Control of transpiration in three coffee cultivars: the role of hydraulic and crown architecture

Water use and hydraulic architecture were studied in the coffee (Coffea arabica) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown architecture, basal sapwood area and total leaf surface area. Transpiration per unit leaf area (E), stomatal conductance (g _s), crown conductance (g _c), total hydraulic conductance of the soil/leaf pathway (G _t) and the stomatal decoupling coefficient, omega (Ω) (Jarvis and McNaughton 1986) were assessed over a range of soil moisture and during partial defoliation treatments. The relationship between sap flow and sapwood area was linear and appeared to be similar for the three cultivars. Variation in g _c, E, and G _t of intact plants and leaf area-specific hydraulic conductivity (k _l) of excised lateral branches was negatively correlated with variation in the ratio of leaf area to sapwood area. Transpiration, g _c, and g _s were positively correlated with G _t. Transpiration and G _t varied with total leaf area and were greatest at intermediate values (10 m²) of leaf area. Omega was greatest in Yellow Caturra, the cultivar with the greatest leaf area and a dense crown, and was smallest in Typica, the cultivar with an open crown. Differences in omega were attributable primarily to differences in leaf boundary layer conductance among the cultivars. Plants of each cultivar that were 40% defoliated maintained sap flows comparable to pretreatment plants, but expected compensatory increases in g _s were not consistently observed. Despite their contrasting crown morphologies and hydraulic architecture, the three cultivars shared common relationships between water use and hydraulic architectural traits. Received: 17 February 1999 / Accepted: 28 July 1999     

The upscaling of transpiration from individual trees to areal transpiration in tree belts

Sap flow measurements have long been used to measure transpiration in individual trees and there exist some well established methods for upscaling individual tree volumetric transpiration to areal transpiration in plantation and forest plots. However, where edge effects are significant, such as in tree belts, the area the volumetric transpiration is to be projected upon is unknown. This paper provides a methodology for estimaing the area that a tree belt hydrologically occupies by using either measurements of tree root density or soil moisture distribution. An application of the proposed methodology shows that simply assuming that the area of the tree belt is the crown projected area could lead to an overestimation of the areal transpiration of 100%. An erratum to this article can be found at     

Sap flow rates of mangrove trees are not unusually low

 In contrast with previous reports, we observed high transpiration rates in mangrove trees. Maximum sap velocities and mean daytime sap flow rates were estimated from heat pulse velocity in entire, field grown trees of Avicennia cf. alba Blume and Rhizophora apiculata Blume. Results were within the range of values measured by identical techniques for trees in lowland dipterocarp and tropical heath forests with a similar climate in Brunei Darussalam (north Borneo). High stomatal conductance (400 mmol m^{–  2} s^{–  1}) was also measured for well insolated leaves of A. cf. alba, with midday water potentials reaching about  – 3 MPa in both species. Received: 11 September 1996 / Accepted: 27 January 1997     

A simple field validation of daily transpiration derived from sapflow using a porometer and minimal meteorological data

Heat-pulse techniques are routinely used to estimate transpiration from canopies of woody plants typically without any local calibration, mainly because of the difficulty of doing so in the field and, frequently, lack of detailed weather data. This is despite concerns that the techniques may produce erroneous values under certain conditions, such as when evaporative demand is high. In this study, we used a micrometeorological approach to validate transpiration from irrigated olives deduced from heat-pulse technique by ascertaining precise values for the parameters that are critical for converting heat-pulse velocity to sapflow. The micrometeorological approach involved limited data on stomatal conductance (g_s), obtained hourly with a porometer on four contrasting days, and was used to calibrate a simple model for predicting conductance. Predicted stomatal conductance (g_sm) agreed well with that measured, and when both were used to calculate hourly transpiration, they produced values that were within 10% of each other. This was despite brief underestimations of transpiration based on g_sm (T_m) in the early hours of the day that arose from poor determination of incident radiation at this time. We then used T_m to iteratively set the values for the various parameters, including the time-out value that accounts for zero-flow conditions, needed to convert heat-pulse velocity to sapflow, for the four days. The best fit between T_m and transpiration from sapflow (T_s) was obtained with time-out value set to 120 s. All heat-pulse velocity data were therefore analysed with this time-out value to obtain sapflow and, hence, transpiration (T_s). Comparison of T_m and T_s for the whole season showed that the former tended to produce higher values on certain days when vapour pressure deficit (D) was high in summer (December–February). While T_s occasionally produced larger values than T_m under the mild conditions of autumn (March–April). Totals of the daily transpiration during the 190-day period were within 10% of each other. Responsible Editor: Yan Li     

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.     

Leaf age and salinity influence water relations of pepper leaves

Plant growth is reduced under saline conditions even when turgor in mature leaves is maintained by osmotic adjustment. The objective of this study was to determine if young leaves from salt-affected plants were also osmotically adjusted. Pepper plants (Capsicum annuum L. cv. California Wonder) were grown in several levels of solution osmotic potential and various components of the plants' water relations were measured to determine if young, rapidly growing leaves could accumulate solutes rapidly enough to maintain turgor for normal cell enlargement. Psychrometric measurements indicated that osmotic adjustment is similar for both young and mature leaves although osmotic potential is slightly lower for young leaves. Total water potential is also lower for young leaves, particularly at dawn for the saline treatments. The result is reduced turgor under saline conditions at dawn for young but not mature leaves. This reduced turgor at dawn, and presumably low night value, is possibly a cause of reduced growth under saline conditions. No differences in leaf turgor occur at midday. Porometer measurements indicated that young leaves at a given salinity level have a higher stomatal conductance than mature leaves, regardless of the time of day. The result of stomatal closure is a linear reduction of transpiration.     

Stomatal control of transpiration in the canopy of a clonal Eucalyptus grandis plantation

 Predawn leaf water potential, stomatal conductance and microclimatic variables were measured on 13 sampling days from November 1995 through August 1996 to determine how environmental and physiological factors affect water use at the canopy scale in a plantation of mature clonal Eucalyptus grandis Hill ex-Maiden hybrids in the State of Espirito Santo, Brazil. The simple ”big leaf” Penman-Monteith model was used to estimate canopy transpiration. During the study period the predawn leaf water potential varied from –0.4 to –1.3 MPa, with the minimum values observed in the winter months (June and August 1996), while the average estimated values for canopy conductance and canopy transpiration fell from 17.3 to 5.8 mm s^–1 and from 0.54 to 0.18 mm h^–1, respectively. On the basis of all measurements, the average value of the decoupling coefficient was 0.25. During continuous soil water shortage a proportional reduction was observed in predawn leaf water potential and in daily maximum values of stomatal conductance, canopy transpiration and decoupling coefficient. The results showed that water vapour exchange in this canopy is strongly dominated by the regional vapour pressure deficit and that canopy transpiration is controlled mainly by stomatal conductance. On a seasonal basis, stomatal conductance and canopy transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture and rainfall. Good results were obtained with a multiplicative empirical model that uses values of photosynthetically active radiation, vapour pressure deficit and predawn leaf water potential to estimate stomatal conductance. Received: 10 June 1998 / Accepted: 20 July 1998     

Seasonal changes in apparent hydraulic conductance and their implications for water use of European beech (<Emphasis Type=Italic>Fagus sylvatica</Emphasis> L.) and sessile oak [<Emphasis Type=Italic>Quercus petraea</Emphasis> (Matt.) Liebl] in South Europe

The water status of Fagus sylvatica L. and Quercus petraea (Matt) Liebl. was analysed during a cycle of progressive natural drought in southern Europe. Predawn (Ψ_pd) and midday water potential were measured in transpiring (Ψ_leaf) and non-transpiring leaves (Ψ_xyl). Furthermore, photosynthesis (A), stomatal conductance to water vapour (g_s) and sap flow (F_d) were recorded on the same dates. Apparent leaf specific hydraulic conductance in the soil–plant–air continuum (K_h) and whole tree hydraulic conductance (K_hsf) were calculated by using the simple analogy of the Ohm’s law. K_h was estimated at different points in the pathway as the ratio between transpiration (E) in the uppermost canopy leaves at midday and the gradient of water potential in the different compartments of the continuum soil–roots–stem–branches–leaves. There was a progressive decrease in water potential measured on non-transpiring leaves at the base of tree crown in both species (Ψ^l_xyl) from the beginning of the growing season to the end of summer. A similar decrease was shown in shoot water potential (Ψ^u_xyl) at the uppermost canopy. Predawn water potential (Ψ_pd) was high in both species until late July (28 July); afterwards, a significant decrease was registered in F. sylvatica and Q. petraea with minimum values of −0.81±0.03 and −0.75±0.06 MPa, respectively, by 15 September. In both species, leaf specific hydraulic conductance in the overall continuum soil–plant–air (K_h) decreased progressively as water stress increases. Minimum values of K_h and K_hsf were recorded when Ψ_pd was lower. However, Q. petraea showed higher K_h than F. sylvatica for the same Ψ_pd. The decrease in K_h with water stress was mainly linked to its fall from the soil to the lowermost canopy (K_srs). Nevertheless, a significant resistance in the petiole–leaf lamina (K_pl) was also recorded because significant differences in all dates were found on Ψ between transpiring and non-transpiring leaves from the same shoot. The decline in K_h was followed by an increase in stomatal control of daily water losses through the decrease of stomatal conductance to water vapour (g_s) during the day. It promoted a seasonal increase in the stomatal limitation to carbon dioxide uptake for photosynthesis (A). These facts were more relevant in F. sylvatica, which had concurrently a higher decline in water use at the tree level than Q. petraea. The results showed a strong coupling in F. sylvatica and Q. petraea between processes at leaf and tree level. It may be hypothesised a role of specific hydraulic conductance not only in the regulation of water losses by transpiration but also of carbon uptake.     

Water relations and gas exchange in olive trees under regulated deficit irrigation and partial rootzone drying

It is widely believed that partial root drying (PRD) reduces water losses by transpiration without affecting yield. However, experimental work carried out to date does not always support this hypothesis. In many cases a PRD treatment has been compared to a full irrigated treatment, so doubt remains on whether the observed benefits correspond to the switching of irrigation or just to PRD being a deficit irrigation treatment. In addition, not always a PRD treatment has been found advantageous as compared to a companion regulated deficit irrigation (RDI) treatment. In this work we have compared the response of mature ‘Manzanilla‘ olive trees to a PRD and an RDI treatment in which about 50% of the crop evapotranspiration (ET_c) was supplied daily by localised irrigation. We alternated irrigation in the PRD treatment every 2 weeks in 2003 and every 3 weeks in 2004. Measurements of stem water potential (Ψ_stem), stomatal conductance (g _s) and net CO₂ assimilation rate (A) were made in trees of both treatments, as well as in trees irrigated to 100% of ET_c (Control trees) and in Rain-fed trees. Sap flow was also measured in different conductive organs of trees under both PRD and RDI treatments, to evaluate the influence of alternating irrigation on root water uptake and tree water consumption. We found small and random differences in Ψ_stem, g _s and A, which gave no evidence of PRD causing a positive effect on the olive tree performance, as compared to RDI. Stomatal conductance decreased in PRD trees as compared to Control trees, but a similar decrease in g _s was also recorded in the RDI trees. Sap flow measurements, which reflected water use throughout the irrigation period, also showed no evidence of g _s being more reduced in PRD than in RDI trees. Daily water consumption was also similar in the trees of the deficit irrigation treatments, for most days, throughout the irrigation period. Alternating irrigation in PRD trees did not cause a change in either water taken up by main roots at each side of the trees, or in the sap flow of both trunk locations and main branches of each side. Results from this work, and from previous work conducted in this orchard, suggest that transpiration is restricted in trees under deficit irrigation, in which roots are left in drying soil when water is applied by localised irrigation, and that there is no need to alternate irrigation for achieving this effect. Section Editor: R. E. Munns     

The effect of short-term flooding on the sap flow, gas exchange and hydraulic conductivity of young apricot trees

The effect of short-term flooding was examined in 2-year-old apricot trees (Prunus armeniaca cv. Búlida). Six apricot trees of similar appearance were submitted to two treatments: three were irrigated daily, while the others were flooded for a period of 50 h by submerging the pots in plastic water tanks. The trees were removed from the water, drained and then placed in the same conditions as the control plants. A decrease in transpiration in the flooded trees with respect to the control plants was evident. The daily pattern of soil O₂ concentration and plant hydraulic resistance followed a similar trend during the flooding. However, this relationship was not maintained throughout the experiment, since the O₂ values increased rapidly when the waterlogging ceased, while plant hydraulic resistance only recovered at the end of the experiment when the original root system, damaged by flooded conditions, was replaced with new roots. In flooded trees, the midday leaf water potential decreased progressively from the beginning of flooding, but gradually recovered when the waterlogging ceased. Leaf conductance values of treated plants were slow to recover, reaching values of the control plants 8 days after the leaf water potential had recovered. The close relationship observed during most of the experiment between the leaf water parameters, leaf conductance and plant hydraulic conductance indicate that hydraulic messages are likely to play a dominant role in co-ordinating the observed responses of the shoot.     

Influence of stomatal distribution on transpiration in low-wind environments

Abstract According to computer energy balance simulations of horizontal thin leaves, the quantitative effects of stomatal distribution patterns (top vs. bottom surfaces) on transpiration (E) were maximal for sunlit leaves with high stomatal conductances (g_s) and experiencing low windspeeds (free or mixed convection regimes). E of these leaves decreased at windspeeds > 50 cm s^?1, despite increases in the leaf-to-air vapour density deficit. At 50 cm s^?1 wind-speed, rapidly transpiring leaves had greater E when one-half of the stomata were on each leaf surface (amphistomaty; 10.16 mmol H₂O m^?2 s^?1) than when all stomata were on either the top (hyperstomaty; 9.34 mmol m^?2s^?1) or bottom (hypostomaty; 7.02 mmol m^?2s^?1) surface because water loss occurred in parallel from both surfaces. Hyperstomatous leaves had larger E than hypostomatous leaves because free convection was greater on the top than on the bottom surface. Transpiration of leaves with large g, was greatest at windspeeds near zero when ～60–75% of the stomata were on the top surface, while at high windspeeds E was greatest with, 50% of the stomata on top. For leaves with low g_s, stomatal distribution exerted little influence on simulated E values. Laboratory measurements of water loss from simulated hypo-, hyper-, and amphistomatous leaf models qualitatively supported these predictions.     

Plasticity in hydraulic architecture of Scots pine across Eurasia

Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G _s) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G _s to D (m/G _s,ref, where m is the slope and G _s,ref is reference G _s at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G _s,ref, and its absolute sensitivity to D (m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.     

Estimation of hydraulic conductance within field-grown apricot using sap flow measurements

Using the heat pulse and other techniques, the hydraulic architecture of apricot trees was mapped out. The flows (overall flow, flow across the four main branches) and forces (water potential differences between xylem and leaves) measured allowed us to quantify hydraulic conductance of branches and of the root/soil resistance. The experiment was carried out in a commercial orchard of 11-year-old apricot trees (Prunus armeniaca L., cv. Búlida, on Real Fino apricot rootstock) during 1 week (October 27–November 3, 1998). Three representative trees with a cylindrical trunk divided into four main branches of different sizes, orientation and local microclimate were chosen for the experiment. Sap flow was measured throughout the experimental period. Twelve sets of heat-pulse probes were used, one for each main branch. The diurnal course of the environmental conditions, the fraction of the area irradiated and leaf water relations were also considered in each main branch. The relationships between leaf water potential, xylem water potential and transpiration were established for different branches and also for the total plant. Using the slopes of these regressions, total plant conductance, the hydraulic conductance of the stem and root pathway, the hydraulic conductance of the canopy and the hydraulic conductance of each branch were estimated. Our findings show that the root conductance and the canopy hydraulic conductance are similar in magnitude. Leaf hydraulic conductance per leaf area unit was similar for each of the four branch orientations, indicating that, while the light microclimate has a dominant influence on transpiration, in this case it had little effect on the hydraulic properties of the canopy.     

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.     

Temperature and humidity effects on branchlet gas-exchange in white spruce: an explanation for the increase in transpiration with branchlet temperature

In situ gas-exchange data, for branchlets of white spruce [Picea glauca (Moench) Voss.] in a mature mixed-wood boreal forest in central Canada (53°44′N 105°14′W), were subjected to a multiple regression analysis. Vapor pressure deficit (VPD) and branchlet temperature (t_leaf) were both significant predictors (P<0.0001) of stomatal conductance to water vapor (g_sw) and net photosynthesis (A_n), together explaining 67 and 64% of the variation in g_sw and A_n, respectively. Since VPD and t_leaf were autocorrelated in these field data, but also to further explore the nature of independent effects of temperature and humidity on water and CO₂ exchange in white spruce, steady-state gas-exchange was performed on well-watered greenhouse-grown seedlings of white spruce. Results from laboratory experiments supported the following conclusions: (1) Transpiration (E) increases with VPD to an inflection point that increases linearly with t_leaf. This t_leaf effect on E could not be explained by trends in VPD, RH, A_n or PFD. Rather, our data support a model in which E and g_sw are influenced by the balance between ’supply’ and ’loss’ of water to and from leaf tissue, respectively. The supply of water appears to be in accordance with Darcy’s law, where supply of water is proportional to the driving gradient in pressure/ tension, specific permeability (k), and inverse of water viscosity (n ^–1). Approximately half of the increase in E could be explained by the linear increase in n ^–1 with increasing t_leaf. We propose that increases in k explain the remainder of the increase in E with t_leaf. (2) VPD and t_leaf appear to have independent effects on g_sw. In contrast, RH effects on g_sw or E were subtle and could be explained by a combination of effects of t_leaf and VPD. (3) A_n was affected primarily by t_leaf, being reduced at low (10°C) and high (40°C) temperatures, and only indirectly by humidity parameters via stomatal conductance, viz. intercellular CO₂ concentrations. Our results have implications for the prediction of water fluxes from plants and canopies in areas where plant temperatures vary diurnally or seasonally. Received: 24 September 1998 / Accepted: 20 July 1999     

Regional variation in canopy transpiration of Central European beech forests

Forest hydrologists have hypothesised that canopy transpiration (E_c) of European temperate forests occurs at rather similar rates in stands with different tree species and hydrologic regimes. We tested this hypothesis by synchronously measuring xylem sap flow in four mature stands of Fagus sylvatica along a precipitation gradient with the aim (1) of exploring the regional variability of annual canopy transpiration (E_c(t)) in this species, and (2) of analysing the relationship between precipitation (P) and E_c(t). E_c(t) rates of 216, 225, 272 and 303 mm year^–1 corresponded to precipitation averages of 520, 710, 801 and 1,040 mm year^–1 in the four stands. We explored the regional variability of E_c(t) in Central European colline to sub-montane beech stands in two meta-analyses based on (1) existing sap flow data on beech (n=5 observations), or (2) all canopy transpiration data on beech obtained by different techniques (sap flow, micrometeorological or soil water budget approaches, n=25). With a coefficient of variation (CV) of 20%, the regional variability of E_c(t) (213–421 mm year^–1) was smaller than the variation in corresponding precipitation (550–1,480 mm year^–1). The mean E_c(t) for beech was 289 (±58) mm year^–1 (n=25). A humped-shaped relationship between E_c(t) and P, with a broad transpiration maximum in the precipitation range from ca. 700 to 1,000 mm year^–1, was found which may indicate soil moisture limitation of transpiration for P –1, and reduced transpiration by increased cloudiness or leaf wetness for P>1,000 mm year^–1. Thus, the precipitation level significantly influences canopy transpiration of humid temperate forests; however, the size of the P influence on E_c(t) and, in part, the direction of its effect differ from forests in semi-arid or arid climates. European beech has the capacity to maintain high E_c rates in both humid and partly dry summer climates (P<550 mm year^–1).     

Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO<Subscript>2</Subscript>

Using a free-air CO₂ enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO₂] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm⁻²) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g _s, μmol m⁻² s⁻¹), photosynthetic CO₂ fixation (A, mmol m⁻² s⁻¹), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO₂ (C_i:C_a). Elevated [CO₂] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO₂] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO₂], g _s was reduced, A/E was stimulated, and C_i:C_a was reduced relative to ambient [CO₂]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO₂]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO₂], although the mechanism for this remains obscure.     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

Mechanisms contributing to seasonal homeostasis of minimum leaf water potential and predawn disequilibrium between soil and plant water potential in Neotropical savanna trees

Seasonal regulation of leaf water potential (_L) was studied in eight dominant woody savanna species growing in Brazilian savanna (Cerrado) sites that experience a 5-month dry season. Despite marked seasonal variation in precipitation and air saturation deficit (D), seasonal differences in midday minimum _L were small in all of the study species. Water use and water status were regulated by a combination of plant physiological and architectural traits. Despite a nearly 3-fold increase in mean D between the wet and dry season, a sharp decline in stomatal conductance with increasing D constrained seasonal variation in minimum _L by limiting transpiration per unit leaf area (E). The leaf surface area per unit of sapwood area (LA/SA), a plant architectural index of potential constraints on water supply in relation to transpirational demand, was about 1.5–8 times greater in the wet season compared to the dry season for most of the species. The changes in LA/SA from the wet to the dry season resulted from a reduction in total leaf surface area per plant, which maintained or increased total leaf-specific hydraulic conductance (G_t) during the dry season. The isohydric behavior of Cerrado tree species with respect to minimum _L throughout the year thus was the result of strong stomatal control of evaporative losses, a decrease in total leaf surface area per tree during the dry season, an increase in total leaf-specific hydraulic conductance, and a tight coordination between gas and liquid phase conductance. In contrast with the seasonal isohydric behavior of minimum _L, predawn _L in all species was substantially lower during the dry season compared to the wet season. During the dry season, predawn _L was more negative than bulk soil estimated by extrapolating plots of E versus _L to E=0. Predawn disequilibrium between plant and soil was attributable largely to nocturnal transpiration, which ranged from 15 to 22% of the daily total. High nocturnal water loss may also have prevented internal water storage compartments from being completely refilled at night before the onset of transpiration early in the day.