Climate-driven changes in biomass allocation in pines

Future increases in air temperature resulting from human activities may increase the water vapour pressure deficit (VPD) of the atmosphere. Understanding the responses of trees to spatial variation in VPD can strengthen our ability to predict how trees will respond to temporal changes in this important variable. Using published values, we tested the theoretical prediction that conifers decrease their investment in photosynthetic tissue (leaves) relative to water‐conducting tissue in the stem (sapwood) as VPD increases. The ratio of leaf/sapwood area (A_L/A_S) decreased significantly with increasing VPD in Pinus species but not in Abies, Pseudotsuga, Tsuga and Picea, and the average A_L/A_S was significantly lower for pines than other conifers (pines: 0.17 m² cm^?2; nonpines: 0.44 m² cm^?2). Thus, pines adjusted to increasing aridity by altering above‐ground morphology while nonpine conifers did not. The average water potential causing a 50% loss of hydraulic conductivity was ?3.28 MPa for pines and ?4.52 MPa for nonpine conifers, suggesting that pines are more vulnerable to xylem embolism than other conifers. For Pinus ponderosa the decrease in A_L/A_S with high VPD increases the capacity to provide water to foliage without escalating the risk of xylem embolism. Low A_L/A_S and plasticity in this variable may enhance drought tolerance in pines. However, lower A_L/A_S with increasing VPD and an associated shift in biomass allocation from foliage to stems suggests that pines may expend more photosynthate constructing and supporting structural mass and carry less leaf area as the climate warms.     

Size-dependent variability of leaf and shoot hydraulic conductance in silver birch

Variation in leaf and shoot hydraulic conductance was examined on detached shoots of silver birch (Betula pendula Roth), cut from the lower third (shade leaves) and upper third of the crown (sun leaves) of large trees growing in a natural temperate forest stand. Hydraulic conductances of whole shoots (K _S), leaf blades (K _lb), petioles (K _P) and branches (i.e. leafless stem; K _B) were determined by water perfusion using a high-pressure flow meter in quasi-steady state mode. The shoots were exposed to irradiance of photosynthetic photon flux density of 200–250 μmol m⁻² s⁻¹, using different light sources. K _lb depended significantly (P < 0.001) on light quality, canopy position and leaf blade area (A _L). K _lb increased from crown base to tree top, in parallel with vertical patterns of A _L. However, the analysis of data on shade and sun leaves separately revealed an opposite trend: the bigger the A _L the higher K _lb. Leaf anatomical study of birch saplings revealed that this trend is attributable to enhanced vascular development with increasing leaf area. Hydraulic traits (K _S, K _B, K _lb) of sun shoots were well co-ordinated and more strongly correlated with characteristics of shoot size than those of shade shoots, reflecting their greater evaporative load and need for stricter adjustment of hydraulic capacity with shoot size. K _S increased with increasing xylem cross-sectional area to leaf area ratio (Huber value; P < 0.01), suggesting a preferential investment in water-conducting tissue (sapwood) relative to transpiring tissue (leaves), and most likely contributing to the functional stability of the hydraulic system, essential for fast-growing pioneer species.     

Influence of nutrient versus water supply on hydraulic architecture and water balance in Pinus taeda

We investigated the hydraulic consequences of a major decrease in root‐to‐leaf area ratio (A_R:A_L) caused by nutrient amendments to 15‐year‐old Pinus taeda L. stands on sandy soil. In theory, such a reduction in A_R:A_L should compromise the trees’ ability to extract water from drying sand. Under equally high soil moisture, canopy stomatal conductance (G_S) of fertilized trees (F) was 50% that of irrigated/fertilized trees (IF), irrigated trees (I), and untreated control trees (C). As predicted from theory, F trees also decreased their stomatal sensitivity to vapour pressure deficit by 50%. The lower G_S in F was associated with 50% reduction in leaf‐specific hydraulic conductance (K_L) compared with other treatments. The lower K_L in F was in turn a result of a higher leaf area per sapwood area and a lower specific conductivity (conducting efficiency) of the plant and its root xylem. The root xylem of F trees was also 50% more resistant to cavitation than the other treatments. A transport model predicted that the lower A_R:A_L in IF trees resulted in a considerably restricted ability to extract water during drought. However, this deficiency was not exposed because irrigation minimized drought. In contrast, the lower A_R:A_L in F trees caused only a limited restriction in water extraction during drought owing to the more cavitation resistant root xylem in this treatment. In both fertilized treatments, approximate safety margins from predicted hydraulic failure were minimal suggesting increased vulnerability to drought‐induced dieback compared with non‐fertilized trees. However, IF trees are likely to be so affected even under a mild drought if irrigation is withheld.     

Interactive effects of elevated CO2 and temperature on water transport inponderosa pine

Many studies report that water flux through trees declines in response to elevated CO₂, but this response may be modified by exposure to increased temperatures. To determine whether elevated CO₂ and temperature interact to affect hydraulic conductivity, we grew ponderosa pine seedlings for 24 wk in growth chambers with one of four atmospheric CO₂ concentrations (350, 550, 750, and 1100 ppm) and either a low (15°C nights, 25°C days) or high (20°C nights, 30°C days) temperature treatment. Vapor pressure deficits were also higher in the elevated temperature treatment. Seedling biomass increased with CO₂ concentration but was not affected by temperature. Root : shoot ratio was unaffected by CO₂ and temperature. Leaf : sapwood area ratio (A_L/A_S) declined in response to elevated temperature but was not influenced by CO₂. Larger tracheid diameters at elevated temperature caused an increase in xylem-specific hydraulic conductivity (K_S). The increase in K_S and decrease in A_L/A_S led to higher leaf-specific hydraulic conductivity (K_L) at elevated temperature. Stomatal conductance (g_S) was correlated with K_L across all treatments. Neither K_S, K_L, nor g_S were affected by elevated CO₂ concentrations. High K_L in response to elevated temperature may support increased transpiration or reduce the incidence of xylem cavitation in ponderosa pine in future, warmer climates.     

Influence of soil porosity on water use in<Emphasis Type="Italic"> Pinus taeda</Emphasis>

We analyzed the hydraulic constraints imposed on water uptake from soils of different porosities in loblolly pine (Pinus taeda L.) by comparing genetically related and even-aged plantations growing in loam versus sand soil. Water use was evaluated relative to the maximum transpiration rate (E _crit) allowed by the soil-leaf continuum. We expected that trees on both soils would approach E _crit during drought. Trees in sand, however, should face greater drought limitation because of steeply declining hydraulic conductivity in sand at high soil water potential (Ψ _S). Transport considerations suggest that trees in sand should have higher root to leaf area ratios (A _R:A _L), less negative leaf xylem pressure (Ψ _L), and be more vulnerable to xylem cavitation than trees in loam. The A _R:A _L was greater in sand versus loam (9.8 vs 1.7, respectively). This adjustment maintained about 86% of the water extraction potential for both soils. Trees in sand were more deeply rooted (>1.9 m) than in loam (95% of roots <0.2 m), allowing them to shift water uptake to deeper layers during drought and avoid hydraulic failure. Midday Ψ _L was constant for days of high evaporative demand, but was less negative in sand (–1.6 MPa) versus loam (–2.1 MPa). Xylem was more vulnerable to cavitation in sand versus loam trees. Roots in both soils were more vulnerable than stems, and experienced the greatest predicted loss of conductivity during drought. Trees on both soils approached E _crit during drought, but at much higher Ψ _S in sand (<–0.4 MPa) than in loam (<–1.0 MPa). Results suggest considerable phenotypic plasticity in water use traits for P. taeda which are adaptive to differences in soil porosity. Received: 28 December 1999 / Accepted: 31 March 2000     

Inter-species variation of photosynthetic and xylem hydraulic traits in the deciduous and evergreen Euphorbiaceae tree species from a seasonally tropical forest in south-western China

The objective of the present study was to examine the functional coordination among hydraulic traits, xylem characteristics and gas exchange rates across three deciduous Euphorbiaceae tree species (Hevea brasiliensis, Macaranga denticulata and Bischofia javanica) and three evergreen Euphorbiaceae tree species (Drypetes indica, Aleurites moluccana and Codiaeum variegatum) from a seasonally tropical forest in south-western China. The deciduous tree species were more vulnerable to water stress-induced embolism than the evergreen tree species. However, the deciduous tree species generally had higher maximal rates of sapwood and leaf-specific hydraulic conductivity (K _S and K _L), respectively. Compared with the evergreen tree species, the deciduous tree species, however, possessed a lower density of sapwood and a wider diameter of xylem vessels. Regardless of leaf phenology, the hydraulic vulnerability and conductivity were significantly correlated with sapwood density and mean vessel diameter. Furthermore, the hydraulic vulnerability was positively correlated with water transport efficiency. In addition, the deciduous tree species exhibited higher maximal photosynthetic rates (A _max) and stomatal conductance (g _max), but lower water use efficiency (WUE). Interestingly, the A _max, g _max and WUE were strongly correlated with K _S and K _L across the deciduous and evergreen tree species. These results suggest that xylem structure, rather than leaf phenology, accounts for the difference in hydraulic traits between the deciduous tree species and the evergreen tree species. Meanwhile, our results show that there is a significant trade-off between hydraulic efficiency and safety, and a strong functional correlation between the hydraulic capacity and gas exchange rates across the deciduous and evergreen tree species.     

The hydraulic architecture of Pinaceae – a review

We reviewed the literature to examine the vulnerability to water stress-induced embolism of Pinaceae relative to other conifers and to study the inter-relationships among the main traits involved in the hydraulic function within the Pinaceae. Results showed that Pinaceae (particularly the genus Pinus) are more vulnerable to xylem embolism, and show less variability in this character, than other conifers. Detailed data from 12 populations of Pinaceae (11 species) from three different areas (Piñol and Sala 2000; Martínez-Vilalta and Piñol 2002; Oliveras et al. 2003) was used to study the relationships among hydraulic properties of stems. These included: leaf-to-wood area ratio (A_L:A _W), wood- and leaf-specific hydraulic conductivity (K_W and K_L, respectively), vulnerability to xylem embolism (Ψ_50PLC), carbon isotope composition of needles (δ¹³C) and minimum needle water potential (minimum Ψ_L). Results showed that hydraulic properties tended to be more correlated among each other than with indicators of environmental (precipitation to potential evapotranspiration ratio, P/E) or physiological water stress (minimum Ψ_L). The only exception was an increase of δ¹³C with decreasing minimum Ψ_L and P/E. Overall, A_L:A _W ratio decreased with increasing vulnerability to xylem embolism, and with increasing K_W and K_L (P<0.05). We found a strong positive relationship between carbon isotope composition and the estimated maximum loss of conductivity due to xylem embolism under field conditions, suggesting stronger stomatal control in more vulnerable species with higher levels of native embolism. Overall, results are consistent with a range of drought-avoidance strategies to minimise the gradient of water potential through the xylem, and show that different relationships among traits are possible depending on the scale of study (individual vs. species or populations). The strong interdependence among hydraulic traits implies that no single trait is a sufficient predictor of drought-resistance in Pinaceae. Finally, it is hypothesised that the intrinsically vulnerable xylem of pines may limit their survival under extremely dry conditions.     

Photosynthetic characteristics of dipterocarp seedlings in three tropical rain forest light environments: a basis for niche partitioning?

In three tropical rain forest light environments in Sabah, Malaysia, we compared photosynthesis in seedlings of ten climax tree species with putatively differing shade tolerances. The objectives of the study were (a) to characterise the range of photosynthetic responses in ten species of the Dipterocarpaceae and (b) to elucidate those photosynthetic characteristics that might provide a basis for niche partitioning. Seedlings were acclimated (c. 7 months) in three light environments; understorey, partial shade and a gap (140 m²). The light environments represented a gradation in median diurnal (0630–1830 hours) photon flux density (PFD) ranging from understorey (4.7 μmol m⁻² s⁻¹), through partial shade (21.2 μmol m⁻² s⁻¹) to gap (113.7 μmol m⁻² s⁻¹). Integrated diurnal PFD were in the sequence gap > partial shade > understorey (15.2, 4.7, 1.3 mol m⁻² day⁻¹, respectively). In gap-acclimated plants, species differed in the photosynthetic light-response variables apparent quantum yield, dark respiration rate, light compensation point, net saturated leaf assimilation rate (A _sat), and in stomatal conductance (g _s sat) when assimilation rate (A) was saturated. A light-demanding pioneer species (Macaranga hypoleuca) and a shade-demanding understorey species (Begonia sp.) had, respectively, higher and lower A _sat and g _s sat than the dipterocarp species. In high-light conditions A _sat and g _s sat were strongly positively correlated in dipterocarp species. Differing photosynthetic characteristics of gap-acclimated plants suggest that, in these dipterocarp species, different rates of carbon fixation may be an important factor contributing towards niche partitioning. Mean integrated diurnal A (A _diurnal) in the gap, partial shade and understory were, respectively, 122.9, 52.7, 20.5 mmol m⁻² day⁻¹. Differences occurred in A _diurnal of dipterocarp species between light environments. When Macaranga was included, differences in A _diurnal were evident in the gap and partial shade, and in both cases were attributed to the pioneer. For the variable A _diurnal, there was of a shift in the rank position of Macaranga among light environments, but a shift did not occur among the dipterocarp species. Results from this study are consistent with the idea that rates of carbon fixation per unit leaf area may contribute towards niche differentiation between the climax and single pioneer species, but not within the group of climax species. Other physiological and/or carbon allocation factors may be involved in any niche partitioning; dipterocarp species often have inherently different growth rates and susceptibility to herbivory. As an alternative to niche partitioning, dipterocarp species may co-exist in natural light environments as a result of habitat disequilibrium or purely stochastic processes. Received: 2 April 1997 / Accepted: 13 July 1997     

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity (k_L) of upper branches was positively correlated with maximum rates of net CO₂ assimilation per unit leaf area (A_area) and stomatal conductance (g_s) across 20 species of canopy trees. Maximum k_L showed stronger correlation with A_area than initial k_L suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k_L was negatively correlated with A_area/g_s and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics (k_theoretical) was positively related to A_area and k_L, consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k_L, A_area, net CO₂ assimilation per unit leaf mass (A_mass), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.     

The fraction of expanding to expanded leaves determines the biomass response of Populus to elevated CO2

We examined whether the effects of elevated CO₂ on growth of 1-year old Populus deltoides saplings was a function of the assimilation responses of particular leaf developmental stages. Saplings were grown for 100 days at ambient (approximately 350 ppm) and elevated (ambient + 200 ppm) CO₂ in forced-air greenhouses. Biomass, biomass distribution, growth rates, and leaf initiation and expansion rates were unaffected by elevated CO₂. Leaf nitrogen (N), the leaf C:N ratio, and leaf lignin concentrations were also unaffected. Carbon gain was significantly greater in expanding leaves of saplings grown at elevated compared to ambient CO₂. The Rubisco content in expanding leaves was not affected by CO₂ concentration. Carbon gain and Rubisco content were significantly lower in fully expanded leaves of saplings grown at elevated compared to ambient CO₂, indicating CO₂-induced down-regulation in fully expanded leaves. Elevated CO₂ likely had no overall effect on biomass accumulation due to the more rapid decline in carbon gain as leaves matured in saplings grown at elevated compared to ambient CO₂. This decline in carbon gain has been documented in other species and shown to be related to a balance between sink/source balance and acclimation. Our data suggest that variation in growth responses to elevated CO₂ can result from differences in leaf assimilation responses in expanding versus expanded leaves as they develop under elevated CO₂. Received: 28 September 1998 / Accepted: 23 June 1999     

High vapour pressure deficit exacerbates xylem cavitation and photoinhibition in shade-grown Piper auritum H.B. & K. during prolonged sunflecks

Water relations dynamics during simulated sunflecks at high (36°C) and medium (27°C) temperatures and high and low vapour pressure deficits beween leaf and air (VPD) were studied on shade-grown Piper auritum H.B. & K. plants, a pioneer tree, common in gaps and clearings of tropical rain forests. The leaves of P. auritum wilt rapidly when exposed to high light. Exposure to high VPD and high light caused substantial and rapid dehydration of leaves. Dehydration could be prevented under high humidity irrespective of temperature. Water stored in leaf cells served as initial source for transpiration upon high light exposure. This effect increased with increasing VPD and temperature. The pronounced decrease in leaf water content over time in high light caused a rapid decrease in leaf water potential (Ψ_l) and a concomitant increase in water potential gradient (ΔΨ/Δx) between trunk and leaf, yet the high leaf elasticity (small bulk elastic modulus, ε) allowed turgor maintenance under most conditions. Under high VPD and high temperature, stomata remained open and ΔΨ/Δx frequently exceeded 0.95 MPa · m⁻¹, the cavitation-inducing threshold (ΔΨ/Δx _cav) causing high rates of acoustic emissions from stems and leaf petioles and leading to concomitant losses in hydraulic conductance per leaf area (k _l). At medium temperature (high VPD), stomatal closure contained xylem embolism by keeping ΔΨ/Δx at or below this threshold. We argue that wilting substantially contributes to creating a sufficient driving force for water uptake from the soil, and reducing the VPD (through a decrease in radiation load and thus leaf temperature) to avoid excessive dehydration. Received: 3 March 1996 / Accepted: 10 November 1996     

Photosynthetic pathway alters xylem structure and hydraulic function in herbaceous plants

Plants using the C₄ photosynthetic pathway have greater water use efficiency (WUE) than C₃ plants of similar ecological function. Consequently, for equivalent rates of photosynthesis in identical climates, C₄ plants do not need to acquire and transport as much water as C₃ species. Because the structure of xylem tissue reflects hydraulic demand by the leaf canopy, a reduction in water transport requirements due to C₄ photosynthesis should affect the evolution of xylem characteristics in C₄ plants. In a comparison of stem hydraulic conductivity and vascular anatomy between eight C₃ and eight C₄ herbaceous species, C₄ plants had lower hydraulic conductivity per unit leaf area (K_L) than C₃ species of similar life form. When averages from all the species were pooled together, the mean K_L for the C₄ species was 1.60 × 10^?4 kg m^?1 s^?1 MPa^?1, which was only one‐third of the mean K_L of 4.65 × 10^?4 kg m^?1 s^?1 MPa^?1 determined for the C₃ species. The differences in K_L between C₃ and C₄ species corresponded to the two‐ to three‐fold differences in WUE observed between C₃ and C₄ plants. In the C₄ species from arid regions, the difference in K_L was associated with a lower hydraulic conductivity per xylem area, smaller and shorter vessels, and less vulnerable xylem to cavitation, indicating the C₄ species had evolved safer xylem than the C₃ species. In the plants from resource‐rich areas, such as the C₄ weed Amaranthus retroflexus, hydraulic conductivity per xylem area and xylem anatomy were similar to that of the C₃ species, but the C₄ plants had greater leaf area per xylem area. The results indicate the WUE advantage of C₄ photosynthesis allows for greater flexibility in hydraulic design and potential fitness. In resource‐rich environments in which competition is high, an existing hydraulic design can support greater leaf area, allowing for higher carbon gain, growth and competitive potential. In arid regions, C₄ plants evolved safer xylem, which can increase survival and performance during drought events.     

Hydraulic properties and photosynthetic rates in co-occurring lianas and trees in a seasonal tropical rainforest in southwestern China

In this study, we examined wood anatomy, hydraulic properties, photosynthetic rate, and water status and osmotic regulation in three liana species and three tree species co-occurring in a seasonal tropical rain forest. Our results showed that the three liana species had larger vessel diameter, lower sapwood density, and consequently higher branch sapwood specific hydraulic conductivity (K _S) than the three tree species. Across species, K _S was positively correlated with leaf nitrogen concentration and maximum net CO₂ assimilation rate. However, it was also positively correlated with xylem water potential at 50% loss of hydraulic conductivity, indicating a trade-off between hydraulic efficiency and safety. Compared to the tree species, the liana species had higher predawn leaf water potential and lower osmotic adjustment in the dry season. The combination of more efficient water transport, higher photosynthetic rates, and their ability to access to more reliable water source at deeper soil layers in the dry season in the lianas should contribute to their fast growth.     

Hydraulic constraints on photosynthesis in subtropical evergreen broad leaf forest and pine woodland trees of the Florida Everglades

The relationship between water transport and photosynthesis represents the trade-off between carbon gain and water loss and was used to evaluate potential differences in water resource utilization among two dominant vegetation types of south Florida: subtropical evergreen broad leaf forests (hardwood hammocks) and pine woodlands (pine rocklands). We found consistent linear positive relationships between the quantum yield of photosystem II (ϕ _PSII), an index of photosynthetic capacity, and hydraulic conductivity per sapwood area (k _S) and per leaf area (k _L) across all species. The slope of the ϕ _PSII–k _S relationship was steeper for hardwood hammock than for pine rockland species. Mean ϕ _PSII was greater in pine rockland species and was greater for a given k _L than in hardwood hammock species. These results are consistent with previous observations demonstrating that pine rocklands tend to have better access to stable water sources than hardwood hammocks. We also found greater photosynthetic carbon isotope discrimination with increasing k _S and k _L in pine rockland species, but not in hardwood hammock species, suggesting increased stomatal conductance with increasing k _S and k _L, consistent with greater water availability in pine rockland habitats. Our study thus utilizes relationships between water transport and photosynthesis to evaluate hydraulic constraints on physiological function between two contrasting vegetation types with contrasting stability of water sources.     

Pentachlorophenol biodegradation kinetics of an oligotrophic fluidized-bed enrichment culture

A fluidized-bed reactor (FBR) was used to enrich an aerobic chlorophenol-degrading microbial culture. Long-term continuous-flow operation with low effluent concentrations selected oligotrophic microorganisms producing good-quality effluent for pentachlorophenol(PCP)-contaminated water. PCP biodegradation kinetics was studied using this FBR enrichment culture. The results from FBR batch experiments were modeled using a modified Haldane equation, which resulted in the following kinetic constants: q _max = 0.41 mg PCP mg protein⁻¹ day⁻¹, K _S = 16 μg l⁻¹, K _i = 5.3 mg l⁻¹, and n = 3.5. These results show that the culture has a high affinity for PCP but is also inhibited by relatively low PCP concentrations (above 1.1 mg PCP l⁻¹). This enrichment culture was maintained over 1 year of continuous-flow operation with PCP as the sole source of carbon and energy. During continuous-flow operation, effluent concentrations below 2 μg l⁻¹ were achieved at 268 min hydraulic retention time (t _HR) and 2.5 mg PCP l⁻¹ feed concentration. An increase in loading rate by decreasing t _HR did not significantly deteriorate the effluent quality until a t _HR decrease from 30 min to 21 min resulted in process failure. Recovery from process failure was slow. Decreasing the feed PCP concentration and increasing t _HR resulted in an improved process recovery. Received: 10 October 1996 / Received revision: 21 January 1997 / Accepted: 24 January 1997     

Hydraulic responses to height growth in maritime pine trees

As trees grow taller, decreased xylem path conductance imposes a major constraint on plant water and carbon balance, and is thus a key factor underlying forest productivity decline with age. The responses of stomatal conductance, leaf area: sapwood area ratio (A_L : A_S) and soil–leaf water potential gradient (ΔΨ_S–L) to height growth were investigated in maritime pine trees. Extensive measurements of in situ sap flow, stomatal conductance and (non‐gravitational) needle water potential (_L = Ψ_L ? ρ_wgh) were made during 2 years in a chronosequence of four even‐aged stands, under both wet and dry soil conditions. Under wet soil conditions, _L was systematically lower in taller trees on account of differences in gravitational potential. In contrast, under dry soil conditions, our measurements clearly showed that _L was maintained above a minimum threshold value of ?2.0 MPa independently of tree height, thus limiting the range of compensatory change in ΔΨ_S–L. Although a decrease in the A_L : A_S ratio occurred with tree height, this compensation was not sufficient to prevent a decline in leaf‐specific hydraulic conductance, K_L (50% lower in 30 m trees than in 10 m trees). An associated decline in stomatal conductance with tree height thus occurred to maintain a balance between water supply and demand. Both the increased investment in non‐productive versus productive tissues (A_S : A_L) and stomatal closure may have contributed to the observed decrease in tree growth efficiency with increasing tree height (by a factor of three from smallest to tallest trees), although other growth‐limiting responses (e.g. soil nutrient sequestration, increased respiratory costs) cannot be excluded.     

Impact of phloem girdling on leaf gas exchange and hydraulic conductance in hybrid aspen

We investigated phloem-xylem interactions in relation to leaf hydraulic capacity in hybrid aspen (Populus tremula L. × P. tremuloides Michx.) by using phloem girdling method. Removal of bark tissues (phloem girdling) at the branch base resulted in a substantial decline in stomatal conductance (g_S), net photosynthetic rate (P_N), and leaf hydraulic efficiency, and in increase of leaf water potential (Ψ_L). Although gS declined more than P_N (83 versus 78 %), the ratio of intercellular to ambient CO₂ concentrations (c_i/c_a) increased from 0.67 to 0.87 in three days after girdling. Girdling induced a decrease in leaf hydraulic conductance (K_L) on average by 43 % (P = 0.006). The changes in gS and leaf conductance to water vapour were co-ordinated with K_L only in girdled branches whereas intrinsic water-use efficiency was invariant to K_L. The declines in K_L with girdling were not accompanied by changes in potassium ion concentration ([K⁺]), electrical conductivity, or pH of xylem sap. The results suggest that phloem girdling at the branch base does not influence the recirculation of ions between the phloem and xylem in hybrid aspen and the decrease of K_L in response to the manipulation is not related to changes in [K⁺] and total ionic content of xylem sap.     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Recent improvements in the development of A2B adenosine receptor agonists

Adenosine is known to exert most of its physiological functions by acting as local modulator at four receptor subtypes named A₁, A_2A, A_2B and A₃ (ARs). Principally as a result of the difficulty in identifying potent and selective agonists, the A_2B AR is the least extensively characterised of the adenosine receptors family. Despite these limitations, growing understanding of the physiological meaning of this target indicates promising therapeutic perspectives for specific ligands. As A_2B AR signalling seems to be associated with pre/postconditioning cardioprotective and anti-inflammatory mechanisms, selective agonists may represent a new therapeutic group for patients suffering from coronary artery disease. Herein we present an overview of the recent advancements in identifying potent and selective A_2B AR agonists reported in scientific and patent literature. These compounds can be classified into adenosine-like and nonadenosine ligands. Nucleoside-based agonists are the result of modifying adenosine by substitution at the N ⁶-, C²-positions of the purine heterocycle and/or at the 5′-position of the ribose moiety or combinations of these substitutions. Compounds 1-deoxy-1-{6-[N′-(furan-2-carbonyl)-hydrazino]-9H-purin-9-yl}-N-ethyl-β-D-ribofuranuronamide (19, hA₁ K _i = 1050 nM, hA_2A K _i = 1550 nM, hA_2B EC₅₀ = 82 nM, hA₃ K _i > 5 μM) and its 2-chloro analogue 23 (hA₁ K _i = 3500 nM, hA_2A K _i = 4950 nM, hA_2B EC₅₀ = 210 nM, hA₃ K _i > 5 μM) were confirmed to be potent and selective full agonists in a cyclic adenosine monophosphate (cAMP) functional assay in Chinese hamster ovary (CHO) cells expressing hA_2B AR. Nonribose ligands are represented by conveniently substituted dicarbonitrilepyridines, among which 2-[6-amino-3,5-dicyano-4-[4-(cyclopropylmethoxy)phenyl]pyridin-2-ylsulfanyl]acetamide (BAY-60–6583, hA₁, hA_2A, hA₃ EC₅₀ > 10 μM; hA_2B EC₅₀ = 3 nM) is currently under preclinical-phase investigation for treating coronary artery disorders and atherosclerosis. This article has previously been published in issue 4/4, under doi:.     

Functional relationships between hydraulic traits and the timing of diurnal depression of photosynthesis

The hydraulic coordination along the water transport pathway helps trees provide adequate water supply to the canopy, ensuring that water deficits are minimized and that stomata remain open for CO₂ uptake. We evaluated the stem and leaf hydraulic coordination and the linkages between hydraulic traits and the timing of diurnal depression of photosynthesis across seven evergreen tree species in the southern Andes. There was a positive correlation between stem hydraulic conductivity (k_s) and leaf hydraulic conductance (K_Leaf) across species. All species had similar maximum photosynthetic rates (A_max). The species with higher k_s and K_Leaf attained A_max in the morning, whereas the species with lower k_s and K_Leaf exhibited their A_max in the early afternoon concurrently with turgor loss. These latter species had very negative leaf water potentials, but far from the pressure at which the 88% of leaf hydraulic conductance is lost. Our results suggest that diurnal gas exchange dynamics may be determined by leaf hydraulic vulnerability such that a species more vulnerable to drought restrict water loss and carbon assimilation earlier than species less vulnerable. However, under stronger drought, species with earlier CO₂ uptake depression may increase the risk of hydraulic failure, as their safety margins are relatively narrow.