESIS OF DISTICHLIS SPICATA (L.) GREENE

The effects of light, temperature, and salinity on growth, net CO₂ exchange and leaf anatomy of Distichlis spicata were investigated in controlled environment chambers. When plants were grown at low light, growth rates were significantly reduced by high substrate salinity or low temperature. However, when plants were grown at high light, growth rates were not significantly affected by temperature or salinity. The capacity for high light to overcome depressed growth at high salinity cannot be explained completely by rates of net photosynthesis, since high salinity caused decreases in net photosynthesis at all environmental conditions. This salinity-induced decrease in net photosynthesis was caused largely by stomatal closure, although plants grown at low temperature and low light showed significant increases in internal leaf resistance to CO₂ exchange. Increased salinity resulted in generally thicker leaves with lower stomatal density but no significant differences in the ratio of mesophyll cell surface area to leaf area. Salinity and light during growth did not significantly affect rates of dark respiration. The mechanisms by which Distichlis spicata tolerates salt appear to be closely coulpled to the utilization of light energy. Salt-induced leaf succulence is of questionable importance to gas exchange at high salinity in this C₄ species.