Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Diurnal depression of leaf hydraulic conductance in a tropical tree species

Diurnal patterns of hydraulic conductance of the leaf lamina (K_leaf) were monitored in a field‐grown tropical tree species in an attempt to ascertain whether the dynamics of stomatal conductance (g_s) and CO₂ uptake (A_leaf) were associated with short‐term changes in K_leaf. On days of high evaporative demand mid‐day depression of K_leaf to between 40 and 50% of pre‐dawn values was followed by a rapid recovery after 1500 h. Leaf water potential during the recovery stage was less than ?1 MPa implying a refilling mechanism, or that loss of K_leaf was not linked to cavitation. Laboratory measurement of the response of K_leaf to Ψ_leaf confirmed that leaves in the field were operating at water potentials within the depressed region of the leaf ‘vulnerability curve’. Diurnal courses of K_leaf and Ψ_leaf predicted from measured transpiration, xylem water potential and the K_leaf vulnerability function, yielded good agreement with observed trends in both leaf parameters. Close correlation between depression of K_leaf, g_s and A_leaf suggests that xylem dysfunction in the leaf may lead to mid‐day depression of gas exchange in this species.     

The 'hydrology' of leaves: co-ordination of structure and function in temperate woody species

The hydraulic conductance of the leaf lamina (K_lamina) substantially constrains whole‐plant water transport, but little is known of its association with leaf structure and function. K_lamina was measured for sun and shade leaves of six woody temperate species growing in moist soil, and tested for correlation with the prevailing leaf irradiance, and with 22 other leaf traits. K_lamina varied from 7.40 × 10^?5 kg m^?2 s^?1 MPa^?1 for Acer saccharum shade leaves to 2.89 × 10^?4 kg m^?2 s^?1 MPa^?1 for Vitis labrusca sun leaves. Tree sun leaves had 15–67% higher K_lamina than shade leaves. K_lamina was co‐ordinated with traits associated with high water flux, including leaf irradiance, petiole hydraulic conductance, guard cell length, and stomatal pore area per lamina area. K_lamina was also co‐ordinated with lamina thickness, water storage capacitance, 1/mesophyll water transfer resistance, and, in five of the six species, with lamina perimeter/area. However, for the six species, K_lamina was independent of inter‐related leaf traits including leaf dry mass per area, density, modulus of elasticity, osmotic potential, and cuticular conductance. K_lamina was thus co‐ordinated with structural and functional traits relating to liquid‐phase water transport and to maximum rates of gas exchange, but independent of other traits relating to drought tolerance and to aspects of carbon economy.     

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

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

Prolonged experimental drought reduces plant hydraulic conductance and transpiration and increases mortality in a piñon–juniper woodland

Plant hydraulic conductance (k_s) is a critical control on whole‐plant water use and carbon uptake and, during drought, influences whether plants survive or die. To assess long‐term physiological and hydraulic responses of mature trees to water availability, we manipulated ecosystem‐scale water availability from 2007 to 2013 in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) woodland. We examined the relationship between k_s and subsequent mortality using more than 5 years of physiological observations, and the subsequent impact of reduced hydraulic function and mortality on total woody canopy transpiration (E_C) and conductance (G_C). For both species, we observed significant reductions in plant transpiration (E) and k_s under experimentally imposed drought. Conversely, supplemental water additions increased E and k_s in both species. Interestingly, both species exhibited similar declines in k_s under the imposed drought conditions, despite their differing stomatal responses and mortality patterns during drought. Reduced whole‐plant k_s also reduced carbon assimilation in both species, as leaf‐level stomatal conductance (g_s) and net photosynthesis (A_n) declined strongly with decreasing k_s. Finally, we observed that chronically low whole‐plant k_s was associated with greater canopy dieback and mortality for both piñon and juniper and that subsequent reductions in woody canopy biomass due to mortality had a significant impact on both daily and annual canopy E_C and G_C. Our data indicate that significant reductions in k_s precede drought‐related tree mortality events in this system, and the consequence is a significant reduction in canopy gas exchange and carbon fixation. Our results suggest that reductions in productivity and woody plant cover in piñon–juniper woodlands can be expected due to reduced plant hydraulic conductance and increased mortality of both piñon pine and juniper under anticipated future conditions of more frequent and persistent regional drought in the southwestern United States.     

Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought

A comparative study on stomatal control under water deficit was conducted on grapevines of the cultivars Grenache, of Mediterranean origin, and Syrah of mesic origin, grown near Montpellier, France and Geisenheim, Germany. Syrah maintained similar maximum stomatal conductance (g_max) and maximum leaf photosynthesis (A_max) values than Grenache at lower predawn leaf water potentials, Ψ_leaf, throughout the season. The Ψ_leaf of Syrah decreased strongly during the day and was lower in stressed than in watered plants, showing anisohydric stomatal behaviour. In contrast, Grenache showed isohydric stomatal behaviour in which Ψ_leaf did not drop significantly below the minimum Ψ_leaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, K_l, incorporating leaf transpiration rate and whole‐plant water potential gradients, previous differences between varieties disappeared both on a seasonal and diurnal scale. This suggested that isohydric and anisohydric behaviour could be regulated by hydraulic conductance. Pressure‐flow measurements on excised organs from plants not previously stressed revealed that Grenache had a two‐ to three‐fold larger hydraulic conductance per unit path length (K_h) and a four‐ to six‐fold larger leaf area specific conductivity (LSC) in leaf petioles than Syrah. Differences between internodes were only apparent for LSC and were much smaller. Cavitation detected as ultrasound acoustic emissions on air‐dried shoots showed higher rates for Grenache than Syrah during the early phases of the dry‐down. It is hypothesized that the differences in water‐conducting capacity of stems and especially petioles may be at the origin of the near‐isohydric and anisohydric behaviour of g.     

Hydraulic characteristics and leaf water use efficiency in trees from tropical montane habitats

This study was carried out in pioneer and successional forest tree species in a lower montane tropical forest with seasonal rains. We tested whether pioneer species feature high hydraulic conductance allowing them to use water profusely at leaf level. Conversely, forest species may have relatively low hydraulic conductance accompanied with better control over water use. This may lead in turn to pioneer species being at a relatively higher risk of shoot water potential falling below the threshold value at which cavitations occur compared to forest. Specific hydraulic conductance ( K _s) measured during the wet season was comparable between pioneers and forest species. During drought, K _s was significantly reduced, and species of both plant groups responded to this by modifying the relationship between conducting area and leaf area (Huver value), such that leaf specific conductivity ( K _l) was unaffected. Thus, leaf area seemed to be adjusted to maintain constant hydraulic sufficiency during drought. Pioneer species were more efficient in conducting water to their leaves but had low control over water use compared to forest species. A trade-off between water transport and leaf water use efficiency was suggested. These ecophysiological differences may have an impact on the performance of the species occupying contrasting habitats. Nonetheless, drought-induced embolisms occurred in trees growing in both open and forest habitats. Overall, during drought, adjustment of leaf area occurred in order to maintain a homeostasis of some physiological traits (leaf-specific conductivity and carbon assimilation).     

Midday stomatal conductance is more related to stem rather than leaf water status in subtropical deciduous and evergreen broadleaf trees

Midday depressions in stomatal conductance (g_s) and photosynthesis are common in plants. The aim of this study was to understand the hydraulic determinants of midday g_s, the coordination between leaf and stem hydraulics and whether regulation of midday g_s differed between deciduous and evergreen broadleaf tree species in a subtropical cloud forest of Southwest (SW) China. We investigated leaf and stem hydraulics, midday leaf and stem water potentials, as well as midday g_s of co‐occurring deciduous and evergreen tree species. Midday g_s was correlated positively with midday stem water potential across both groups of species, but not with midday leaf water potential. Species with higher stem hydraulic conductivity and greater daily reliance on stem hydraulic capacitance were able to maintain higher stem water potential and higher g_s at midday. Deciduous species exhibited significantly higher stem hydraulic conductivity, greater reliance on stem capacitance, higher stem water potential and g_s at midday than evergreen species. Our results suggest that midday g_s is more associated with midday stem than with leaf water status, and that the functional significance of stomatal regulation in these broadleaf tree species is probably for preventing stem xylem dysfunction.     

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.     

Aquaporin regulation in roots controls plant hydraulic conductance,stomatal conductance,and leaf water potential in Pinus radiata under water stress

Stomatal regulation is crucial for forest species performance and survival on drought‐prone sites. We investigated the regulation of root and shoot hydraulics in three Pinus radiata clones exposed to drought stress and its coordination with stomatal conductance (g_s) and leaf water potential (Ψ_leaf). All clones experienced a substantial decrease in root‐specific root hydraulic conductance (K_root‐r) in response to the water stress, but leaf‐specific shoot hydraulic conductance (K_shoot‐l) did not change in any of the clones. The reduction in K_root‐r caused a decrease in leaf‐specific whole‐plant hydraulic conductance (K_plant‐l). Among clones, the larger the decrease in K_plant‐l, the more stomata closed in response to drought. Rewatering resulted in a quick recovery of K_root‐r and g_s. Our results demonstrated that the reduction in K_plant‐l, attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψ_leaf as water stress started. We concluded that higher K_plant‐l is associated with water stress resistance by sustaining a less negative Ψ_leaf and delaying stomatal closure.     

Leaf hydraulic conductance and mesophyll conductance are not closely related within a single species

Stomata represent one resistor in a series of resistances for carbon and water exchange between the leaf and the atmosphere; the remaining resistors occurring within the leaf, commonly represented as mesophyll conductance to CO₂, g_m, and leaf hydraulic conductance, k_Leaf. Recent studies have proposed that g_m and k_Leaf may be coordinated across species because of shared pathways. We assessed the correlation between g_m and k_Leaf within cotton, under growth CO₂ partial pressure and irradiance treatments and also with short‐term variation in irradiance and humidity. g_m was estimated using two isotopic techniques that allowed partitioning of total g_m (Δ¹³C‐g_m) into cell wall plus plasma membrane conductance (Δ¹⁸O‐g_m) and chloroplast membrane conductance (g_cm). A weak correlation was found between Δ¹³C‐g_m and k_Leaf only when measured under growth conditions. However, Δ¹⁸O‐g_m was related to k_Leaf under both short‐term environmental variation and growth conditions. Partitioning g_m showed that g_cm was not affected by short‐term changes in irradiance or correlated with k_Leaf, but was strongly reduced at high growth CO₂ partial pressure. Thus, simultaneous measurements of g_m, k_Leaf and g_cm suggest independent regulation of carbon and water transport across the chloroplast membrane with limited coordinated regulation across the cell wall and plasma membrane.     

Soybean leaf hydraulic conductance does not acclimate to growth at elevated [CO2] or temperature in growth chambers or in the field

Background and Aims

Leaf hydraulic properties are strongly linked with transpiration and photosynthesis in many species. However, it is not known if gas exchange and hydraulics will have co-ordinated responses to climate change. The objective of this study was to investigate the responses of leaf hydraulic conductance (K_leaf) in Glycine max (soybean) to growth at elevated [CO₂] and increased temperature compared with the responses of leaf gas exchange and leaf water status.

Methods

Two controlled-environment growth chamber experiments were conducted with soybean to measure K_leaf, stomatal conductance (g_s) and photosynthesis (A) during growth at elevated [CO₂] and temperature relative to ambient levels. These results were validated with field experiments on soybean grown under free-air elevated [CO₂] (FACE) and canopy warming.

Key results

In chamber studies, K_leaf did not acclimate to growth at elevated [CO₂], even though stomatal conductance decreased and photosynthesis increased. Growth at elevated temperature also did not affect K_leaf, although g_s and A showed significant but inconsistent decreases. The lack of response of K_leaf to growth at increased [CO₂] and temperature in chamber-grown plants was confirmed with field-grown soybean at a FACE facility.

Conclusions

Leaf hydraulic and leaf gas exchange responses to these two climate change factors were not strongly linked in soybean, although g_s responded to [CO₂] and increased temperature as previously reported. This differential behaviour could lead to an imbalance between hydraulic supply and transpiration demand under extreme environmental conditions likely to become more common as global climate continues to change.     

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.     

Disentangling the contributions of ontogeny and water stress to photosynthetic limitations in almond trees

Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (W_a), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (N_w). Concomitantly, area‐based maximum carboxylation rate (V_cmax), maximum electron transport rate (J_max), mesophyll conductance to CO₂ diffusion (gm)′ and light‐saturated net photosynthesis (A_max) declined in both well‐watered and water‐stressed almond leaves. Although g_m and stomatal conductance (g_s) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between g_m and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of A_max was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, A_max was mainly limited by g_s and, to a lesser extent, by photosynthetic capacity and g_m. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.     

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.     

Isometric partitioning of hydraulic conductance between leaves and stems: balancing safety and efficiency in different growth forms and habitats

Recent advances in modelling the architecture and function of the plant hydraulic network have led to improvements in predicting and interpreting the consequences of functional trait variation on CO₂ uptake and water loss. We build upon one such model to make novel predictions for scaling of the total specific hydraulic conductance of leaves and shoots (k_L and k_SH, respectively) and variation in the partitioning of hydraulic conductance. Consistent with theory, we observed isometric (slope = 1) scaling between k_L and k_SH across several independently collected datasets and a lower ratio of k_L and k_SH, termed the leaf‐to‐shoot conductance ratio (C_LSCR), in arid environments and in woody species. Isometric scaling of k_L and k_SH supports the concept that hydraulic design is coordinated across the plant. We propose that C_LSCR is an important adaptive trait that represents the trade‐off between efficiency and safety at the scale of the whole plant.     

Stem water storage capacity and efficiency of water transport: their functional significance in a Hawaiian dry forest

We investigated the contribution of internal water storage and efficiency of water transport to the maintenance of water balance in six evergreen tree species in a Hawaiian dry forest. Wood‐saturated water content, a surrogate for relative water storage capacity, ranged from 70 to 105%, and was inversely related to its morphological correlate, wood density, which ranged between 0·51 and 0·65 g cm^?3. Leaf‐specific conductivity (k_L) measured in stem segments from terminal branches ranged from 3 to 18 mmol m^?1 s^?1 MPa^?1, and whole‐plant hydraulic efficiency calculated as stomatal conductance (g) divided by the difference between predawn and midday leaf water potential (Ψ_L), ranged from 70 to 150 mmol m^?2 s^?1 MPa^?1. Hydraulic efficiency was positively correlated with k_L (r² = 0·86). Minimum annual Ψ_L ranged from ? 1·5 to ? 4·1 MPa among the six species. Seasonal and diurnal variation in Ψ_L were associated with differences among species in wood‐saturated water content, wood density and k_L. The species with higher wood‐saturated water content were more efficient in terms of long‐distance water transport, exhibited smaller diurnal variation in Ψ_L and higher maximum photosynthetic rates. Smaller diurnal variation in Ψ_L in species with higher wood‐saturated water content, k_L and hydraulic efficiency was not associated with stomatal restriction of transpiration when soil water deficit was moderate, but avoidance of low minimum seasonal Ψ_L in these species was associated with a substantial seasonal decline in g. Low seasonal minimum Ψ_L in species with low k_L, hydraulic efficiency, and wood‐saturated water content was associated with higher leaf solute content and corresponding lower leaf turgor loss point. Despite the species‐specific differences in leaf water relations characteristics, all six evergreen tree species shared a common functional relationship defined primarily by k_L and stem water storage capacity.     

Different responses to drought and freeze stress of Quercus ilex L. growing along a latitudinal gradient

The vulnerability to drought and freeze stress was measured in young plants of Quercus ilex L. growing in the field in two natural sites within the Italian distribution area of this species, i.e. Sicily (Southern Mediterranean Basin) and Venezia Giulia (Northeastern Italy), respectively. In particular, the resistance strategies adopted by Q. ilex to withstand the two stresses were estimated in terms of seasonal and/or diurnal changes in leaf conductance to water vapour (g_L), water potential (_L) and relative water content (RWC) as well as of xylem embolism in the stem and root hydraulic conductance (K_RL). Sicilian (SI) plants showed to reduce water loss by stomatal closure (g_L decreased) in summer, thus maintaining average RWCs at 88–90%. Moreover, SI plants showed considerable resistance to xylem cavitation in the stem (the loss of hydraulic conductance, PLC, was less than 12% throughout the year) and to maintain the hydraulic conductance of their roots (K_RL), constantly high even in summer. Plants growing in Venezia Giulia (VG plants), on the contrary, underwent leaf dehydration in the winter due to freeze stress so that RWC measured in April was still 78% on a diurnal basis. This was apparently due to consistent xylem embolism in the stem. In fact, PLC was as high as 40% between November and March. Only in the summer was PLC similar to that recorded in SI plants. Moreover, K_RL of VG plants decreased in November from about 1.5 to 0.8×10^–4 kg s^–1 m^–2 MPa^–1, i.e. about 50%, and in February K_RL dropped further to 0.4×10^–4 kg s^–1 m^–2 MPa^–1. On the basis of the above, we conclude that: (a) Q. ilex was more sensitive to freeze than to drought stress so that freeze stress can be considered as a factor limiting the distribution area of this species; (b) drought and freeze stress were faced by Q. ilex adopting two different resistance strategies, i.e. drought avoidance based on water saving in Sicily and freeze tolerance in Venezia Giulia.     

Are Sclerophylls and Malacophylls Hydraulically Different?

This work tests the hypothesis that sclerophylls (i.e. hard-leaved species) would be less efficient than malacophylls (i.e. soft-leaved species) in terms of water transport through the stem as well as within the leaf blade. Mean leaf surface area (A_L), leaf specific mass (LSM) as well as shoot (K_WL), stem (K_SL) and leaf (K_LL) hydraulic conductances were measured in eight Mediterranean evergreen sclerophylls and eight temperate deciduous malacophylls. No difference was observed between the two groups in terms of K_LL and of the contribution of leaves to the overall shoot hydraulic resistance. Leaves represented in all cases 48 to 90 % of the shoot hydraulic resistance, suggesting that the sclerophyllous habitus does not per se lead to low efficiency in water transport within the leaf blade. A weak negative relationship (r² = 0.252) appeared to exist between K_SL and LSM. This might provide an explanation for the lower growth rates of sclerophylls with respect to malacophylls.     

Neither xylem collapse,cavitation, or changing leaf conductance drive stomatal closure in wheat

Identifying the drivers of stomatal closure and leaf damage during stress in grasses is a critical prerequisite for understanding crop resilience. Here, we investigated whether changes in stomatal conductance (g_s) during dehydration were associated with changes in leaf hydraulic conductance (K_leaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of g_s was concomitant with declining K_leaf under mild water stress. This early decline of leaf hydraulic conductance was not driven by cavitation, as the first cavitation events in leaf and stem were detected well after K_leaf had declined. Xylem vessel deformation could only account for <5% of the observed decline in leaf hydraulic conductance during dehydration. Thus, we concluded that changes in the hydraulic conductance of tissues outside the xylem were responsible for the majority of K_leaf decline during leaf dehydration in wheat. However, the contribution of leaf resistance to whole plant resistance was less than other tissues (<35% of whole plant resistance), and this proportion remained constant as plants dehydrated, indicating that K_leaf decline during water stress was not a major driver of stomatal closure.