Assessing transpiration in the tussock grass Stipa tenacissima L.: the crucial role of the interplay between morphology and physiology

Plant transpiration has a key role on both plant performance and ecosystem functioning in arid zones, but realistic estimates at appropriate spatial-temporal scales are scarce. Leaf and tiller morphology and crown architecture were studied together with leaf physiology and whole plant water balance in four individual plants of Stipa tenacissima of different sizes to determine the relative influence of processes taking place at different spatial and temporal scales on whole plant transpiration. Transpiration was estimated in potted plants by leaf-level gas exchange techniques (infrared gas analyzer and porometer), by sap flow measurements, and by integrating leaf physiology and crown architecture with the 3-D computer model Yplant. Daily transpiration of each individual plant was monitored using a gravimetric method, which rendered the reference values. Leaves on each individual plant significantly varied in their physiological status. Young and green parts of the leaves showed five times higher chlorophyll concentration and greater photosynthetic capacity than the senescent parts of the foliage. Instantaneous leaf-level transpiration measurements should not be used to estimate plant transpiration, owing to the fact that extrapolations overestimated individual transpiration by more than 100%. Considering leaf age effects and scaling the estimates according to the relative amount of each foliage category reduced this difference to 46% though it was still significantly higher than gravimetric measurements. Sap flow calculations also overestimated tussock transpiration. However, 3-D reconstruction of plants with Yplant and transpiration estimates, considering both the physiological status and the daily pattern of radiation experienced by each individual leaf section within the crown, matched the gravimetric measurements (differences were only 4.4%). The complex interplay of leaf physiology and crown structure must be taken into account in scaling up plant transpiration from instantaneous, leaf-level measurements, and our study indicates that transpiration of complex crowns is easily overestimated.     

Effects of tussock size and soil water content on whole plant gas exchange in Stipa tenacissima L.: Extrapolating from the leaf versus modelling crown architecture

In this study we evaluated daily whole plant transpiration and net photosynthetic rates in Stipa tenacissima L. (Poaceae) tussocks of different sizes subjected to three levels of soil moisture. The crown architecture of 12 tussocks was reconstructed with the 3D computer model Yplant taking into account the morphology and physiology of the leaves determined at different soil moisture levels. We also calculated whole plant transpiration by extrapolating leaf transpiration in different senescence conditions measured with a diffusion porometer. This extrapolated transpiration overestimated transpiration, particularly when the soil moisture level was high (>15% of volumetric soil water content). At this high level of soil moisture, large tussocks (>60 cm in diameter), which were sexually mature and had a large leaf surface area, were the most efficient with regard to daily water use efficiency (whole plant net photosynthesis/whole plant transpiration). Whole plant water use efficiency decreased with tussock size primarily because small tussocks exhibited high transpiration rates. Small tussocks were more sensitive to soil drying than large and intermediate ones, presenting a faster rate of leaf senescence as water deficit increased. Leaf acclimation to irradiance, which was significantly influenced by the degree of mutual shading among neighbouring leaves, along with the ontogeny of the tussock and its effect upon leaf senescence were found to be the main mechanisms involved in the different responses to water limitations found in whole plant gas exchange variables. Our results show that the size of each individual plant must be taken into account in processes of scaling-up of carbon gain and transpiration from leaf to stand, as this is a particularly relevant aspect in estimating water use by semiarid vegetation.     

A simulation study on the importance of size-related changes in leaf morphology and physiology for carbon gain in an epiphytic bromeliad

This study addresses the question of how size-related changes in leaf morphology and physiology influence light absorption and carbon gain of the epiphytic bromeliad Vriesea sanguinolenta. A geometrically based computer model, Y-plant, was used for the three-dimensional reconstruction of entire plants and for calculation of whole plant light interception and carbon gain. Plants of different sizes were reconstructed, and morphological and physiological attributes of young and old leaves, and small and large plants were combined to examine the individual effects of each factor on light absorption and carbon gain of the plant. The influence of phyllotaxis on light absorption was also explored. Departure of measured divergence angles between successive leaves from the ideal 137.5 degrees slightly decreased light absorption. The only morphological parameter that consistently changed with plant size was leaf shape: larger plants produced more slender foliage, which substantially reduced self-shading. Nevertheless, self-shading increased with plant size. While the maximum rate of net CO(2) uptake of leaves increased linearly with plant size by a factor of two from the smallest to the largest individual, the potential plant carbon gain (based on total foliage area) showed a curvilinear relationship, but with similar numerical variation. We conclude that leaf physiology has a greater impact on plant carbon gain than leaf and plant morphology in this epiphytic bromeliad.     

小叶锦鸡儿灌丛群落蒸腾耗水量估算方法

为了确定沙地小叶锦鸡儿 (Caragana microphylla) 人工林的蒸腾速率, 于2006年6月运用热平衡茎流测量技术, 对科尔沁沙地一处15a生人工小叶锦鸡儿群落的分枝液流动态进行了监测。根据生物统计结果选取被测标准枝, 标准枝基径在0.4~1cm范围内。同时, 分别用基径总断面积推算法和叶片密度推算法对灌丛叶面积进行了估算。以叶面积为扩展纯量, 利用标准枝液流对灌丛群落耗水量进行尺度转换, 在转换过程中, 假设叶面积与蒸腾耗水量之间具有很强的相关性。该尺度转换方法经与大型称重式Lysimeter测值对比验证, 误差小于14.3%, 可望准确估算小叶锦鸡儿灌丛群落的蒸腾耗水量。     

Structural and physiological sexual dimorphism estimated from three‐dimensional virtual trees of yerba‐mate (Ilex paraguariensis) is modified by cultivation environment

Yerba‐mate is a subtropical, evergreen, dioecious, South American tree. Sexual dimorphism in photosynthesis, leaf allometry and foliage distribution was hypothesised. Virtual trees (constructed in VPlants software from detailed measurements of plant morphogenesis) of the two genders were compared considering two contrasted cultivation environments and three developmental stages. The total crown volume, leaf area per plant (LA), leaf area index (LAI) and leaf area density (LAD) were calculated. The light interception and photosynthesis were computed from mock‐ups in VegeSTAR. Structural sexual dimorphism concerned general plant form, internode length, leaf allometry, leaf surface, pattern of leaf area distribution and LAD. Cultivation environment and developmental stage acted strongly on sex expression of all observed structural parameters and physiological stages. Sexual differentiation in LA and light interception was related to leaves positioned in the lowest layers (150 cm above ground), whereas sexual specialisation in leaf and plant photosynthesis was related to early vegetative and reproductive stages. Several sexual responses strongly depended on the environment, especially light conditions, with opposite effects observed on female and male plants whether they were cultivated in monoculture or in forest understorey, under high‐light condition or low‐light condition, respectively. Optimised foliage structure and physiology in females may compensate for greater reproductive costs in early developmental stages, but females and males equalise in photosynthetic efficiency after 2‐year regrowth.     

Sheathing the blade: Significant contribution of sheaths to daytime and nighttime gas exchange in a grass crop

Despite representing a sizeable fraction of the canopy, very little is known about leaf sheath gas exchange in grasses. Specifically, estimates of sheath stomatal conductance, transpiration and photosynthesis along with their responses to light, CO₂ and vapour pressure deficit (VPD) are unknown. Furthermore, the anatomical basis of these responses is poorly documented. Here, using barley as a model system, and combining leaf-level gas exchange, whole-plant gravimetric measurements, transpiration inhibitors, anatomical observations, and biophysical modelling, we found that sheath and blade stomatal conductance and transpiration were similar, especially at low light, in addition to being genotypically variable. Thanks to high abaxial stomata densities and surface areas nearly half those of the blades, sheaths accounted for up to 17% of the daily whole-plant water use, which -surprisingly- increased to 45% during the nighttime. Sheath photosynthesis was on average 17–25% that of the blade and was associated with lower water use efficiency. Finally, sheaths responded differently to the environment, exhibiting a lack of response to CO₂ but a strong sensitivity to VPD. Overall, these results suggest a key involvement of sheaths in feedback loops between canopy architecture and gas exchange with potentially significant implications on adaptation to current and future climates in grasses.     

浙江天童太白山不同群落植物构型比较

植物构型是植株构件在空间上的分配方式,反映了植物对环境的响应策略。通过对浙江天童太白山海拔差异很小的栲树(Castanopsis fargesii)、小叶青冈(Cyclobalanopsis gracilis)和云山青冈(Cyclobalanopsis sessilifolia)群落类型中所有植株的树高、树冠厚度、树冠面积、叶片聚集度、枝下高和距地45 cm基径等植物构型性状,以及树冠曝光指数、土壤含水率、空气温湿度、土壤p H值和风速6个环境因子的测定,分别分析乔灌木层植物构型性状及性状关系在3个群落间的变化规律。结果表明:(1)从栲树到小叶青冈至云山青冈群落,灌木层的树高、树冠厚度、枝下高和距地45 cm基径增加,叶片聚集度减小;乔木层的树高、树冠厚度、树冠面积、枝下高和距地45 cm基径均减小,叶片聚集度增大;(2)3个群落灌木层构型性状间显著相关(P0.001),而乔木层只在中低海拔群落存在相关性;(3)从栲树到小叶青冈至云山青冈群落,乔灌木层的冠层曝光指数显著增加(P0.05);(4)多元逐步回归表明,树冠曝光指数对灌木层构型性状变异的贡献最大,而风速、土壤含水率和p H值对乔木构型性状的变异起主导作用。综上得知,天童太白山乔灌木植物在不同群落间存在构型分异,植物对光资源的竞争是引起灌木构型在不同群落间变化的主要驱动因子,而对乔木植物,其构型变化更多受到风速和土壤含水量的影响。     

Observed and modelled historical trends in the water‐use efficiency of plants and ecosystems

Plant water‐use efficiency (WUE, the carbon gained through photosynthesis per unit of water lost through transpiration) is a tracer of the plant physiological controls on the exchange of water and carbon dioxide between terrestrial ecosystems and the atmosphere. At the leaf level, rising CO₂ concentrations tend to increase carbon uptake (in the absence of other limitations) and to reduce stomatal conductance, both effects leading to an increase in leaf WUE. At the ecosystem level, indirect effects (e.g. increased leaf area index, soil water savings) may amplify or dampen the direct effect of CO₂. Thus, the extent to which changes in leaf WUE translate to changes at the ecosystem scale remains unclear. The differences in the magnitude of increase in leaf versus ecosystem WUE as reported by several studies are much larger than would be expected with current understanding of tree physiology and scaling, indicating unresolved issues. Moreover, current vegetation models produce inconsistent and often unrealistic magnitudes and patterns of variability in leaf and ecosystem WUE, calling for a better assessment of the underlying approaches. Here, we review the causes of variations in observed and modelled historical trends in WUE over the continuum of scales from leaf to ecosystem, including methodological issues, with the aim of elucidating the reasons for discrepancies observed within and across spatial scales. We emphasize that even though physiological responses to changing environmental drivers should be interpreted differently depending on the observational scale, there are large uncertainties in each data set which are often underestimated. Assumptions made by the vegetation models about the main processes influencing WUE strongly impact the modelled historical trends. We provide recommendations for improving long‐term observation‐based estimates of WUE that will better inform the representation of WUE in vegetation models.     

Scale‐dependent variation in visual estimates of grassland plant cover

Abstract. Plant cover was visually estimated by five observers, independent of each other, in a species‐rich grassland in the Bílé Karpaty Mts., southeastern Czech Republic, in seven plots ranging from 0.001 to 4 m². Variation of total plant cover among the observers was high at small scales: 0.001–0.016 m²; coefficient of variation, CV = 35 to 45%, but much lower at larger scales: 0.06–4 m²; CV = 7 to 15%. Differences between visual estimates of plant cover of individual species made by different observers were affected by plot size, total cover and morphology of particular plants. CV of the cover of individual species ranged from 0 to 225% and decreased with increasing plot size. For abundant plants the CV attained ca. 50%, independent of plot size. In spite of a very high number of sterile plants with similar leaf morphology and colour, the observed variation in cover estimates in the studied grassland was comparable with results reported from other vegetation types. Differences between estimates by individual observers were often larger than usual year to year changes in undisturbed grasslands. Therefore, I suggest that to avoid difficulties in the interpretation of results based on plant cover data obtained from visual estimates, several observers should always work together, adjusting their extreme estimates.     

Competition-dependent modelling of foliage biomass in forest stands

Currently, foliage biomass is estimated based on stem diameter or basal area. However, it is questionable whether the relations between foliage and stem observed from plantations of a single tree species can be applied to stands of different structure or species composition. In this paper, a procedure is presented to simulate foliage and branch biomass of tree crowns relative to crown size and light competition. Crowns are divided into layers and segments and each segment is divided into a foliated and an unfoliated fraction. Depending on the competitive status of the segment, leaf area density, specific leaf area and foliated branch fraction are determined. Based on this information, foliage biomass is calculated. The procedure requires a crown shape function and a measure to characterise competition for light and space of each individual segment within the canopy. Simple solutions are suggested for both requirements to enable an application with data that can be measured non-destructively in the field; these were stem position, tree height, crown base height, crown radii and some general crown shape information. The model was parameterised from single trees of Norway spruce and European beech and partly evaluated with independent data close to the investigation plot. Evaluations showed that the model can attribute the ecology of the different crown forms. Modelled foliage distribution for beech and spruce as well as total needle biomass of spruce agreed well with measurements but foliage biomass of beech was underestimated. The results are discussed in the context of a general model application in structured forests.     

Light interception efficiency explained by two simple variables: a test using a diversity of small- to medium-sized woody plants

? Plant light interception efficiency is a crucial determinant of carbon uptake by individual plants and by vegetation. Our aim was to identify whole-plant variables that summarize complex crown architecture, which can be used to predict light interception efficiency. ? We gathered the largest database of digitized plants to date (1831 plants of 124 species), and estimated a measure of light interception efficiency with a detailed three-dimensional model. Light interception efficiency was defined as the ratio of the hemispherically averaged displayed to total leaf area. A simple model was developed that uses only two variables, crown density (the ratio of leaf area to total crown surface area) and leaf dispersion (a measure of the degree of aggregation of leaves). ? The model explained 85% of variation in the observed light interception efficiency across the digitized plants. Both whole-plant variables varied across species, with differences in leaf dispersion related to leaf size. Within species, light interception efficiency decreased with total leaf number. This was a result of changes in leaf dispersion, while crown density remained constant. ? These results provide the basis for a more general understanding of the role of plant architecture in determining the efficiency of light harvesting.     

Convergence in light capture efficiencies among tropical forest understory plants with contrasting crown architectures: a case of morphological compensation

Leaf and crown characteristics were examined for 24 tree and herbaceous species of contrasting architectures from the understory of a lowland rainforest. Light-capture efficiency was estimated for the crowns of the different species with a three-dimensional geometric modeling program. Causal relationships among traits affecting light absorption at two hierarchical levels (leaf and whole crown) were quantified using path analysis. Light-capture and foliage display efficiency were found to be very similar among the 24 species studied, with most converging on a narrow range of light absorption efficiencies (ratio of absorbed vs. available light of 0.60-0.75). Exceptionally low values were found for the climber vines and, to a lesser extent, for the Bromeliad Aechmea magdalenae. Differences in photosynthetic photon flux density (PFD) absorbed per unit leaf area by individual plants were mostly determined by site to site variation in PFD and not by the differences in crown architecture among individuals or species. Leaf angle, and to a lesser extent also supporting biomass, specific leaf area, and internode length, had a significant effect on foliage display efficiency. Potential constraints on light capture such as the phyllotactic pattern were generally offset by other compensatory adjustments of crown structure such as internode length, arching stems, and plagiotropy. The variety of shoot morphologies capable of efficiently capturing light in tropical forest understories is greater than initially thought, extending over species with very different phyllotactic patterns, crown architectures, leaf sizes, and morphologies.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

A three-dimensional statistical reconstruction model of grapevine (Vitis vinifera) simulating canopy structure variability within and between cultivar/training system pairs

Background and Aims

In grapevine, canopy-structure-related variations in light interception and distribution affect productivity, yield and the quality of the harvested product. A simple statistical model for reconstructing three-dimensional (3D) canopy structures for various cultivar–training system (C × T) pairs has been implemented with special attention paid to balance the time required for model parameterization and accuracy of the representations from organ to stand scales. Such an approach particularly aims at overcoming the weak integration of interplant variability using the usual direct 3D measurement methods.

Model

This model is original in combining a turbid-medium-like envelope enclosing the volume occupied by vine shoots with the use of discrete geometric polygons representing leaves randomly located within this volume to represent plant structure. Reconstruction rules were adapted to capture the main determinants of grapevine shoot architecture and their variability. Using a simplified set of parameters, it was possible to describe (1) the 3D path of the main shoot, (2) the volume occupied by the foliage around this path and (3) the orientation of individual leaf surfaces. Model parameterization (estimation of the probability distribution for each parameter) was carried out for eight contrasting C × T pairs.

Key Results and Conclusions

The parameter values obtained in each situation were consistent with our knowledge of grapevine architecture. Quantitative assessments for the generated virtual scenes were carried out at the canopy and plant scales. Light interception efficiency and local variations of light transmittance within and between experimental plots were correctly simulated for all canopies studied. The approach predicted these key ecophysiological variables significantly more accurately than the classical complete digitization method with a limited number of plants. In addition, this model accurately reproduced the characteristics of a wide range of individual digitized plants. Simulated leaf area density and the distribution of light interception among leaves were consistent with measurements. However, at the level of individual organs, the model tended to underestimate light interception.Key words: Canopy, architecture, hemispherical, picture, light interception, radiative, balance, stochastic, modelling, virtual, plants     

Clouds homogenize shoot temperatures,transpiration, and photosynthesis within crowns of <Emphasis Type="Italic">Abies fraseri</Emphasis> (Pursh.) Poiret

Multiple studies have examined the effects of clouds on shoot and canopy-level microclimate and physiological processes; none have yet done so on the scale of individual plant crowns. We compared incident photosynthetically active radiation (PAR), leaf temperatures, chlorophyll fluorescence, and photosynthetic gas exchange of shoots in three different spatial locations of Abies fraseri crowns on sunny (clear to partly cloudy) versus overcast days. The field site was a Fraser fir farm (1038 m elevation) in the Appalachian mountains, USA. Ten saplings of the same age class were marked and revisited for all measurements. Sunny conditions corresponded with 5–10× greater sunlight incidence on south-facing outer shoots compared to south-facing inner and north-facing outer shoots, which were shaded and received only indirect (diffuse) sunlight. Differences in spatial distribution of irradiance were mirrored in differences in shoot temperatures, photosynthesis, and transpiration, which were all greater in south-facing outer shoots compared to more shaded crown locations. In contrast, overcast conditions corresponded with more homogeneous sunlight distribution between north and south-facing outer shoots, and similar shoot temperatures, chlorophyll fluorescence (ΦPSII), photosynthesis, and transpiration; these effects were observed in south-facing inner shoots as well, but to a lesser extent. There was no significant difference in conductance between different crown locations on sunny or overcast days, indicating spatial differences in transpiration under sunny conditions were likely driven by leaf temperature differences. We conclude that clouds can affect spatial distribution of sunlight and associated physiological parameters not only within forest communities, but within individual crowns as well.     

Foliage randomness and light interception in 3-D digitized trees: an analysis from multiscale discretization of the canopy

Light models for vegetation canopies based on the turbid medium analogy are usually limited by the basic assumption of random foliage dispersion in the canopy space. The objective of this paper was to assess the effect of three possible sources of non-randomness in tree canopies on light interception properties. For this purpose, four three-dimensional (3-D) digitized trees and four theoretical canopies – one random and three built from fractal rules – were used to compute canopy structure parameters and light interception, namely the sky-vault averaged STAR (Silhouette to Total Area Ratio). STAR values were computed from (1) images of the 3-D plants, and (2) from a 3-D turbid medium model using space discretization at different scales. For all trees, departure from randomness was mainly due to the spatial variations in leaf area density within the canopy volume. Indeed STAR estimations, based on turbid medium assumption, using the finest space discretization were very close to STAR values computed from the plant images. At this finest scale, foliage dispersion was slightly clumped, except one theoretical fractal canopy, which showed a marked regular dispersion. Taking into account a non-infinitely small leaf size, whose effect is theoretically to shorten self-shading, had a minor effect on STAR computations. STAR values computed from the 3-D turbid medium were very sensitive to plant lacunarity, a parameter introduced in the context of fractal studies to characterize the distribution of gaps in porous media at different scales. This study shows that 3-D turbid medium models based on space discretization are able to give correct estimation of light interception by 3-D isolated trees, provided that the 3-D grid is properly defined, that is, discretization maximizes plant lacunarity.     

Physiological mechanisms drive differing foliar calcium content in ferns and angiosperms

Recent evidence points to ferns containing significantly lower contents of foliar calcium and other cations than angiosperms. This is especially true of more ancient ‘non-polypod’ fern lineages, which predate the diversification of angiosperms. Calcium is an important plant nutrient, the lack of which can potentially slow plant growth and litter decomposition, and alter soil invertebrate communities. The physiological mechanisms limiting foliar calcium (Ca) content in ferns are unknown. While there is a lot we do not know about Ca uptake and transport in plants, three physiological processes are likely to be important. We measured transpiration rate, cation exchange capacity, and leaching loss to determine which process most strongly regulates foliar Ca content in a range of fern and co-occurring understory angiosperm species from a montane Hawaiian rainforest. We found higher instantaneous and lifetime (corrected for leaf lifespan) transpiration rates in angiosperms relative to ferns. Ferns preferentially incorporated Ca into leaves relative to strontium, which suggests that root or stem cation exchange capacity differs between ferns and angiosperms, potentially affecting calcium transport in plants. There were no differences in foliar Ca leaching loss between groups. Among the physiological mechanisms measured, foliar Ca was most strongly correlated with leaf-level transpiration rate and leaf lifespan. This suggests that inter-specific differences in a leaf’s lifetime transpiration may play a significant role in determining plant nutrition.     

Stem flow and porometer measurements of transpiration from honey mesquite (Prosopis glandulosa)

The objective of this study was to compare stem flow and porometermethods of measuring transpiration of honey mesquite (Prosopisglandulosa) trees on a semiarid site. Stem flow was measuredusing heat balance stem flow gauges. Porometer measurementsof leaf stomatal conductance (g_s) were made within foliage layersof each stem and scaled to transpiration values for the entirestem (E_stem) using stem leaf area. Simultaneous measurementsusing both methods were made diurnally and under artificiallyimposed stem shading or defoliation in June and October 1990.Stem flow and E_stem had similar diumal patterns except on 2d in June when E_stem increased during the afternoon while stemflow declined relative to midday values. During October, E_stemwas greater than stem flow throughout the day. This was attributedto sampling error in which only undamaged leaves were used forporometer measurements yet, by this time in the growing season,many leaves on each stem were damaged from insects or wind andlikely had lower transpiration rates. A regression coefficientbetween E_stem and stem flow of 0.79 in June and 0.91 in Octobersuggested the two methods were comparable, but there was considerablevariation between methods during peak transpiration rates. Bothtechniques detected that artificial shading or defoliation causedsimilar relative declines in transpiration. Results imply thatestimates of stem transpiration can be obtained by scaling porometermeasurements of leaves but accuracy declines at higher transpirationrates. Key words: Sap flow, evapotranspiration, stomatal conductance, scaling, water relations