Stomatal conductance and transpiration of the two faces of Acacia phyllodes

Summary The daily course of stomatal conductance and transpiration was monitored on each separate face of vertical phyllodes of various acacias. The selected phyllodes had a north-south orientation so that one side faced eastwards and the other westwards. The principal measurements were made on Acacia longifolia and A. melanoxylon in Portugal in late summer and autumn, and additional measurements were made on A. ligulata and A. melanoxylon in Australia. In Portugal, irrespective of soil moisture status, conductance showed on early morning maximum with a subsequent gradual decline and sometimes a subsidiary peak in the late afternoon. Maximum conductances appeared to be a function of soil moisture status, whereas the decline in conductance in the late morning and afternoon was correlated with changes in phyllode-to-air vapour pressure deficits rather than changes in phyllode water status. The relationship of transpiration to phyllode water potential did not appear to be influenced by soil moisture status, although transpiration was less in drier soils and in the afternoons, this latter factor contributing to a marked hysteresis in the relationship. The opposing faces of the phyllodes exhibited a high degree of synchrony, showing parallel stomatal opening and closing, despite their large differences in irradiance. Stomatal conductance tended to be higher on the eastern faces in the morning and lower in the afternoon. In A. longifolia the daily average of relative conductance was much the same for both faces, but in A. melanoxylon that of the eastern face was higher and was retained even when the normal orientation of the phyllodes was reversed by turning them through 180°. Synchrony must be achieved by the stomata of both sides responding to common environmental or endogenous signals which are perceived by both surfaces with equal sensitivity.     

Stomatal acclimation to vapour pressure deficit doubles transpiration of small tree seedlings with warming

Future climate change is expected to increase temperature (T) and atmospheric vapour pressure deficit (VPD) in many regions, but the effect of persistent warming on plant stomatal behaviour is highly uncertain. We investigated the effect of experimental warming of 1.9–5.1 °C and increased VPD of 0.5–1.3 kPa on transpiration and stomatal conductance (g_s) of tree seedlings in the temperate forest understory (Duke Forest, North Carolina, USA). We observed peaked responses of transpiration to VPD in all seedlings, and the optimum VPD for transpiration (D_opt) shifted proportionally with increasing chamber VPD. Warming increased mean water use of Carya by 140% and Quercus by 150%, but had no significant effect on water use of Acer. Increased water use of ring‐porous species was attributed to (1) higher air T and (2) stomatal acclimation to VPD resulting in higher g_s and more sensitive stomata, and thereby less efficient water use. Stomatal acclimation maintained homeostasis of leaf T and carbon gain despite increased VPD, revealing that short‐term stomatal responses to VPD may not be representative of long‐term exposure. Acclimation responses differ from expectations of decreasing g_s with increasing VPD and may necessitate revision of current models based on this assumption.     

Stomatal control of transpiration

The role of stomata in regulating transpiration from vegetation has historically been controversial among those working either at the single leaf, or at the extensive canopy scales. Recently, the role of unstirred air layers surrounding leaves and canopies in limiting the impact of stomatal movements on transpiration has received renewed recognition. This has led to notable progress in quantitatively describing the effectiveness of stomata in controlling transpiration and in reconciling contrasting viewpoints concerning the role of stomata at the leaf, stand and regional scales. Considerable progress has also been made in understanding how variations in aerial factors such as evaporative demand and edaphic factors such as soil water availability are sensed and transduced into appropriate stomatal regulatory responses. These developments indicate that studies carried out at multiple scales of observation are needed to understand how external environmental factors and intrinsic plant properties interact to determine the role of stomata in regulating transpiration from different types of vegetation.     

Stomatal conductance and transpiration of Acacia under field conditions: similarities and differences between leaves and phyllodes

Summary Leaf diffusive conductance and transpiration rates in response to situations of high evaporative demand were measured in 40 Acacia species varying widely with regard to the morphological and anatomical characters of their assimilatory organs. The measurements took place in south-eastern and central Australia, central Africa and south-western Europe and included species of all three subgenera of Acacia Mill. Soil moisture conditions and consequently the water status of the experimental plants varied between the different measuring sites, some of which were regularly watered. All the species investigated showed a similar daily pattern of diffusive conductance with a morning peak and a subsequent decrease, which was more pronounced in plants growing under water stress, indicating a decisive stomatal regulation of transpiration. A relationship between the structure of assimilatory organs and leaf diffusive conductance or transpiration rates per unit surface area could not be detected in the Australian acacias. However, there are indications that the leaves of the non-Australian species operate on higher conductances than the foliage of the Australian ones. It is suggested that the observed differences in the performance of African and Australian acacias reflect the deciduous or evergreen nature of foliage rather than structural differences. In regard to taxon-specific differentiation this might implicate an ecophysiological character which separates the evergreen species of the geographically isolated subgenus Heterophyllum from the deciduous species of the subgenera Aculeiferum and Acacia with an overlapping area of distribution.     

Hydraulic acclimation to shading in boreal conifers of varying shade tolerance

The purpose of this study was to determine how shading affects the hydraulic and wood‐anatomical characteristics of four boreal conifers (Pinus banksiana, Pinus contorta, Picea glauca and Picea mariana) that differ in shade tolerance. Plants were grown in an open field and under a deciduous‐dominated overstory for 6 years. Sapwood‐ and leaf‐area specific conductivity, vulnerability curves, and anatomical measurements (light and scanning electron microscopy) were made on leading shoots from six to nine trees of each treatment combination. There was no difference in sapwood‐area specific conductivity between open‐grown and understory conifers, although two of four species had larger tracheid diameters in the open. Shaded conifers appeared to compensate for small diameter tracheids by changes in pit membrane structure. Scanning electron microscopy revealed that understory conifers had thinner margo strands, greater maximum pore size in the margo, and more torus extensions. All of these trends may contribute to inadequate sealing of the torus. This is supported by the fact that all species showed increased vulnerability to cavitation when grown in the understory. Although evaporative demand in an understory environment is low, a rapid change into fully exposed conditions could be detrimental for shaded conifers.     

Responses of transpiration and hydraulic conductance to root temperature in nitrogen- and phosphorus-deficient cotton seedlings

Suboptimal N or P availability and cool temperatures all decrease apparent hydraulic conductance (L) of cotton (Gossypium hirsutum L.) roots. The interaction between nutrient status and root temperature was tested in seedlings grown in nutrient solutions. The depression of L (calculated as the ratio of transpiration rate to absolute value of leaf water potential [Ψ_w]) by nutrient stress depended strongly on root temperature, and was minimized at high temperatures. In fully nourished plants, L was high at all temperatures ≥20°C, but it decreased greatly as root temperature approached the chilling threshold of 15°C. Decreasing temperature lowered Ψ_w first, followed by transpiration rate. In N- or P-deficient plants, L approached the value for fully nourished plants at root temperatures ≥30°C, but it decreased almost linearly with temperature as roots were cooled. Nutrient effects on L were mediated only by differences in transpiration, and Ψ_w was unaffected. The responses of Ψ_w and transpiration to root cooling and nutrient stress imply that if a messenger is transmitted from cooled roots to stomata, the messenger is effective only in nutrient-stressed plants.     

Leaf conductance and transpiration, and water relations of evergreen and deciduous perennials co-occurring in a moist chaparral site

Abstract. Diurnal courses of leaf conductance, water potential and environmental parameters were measured through the year. The seasonal decreases in plant water potentials were greatest in evergreen H. arbutifolia , intermediate in winter deciduous C. occidentalis and least in drought deciduous A. californica. The seasonal patterns of water use were very similar. Estimates of soil root-stem conductances to liquid water flux indicate that during the spring, conductances are high, that during the early summer there is a daily shift from high morning conductances to lower afternoon conductances, and that late summer conductances are low.     

Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest

Density-dependent seedling mortality could increase with a species relative abundance, thereby promoting species coexistence. Differences among species in light-dependent mortality also could enhance coexistence via resource partitioning. These compatible ideas rarely have been considered simultaneously. We developed models of mortality as functions of irradiance and local conspecific density (LCD) for seedlings of 53 tropical woody species. Species varied in mortality responses to these factors, but mortality consistently increased with shading and LCD. Across species, density-dependent mortality on a per-neighbour basis was inversely related to species community abundance, but higher LCD in more common species resulted in a weak relationship between species abundance and density-dependent mortality scaled to species maximum LCD. Species mortality responses to shading and maximum LCD were strongly and positively correlated. Our results suggest that species differences in density-dependent mortality are more strongly related to physiologically based life-history traits than biotic feedbacks related to community abundance.     

Stomatal conductance and photosynthesis vary linearly with plant hydraulic conductance in ponderosa pine

Recent work has shown that stomatal conductance (g_s) and assimilation (A) are responsive to changes in the hydraulic conductance of the soil to leaf pathway (K_L), but no study has quantitatively described this relationship under controlled conditions where steady‐state flow is promoted. Under steady‐state conditions, the relationship between g_s, water potential (Ψ) and K_L can be assumed to follow the Ohm's law analogy for fluid flow. When boundary layer conductance is large relative to g_s, the Ohm's law analogy leads to g_s = K_L (Ψ_soil?Ψ_leaf)/D, where D is the vapour pressure deficit. Consequently, if stomata regulate Ψ_leaf and limit A, a reduction in K_L will cause g_s and A to decline. We evaluated the regulation of Ψ_leaf and A in response to changes in K_L in well‐watered ponderosa pine seedlings (Pinus ponderosa). To vary K_L, we systematically reduced stem hydraulic conductivity (k) using an air injection technique to induce cavitation while simultaneously measuring Ψ_leaf and canopy gas exchange in the laboratory under constant light and D. Short‐statured seedlings (< 1 m tall) and hour‐long equilibration times promoted steady‐state flow conditions. We found that Ψ_leaf remained constant near ? 1·5 MPa except at the extreme 99% reduction of k when Ψ_leaf fell to ? 2·1 MPa. Transpiration, g_s, A and K_L all declined with decreasing k (P < 0·001). As a result of the near homeostasis in bulk Ψ_leaf, g_s and A were directly proportional to K_L (R² > 0·90), indicating that changes in K_L may affect plant carbon gain.     

Stomatal conductance of differentially salinized plants

Stomatal resistance was measured daily with a stomatal diffusion porometer during a 4-week period in leaves of bean (Phaseolus vulgaris L., var. Bush Blue Lake) and barley (Hordeum vulgare L., var. Liberty) plants having roots equally split between two differentially salinized nutrient solutions. The stomatal conductance (reciprocal of stomatal resistance) of plants with half their roots in saline solutions was intermediate between the stomatal conductances of plants grown in nonsaline solutions and those grown in saline solutions.     

Effect of partial root excision on transpiration, root hydraulic conductance and leaf growth in wheat seedlings.

Removal of four out of five roots did not lower transpiration and stomatal conductivity of wheat (Triticum durum Desf.) seedlings. Water content of mature expanded leaf lamina remained constant at control levels. The results suggest that the only remaining root was capable to supply the shoot with water. This was evidenced by an increase in hydraulic conductivity of the root system following partial root excision measured at low subatmospheric pressures induced by vacuum. In the absence of a hydrostatic gradient, water flow from reduced root system was initially not higher than from an intact system, but increased subsequently. ABA content was increased in roots 1 h after partial root excision, which might contribute to the increase in hydraulic conductivity.     

Cavitation,stomatal conductance,and leaf dieback in seedlings of two co-occurring Mediterranean shrubs during an intense drought

Seedling shrubs in the Mediterranean semi-arid climate are subjected to intense droughts during summer. Thus, seedlings often surpass their limits of tolerance to water stress, resulting in the loss of hydraulic conductivity due to xylem cavitation. The response in terms of stomatal conductance, vulnerability to cavitation, leaf dieback, and survival were analysed in two co-occurring seedlings of mastic tree (Pistacia lentiscus L.) and kermes oak (Quercus coccifera L.) during an intense drought period. Both species reacted to drought with steep decreases in stomatal conductance before the critical water potential brought about the onset of cavitation events. Q. coccifera showed wider safety margins for avoiding runaway embolism than P. lentiscus and these differences could be related to the particular drought strategy displayed by each species: water saver or water spender. The limits for survival, resprout capacity and leaf dieback were also analysed in terms of loss of conductivity. By contrast with previous studies, the species showing higher seedling survival in the presence of drought also showed higher susceptibility to cavitation and operated with a lower safety margin for cavitation. Both species showed a leaf specific conductivity (LSC) threshold below which leaf biomass had to be regulated to avoid runaway embolism. However, each species displayed a different type of response: P. lentiscus conserved total leaf area up to 100% loss of LSC, whereas Q. coccifera continuously adjusted leaf biomass throughout the drought period in order to maintain the LSC very close to the maximum values recorded without loss of conductivity. Both species maintained the capacity for survival until the loss of conductivity was very nearly 100%.     

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.     

Stomatal conductance, photosynthesis and respiration of temperate deciduous tree seedlings grown outdoors at an elevated concentration of carbon dioxide

Seedlings of temperate deciduous tree species were grown outdoors at ambient and at an elevated concentration of carbon dioxide to examine how aspects of their gas exchange would be altered by growth at elevated carbon dioxide concentration. Leaf conductances to water vapour and net carbon dioxide exchange rates were determined periodically near midday. Whole-plant carbon dioxide efflux rates in darkness were also determined. The stomatal conductance of leaves of plants grown and measured at 700 cm³ m^?3 carbon dioxide did not differ from that of plants grown and measured at 350 cm³ m^?3 in Malus domestica, Quercus prinus and Quercus robur at any measurement time. In Acer saccharinum, lower conductances occurred for plants grown and measured at elevated carbon dioxide concentration only at measurement temperatures above 33°C. Photo-synthetic adjustment to elevated carbon dioxide concentration was evident only in Q. robur. All species examined had lower rates of dark respiration per unit of mass when grown and measured at elevated carbon dioxide concentration.     

Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments

 Light saturated photosynthesis (A) in field saplings of shade tolerant, intermediate, and intolerant tree species was analyzed for stomatal and nonstomatal limitations to test differences between species and sun and shade phenotypes during drought. Throughout the study, photosynthesis was highest and mesophyll limitations of A (L_m) lowest in the intolerant species in both open and understory habitats. The shade tolerant species exhibited the only drought-related decreased A and increased L_m in the open, and the greatest drought-related decreased A and increased L_m in the understory. Few species exhibited significant habitat or drought-related differences in stomatal conductance to CO₂ (g_c), but even slight decreases in g_c during drought were associated with large increases in stomatal limitations to A (L_g). Combined changes in L_m and L_g resulted in increased relative stomatal limitation to A (l _g) in several species during drought. Nevertheless, the overall lack of stomatal closure allowed for nonstomatal limitations to play a major role in reduced A during drought. Higher leaf N was associated with shallower slope of the l _g versus g_c relationship, an indication of greater A capacity. Photosynthetic capacity tended to be greater in the intolerant species than the tolerant species, and it tended to decrease during drought primarily in the shade tolerant species in the understory. Findings in the literature suggest that carbon reduction reactions may be more susceptible to drought than photosynthetic light reactions. If so, reduced carbon reduction capacity of shade tolerant species or shade phenotypes may predispose them to drought conditions, which suggests a mechanism behind the well-recognized tradeoff between drought tolerance and shade tolerance of temperate tree species. Received: 20 October 1995 / Accepted: 20 February 1996     

整树水力导度协同冠层气孔导度调节森林蒸腾

冠层气孔导度决定森林的蒸腾效率,它对驱动水汽移动的水汽应力的响应受树木水力结构的影响,并随水汽压亏缺上升和水力导度下降而降低,维持水势在最低阈值之上,避免出现水力灾变,调控冠层蒸腾。由于叶形和树冠结构的特点,部分脱耦联反映了湿润地区阔叶林冠层与大气的水汽交换特征,单纯以气孔导度的变化难以完整描述水分通量的调节规律,因而,需要考虑冠层气孔导度与水力导度协同控制冠层蒸腾的潜在机理。通过整合叶片气孔气体交换、树干液流、冠层微气象和其他环境因子的野外观测值,估测不同时间尺度的森林冠层气孔导度与大气的脱耦联系数和变异范围,以基于树干液流的冠层蒸腾,结合叶片/土壤水势梯度计算的水力导度,分析水力导度影响冠层气孔导度响应水汽压亏缺的敏感性,可以揭示和阐明水力导度和冠层气孔导度联合调节森林蒸腾的机理,对准确估测全球变化背景下森林对水资源利用的潜在生态效应有明显的理论意义。     

Stomatal and environmental control of transpiration in a lowland tropical forest tree

Stomatal control of crown transpiration was studied in Anacardium excelsum, a large-leaved, emergent canopy species common in the moist forests of Central and northern South America. A construction crane equipped with a gondola was used to gain access to the uppermost level in the crown of a 35-m-tall individual. Stomatal conductance at the single leaf scale, and transpiration and total vapour phase conductance (stomatal and boundary layer) at the branch scale were measured simultaneously using the independent techniques of porometry and stem heat balance, respectively. This permitted the sensitivity of transpiration to a marginal change in stomatal conductance to be evaluated using a dimensionless coupling coefficient (1-ω) ranging from zero to 1, with 1 representing maximal stomatal control of transpiration. Average stomatal conductance varied from 0.09 mol m^?2 s^?1 during the dry season to 0.3 mol m^?2 s^?1 during the wet season. Since boundary layer conductance was relatively low (0.4 mol m^?2 s^?1), 1-ω ranged from 0.46 during the dry season to only 0.25 during the wet season. A pronounced stomatal response to humidity was observed, which strongly limited transpiration as evaporative demand increased. The stomatal response to humidity was apparent only when the leaf surface was used as the reference point for measurement of external vapour pressure. Average transpiration was predicted to be nearly the same during the dry and wet seasons despite a 1 kPa difference in the prevailing leaf-to-air vapour pressure difference. The patterns of stomatal behaviour and transpiration observed were consistent with recent proposals that stomatal responses to humidity are based on sensing the transpiration rate itself.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Stomatal conductance and transpirational responses of field-grown cotton to ozone

Abstract Stomatal conductance and transpiration were measured on normally-irrigated (NI) and water-stressed (WS) field-grown cotton (Gossypium hirsutum L.) exposed throughout the growing season to a gradient of ozone (O₃) concentrations. Environmental conditions during the growing season strongly affected stomatal responses and yield reductions due to O₃ exposure. Maximum stomatal conductance and transpiration decreased with increased O₃ concentration both in NI and WS treatments. Maximum conductance in severely O₃-stressed plants averaged 30% lower than charcoal-filtered (control) plants, but maximum transpiration was only 17% lower. Conductance in WS plots averaged 22% lower than in NI plots but transpiration rates were the same in both treatments. Yield reductions induced by O₃ were highly correlated (r²= 0.84) with daily transpiration. Stomata of O₃-stressed plants opened and closed at the same rate as control plants in response to changes in light intensity, suggesting that the mechanism of stomatal movement had not been impaired by exposure to O₃. Reductions in conductance and transpiration in O₃-stressed plants were attributed to inhibition of photosynthesis by O₃, leading to accumulation of CO₂ in intercellular spaces.     

Stomatal conductance tracks soil-to-leaf hydraulic conductance in faba bean and maize during soil drying

Although regulation of stomatal conductance is widely assumed to be the most important plant response to soil drying, the picture is incomplete when hydraulic conductance from soil to the leaf, upstream of the stomata, is not considered. Here, we investigated to what extent soil drying reduces the conductance between soil and leaf, whether this reduction differs between species, how it affects stomatal regulation, and where in the hydraulic pathway it occurs. To this end, we noninvasively and continuously measured the total root water uptake rate, soil water potential, leaf water potential, and stomatal conductance of 4-week-old, pot-grown maize (Zea mays) and faba bean (Vicia faba) plants during 4 days of water restriction. In both species, the soil–plant conductance, excluding stomatal conductance, declined exponentially with soil drying and was reduced to 50% above a soil water potential of −0.1 MPa, which is far from the permanent wilting point. This loss of conductance has immediate consequences for leaf water potential and the associated stomatal regulation. Both stomatal conductance and soil–plant conductance declined at a higher rate in faba bean than in maize. Estimations of the water potential at the root surface and an incomplete recovery 22 h after rewatering indicate that the loss of conductance, at least partly, occurred inside the plants, for example, through root suberization or altered aquaporin gene expression. Our findings suggest that differences in the stomatal sensitivity among plant species are partly explained by the sensitivity of root hydraulic conductance to soil drying.

The hydraulic conductance between soil and leaf decreases exponentially with decreasing soil water potential, at a species-specific rate related to the decline rate of stomatal conductance.