Relationships between stomatal behavior,xylem vulnerability to cavitation and leaf water relations in two cultivars of Vitis vinifera

Current understanding of physiological mechanisms governing stomatal behavior under water stress conditions is still incomplete and controversial. It has been proposed that coordination of stomatal kinetics with xylem vulnerability to cavitation [vulnerability curve (VC)] leads to different levels of isohydry/anisohydry in different plant species/cultivars. In this study, this hypothesis is tested in Vitis vinifera cultivars displaying contrasting stomatal behavior under drought stress. The cv Montepulciano (MP, near‐isohydric) and Sangiovese (SG, anisohydric) were compared in terms of stomatal response to leaf and stem water potential, as possibly correlated to different petiole hydraulic conductivity (k_petiole) and VC, as well as to leaf water relations parameters. MP leaves showed almost complete stomatal closure at higher leaf and stem water potentials than SG leaves. Moreover, MP petioles had higher maximum k_petiole and were more vulnerable to cavitation than SG. Water potential at the turgor loss point was higher in MP than in SG. In SG, the percentage reduction of stomatal conductance (PLg_s) under water stress was almost linearly correlated with corresponding percentage loss of k_petiole (PLC), while in MP PLg_s was less influenced by PLC. Our results suggest that V. vinifera near‐isohydric and anisohydric genotypes differ in terms of xylem vulnerability to cavitation as well as in terms of k_petiole, and that the coordination of these traits leads to their different stomatal responses under water stress conditions.     

Hydraulic properties of rice and the response of gas exchange to water stress

We investigated the role of xylem cavitation, plant hydraulic conductance, and root pressure in the response of rice (Oryza sativa) gas exchange to water stress. In the field (Philippines), the percentage loss of xylem conductivity (PLC) from cavitation exceeded 60% in leaves even in watered controls. The PLC versus leaf water potential relationship indicated diurnal refilling of cavitated xylem. The leaf water potential causing 50 PLC (P(50)) was -1.6 MPa and did not differ between upland versus lowland rice varieties. Greenhouse-grown varieties (Utah) were more resistant to cavitation with a 50 PLC of -1.9 MPa but also showed no difference between varieties. Six-day droughts caused concomitant reductions in leaf-specific photosynthetic rate, leaf diffusive conductance, and soil-leaf hydraulic conductance that were associated with cavitation-inducing water potentials and the disappearance of nightly root pressure. The return of root pressure after drought was associated with the complete recovery of leaf diffusive conductance, leaf-specific photosynthetic rate, and soil-leaf hydraulic conductance. Root pressure after the 6-d drought (61.2 +/- 8.8 kPa) was stimulated 7-fold compared with well-watered plants before drought (8.5 +/- 3.8 kPa). The results indicate: (a) that xylem cavitation plays a major role in the reduction of plant hydraulic conductance during drought, and (b) that rice can readily reverse cavitation, possibly aided by nocturnal root pressure.     

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

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

Minimum hydraulic safety leads to maximum water-use efficiency in a forage grass

Understanding how water-use regulation relates to biomass accumulation is imperative for improving crop production in water-limited environments. Here, we examine how the vulnerability of xylem to water stress-induced cavitation and the coordination between water transport capacity and assimilation (A) influences diurnal water-use efficiency (WUE) and dry-matter production in Lolium perenne L. - a commercial forage grass. Plants were exposed to a range of water stresses, causing up to 90% leaf death, by withholding water and then rewatering to observe the recovery process. Leaf hydraulic conductance (K(leaf) ) declined to 50% of maximum at a leaf water potential (ψ(leaf) ) of -1 MPa, whereas complete stomatal closure occurred well after this point, at -2.35 MPa, providing no protection against hydraulic dysfunction. Instantaneous A remained maximal until >70% of hydraulic conductivity had been lost. Post-stress rewatering showed that 95% loss of K(leaf) could be incurred before the recovery of gas exchange exceeded 1 d, with a rapid transition to leaf death after this point. Plants exposed to sustained soil water deficits through restricted nightly watering regimes did not suffer cumulative losses in K(leaf) ; instead, ψ(leaf) and gas exchange recovered diurnally. The effect was improved WUE during the day and optimal ψ(leaf) during the night for the maintenance of growth.     

Drought-Induced Xylem Dysfunction in Petioles,Branches, and Roots of Populus balsamifera L. and Alnus glutinosa (L.) Gaertn

Variation in vulnerability to xylem cavitation was measured within individual organs of Populus balsamifera L. and Alnus glutinosa (L.) Gaertn. Cavitation was quantified by three different techniques: (a) measuring acoustic emissions, (b) measuring loss of hydraulic conductance while air-dehydrating a branch, and (c) measuring loss of hydraulic conductance as a function of positive air pressure injected into the xylem. All of these techniques gave similar results. In Populus, petioles were more resistant than branches, and branches were more resistant than roots. This corresponded to the pattern of vessel width: maximum vessel diameter in 1- to 2-year-old roots was 140 [mu]m, compared to 65 and 45 [mu]m in rapidly growing 1-year-old shoots and petioles, respectively. Cavitation in Populus petioles started at a threshold water potential of -1.1 MPa. The lowest leaf water potential observed was -0.9 MPa. In Alnus, there was no relationship between vessel diameter and the cavitation response of a plant organ. Although conduits were narrower in petioles than in branches, petioles were more vulnerable to cavitation. Cavitation in petioles was detected when water potential fell below -1.2 MPa. This value equaled midday leaf water potential in late June. As in Populus, roots were the most vulnerable organ. The significance of different cavitation thresholds in individual plant organs is discussed.     

Xylem cavitation in the leaf of Prunus laurocerasus and its impact on leaf hydraulics

This paper reports how water stress correlates with changes in hydraulic conductivity of stems, leaf midrib, and whole leaves of Prunus laurocerasus. Water stress caused cavitation-induced dysfunction in vessels of P. laurocerasus. Cavitation was detected acoustically by counts of ultrasonic acoustic emissions and by the loss of hydraulic conductivity measured by a vacuum chamber method. Stems and midribs were approximately equally vulnerable to cavitations. Although midribs suffered a 70% loss of hydraulic conductance at leaf water potentials of -1.5 MPa, there was less than a 10% loss of hydraulic conductance in whole leaves. Cutting and sealing the midrib 20 mm from the leaf base caused only a 30% loss of conduction of the whole leaf. A high-pressure flow meter was used to measure conductance of whole leaves and as the leaf was progressively cut back from tip to base. These data were fitted to a model of hydraulic conductance of leaves that explained the above results, i.e. redundancy in hydraulic pathways whereby water can flow around embolized regions in the leaf, makes whole leaves relatively insensitive to significant changes in conductance of the midrib. The onset of cavitation events in P. laurocerasus leaves correlated with the onset of stomatal closure as found recently in studies of other species in our laboratory.     

Hydraulic vulnerability, vessel refilling, and seasonal courses of stem water potential of Sorbus aucuparia L. and Sambucus nigra L.

Differences in the seasonal variation in stem water potential between the two shrub species Sorbus aucuparia and Sambucus nigra were related with their vulnerability to xylem cavitation. It was also demonstrated indirectly that the two species differ in the extent to which they reverse cavitation. Seasonal variation in stem water potential was investigated during three growing seasons with in situ stem psychrometers. Sorbus experienced wide water potential variations and reached a minimum of -4.2 MPa during drought. Under the same microclimatic conditions, Sambucus experienced consistent stem water potentials with a minimum of -1.7 MPa. The relationship between percentage loss in hydraulic conductivity (PLC) and water potential (hydraulic vulnerability curve) of the two species differed in shape: a flat curve with nearly total loss of conductivity at -6 MPa was found for SORBUS: Sambucus showed a steep vulnerability curve with 90% loss conductivity at -2.2 MPa. Thus, Sambucus is extremely vulnerable to cavitation, but Sorbus is an almost invulnerable species. This different cavitation resistance adjusted the ranges of field stem water potential that the species experienced. Finally, seasonal courses of naturally occurring (native) embolism were compared with calculated PLC courses. This comparison indicates that Sorbus did not refill embolized xylem vessels whereas Sambucus reversed embolism. It was concluded that species which are highly vulnerable to cavitation and drought-induced embolism need refilling of embolized vessels as well as isohydric water potential patterns as two strategies of survival.     

Vein recovery from embolism occurs under negative pressure in leaves of sunflower (Helianthus annuus)

Leaf veins undergo cavitation at water potentials (Psi(leaf)) commonly experienced by field-growing plants. Theoretically, embolism reversal should not be possible until xylem pressures rise by several kilopascals of atmospheric pressure, but recent evidence suggests that embolized conduits can be refilled even when surrounded by others at substantial tension (novel refilling). The present study reports 'novel refilling' occurring in leaf veins of sunflower (Helianthus annuus L.) while at Psi(leaf) = -0.33 MPa. Sixty per cent loss of vein hydraulic conductance (K(vein)) was recorded at Psi(leaf) < -0.65 MPa, while stem hydraulic conductance (K(stem)) was unaffected even at Psi(leaf) = -1.1 MPa. Loss of K(vein) was accompanied by stomatal closure. Water-stressed plants (Psi(leaf) = -1.1 MPa) were rehydrated overnight to different target water potentials achieved by using PEG at different concentrations as irrigation medium. K(vein) recovered by 50% at Psi(leaf) = -0.47 MPa and vein refilling was complete at Psi(leaf) = -0.33 MPa, i.e. well below the theoretical limit for conduit refilling (-0.05 MPa as calculated for sunflower minor veins). Mercurials supplied to detached leaves had no effect on the refilling process. Upon rehydration, recovery of K(vein) was not paralleled by recovery of whole-plant hydraulic conductance or leaf conductance to water vapour (g(L)), as a likely consequence of hydraulic failure of other components of the water pathway (root system or extravascular leaf compartments) and/or root-to-leaf chemical signalling. This is the first study providing experimental evidence for 'novel refilling' in a herbaceous dicot and highlighting the importance of this process in the leaf.     

Water loss regulation to soil drought associated with xylem vulnerability to cavitation in temperate ring-porous and diffuse-porous tree seedlings

Key message

Sustainable stomatal opening despite xylem cavitation occurs in ring-porous species and stomatal closure prior to cavitation in diffuse-porous species during soil drought.

Abstract

To elucidate the relationship between water loss regulation and vulnerability to cavitation associated with xylem structure, stomatal conductance (g _s), defoliation, vulnerability curves, and vessel features were measured on seedlings of ring-porous Zelkova serrata and Melia azedarach, and diffuse-porous Betula platyphylla var. japonica, Cerasus jamasakura and Carpinus tschonoskii. Under prolonged drought conditions, the percentage loss of hydraulic conductivity (PLC) increased and g _s decreased gradually with decreasing predawn (Ψ_pd) or xylem water potential (Ψ_xylem) in Z. serrata. During the gentle increase of PLC in M. azedarach, g _s increased in the early stages of dehydration while leaves were partly shed. A sharp reduction in g _s was observed before the onset of an increase in the PLC for drying plants of the three diffuse-porous species, suggesting cavitation avoidance by stomatal regulation. In the ring-porous species, xylem-specific hydraulic conductivity (K _s) was higher, whereas the vessel multiple fractions, the ratio of the number of grouped vessels to total vessels, was lower than that in the diffuse-porous species, suggesting that many were distributed as solitary vessels. This may explain the gradual increase in the PLC with decreasing Ψ_xylem because isolated vessels provide less opportunity for air seeding. Different water loss regulation to soil drought was identified among the species, with potential mechanisms being sustainable gas exchange at the expense of xylem dysfunction or partial leaf shedding, and the avoidance of xylem cavitation by strict stomatal regulation. These were linked to vulnerability to cavitation that appears to be governed by xylem structural properties.     

Long-term acclimatization of hydraulic properties, xylem conduit size, wall strength and cavitation resistance in Phaseolus vulgaris in response to different environmental effects

Phaseolus vulgaris grown under various environmental conditions was used to assess long-term acclimatization of xylem structural characteristics and hydraulic properties. Conduit diameter tended to be reduced and 'wood' density (of 'woody' stems) increased under low moisture ('dry'), increased soil porosity ('porous soil') and low phosphorus ('low P') treatments. Dry and low P had the largest percentage of small vessels. Dry, low light ('shade') and porous soil treatments decreased P50 (50% loss in conductivity) by 0.15-0.25 MPa (greater cavitation resistance) compared with 'controls'. By contrast, low P increased P50 by 0.30 MPa (less cavitation resistance) compared with porous soil (the control for low P). Changes in cavitation resistance were independent of conduit diameter. By contrast, changes in cavitation resistance were correlated with wood density for the control, dry and porous soil treatments, but did not appear to be a function of wood density for the shade and low P treatments. In a separate experiment comparing control and porous soil plants, stem hydraulic conductivity (kh), specific conductivity (ks), leaf specific conductivity (LSC), total pot water loss, plant biomass and leaf area were all greater for control plants compared to porous soil plants. Porous soil plants, however, demonstrated higher midday stomatal conductance to water vapour (gs), apparently because they experienced proportionally less midday xylem cavitation.     

Variation in embolism occurrence and repair along the stem in drought‐stressed and re‐watered seedlings of a poplar clone

Root pressure and plasma membrane intrinsic protein (PIP) availability in the xylem have been recognized to participate in the refilling of embolized conduits, yet integration of the two mechanisms has not been reported in the same plant. In this study, 4‐month‐old seedlings of a hybrid poplar (Populus alba × Populus glandulosa) clone 84K were subjected to two contrasting soil‐water treatments, with the drought treatment involving withholding of water for 17 days to reduce the soil‐water content to 10% of the saturated field capacity, followed by a re‐watering cycle. The percentage loss of stem hydraulic conductance (PLC) sharply increased, and stomatal conductance and photosynthesis declined in response to drought stress; these processes were gradually restored following the subsequent re‐watering. Embolism was most severe in the middle portions of the stem, followed by the basal and top portions of the stems of seedlings subjected to drought stress and subsequent re‐watering. Although drought stress eliminated root pressure, re‐watering partially restored it in a short period of time. The expression of PIP genes in the xylem was activated by drought stress, and some PIP genes were further stimulated in the top portion after re‐watering. The dynamics of root pressure and differential expression of PIP genes along the stem coincided with changes in PLC, suggesting that root pressure and PIPs work together to refill the embolized vessels. On the basis of the recovery dynamics in PLC and g_smax (maximum stomatal conductance) after re‐watering, the stomatal closure and xylem cavitation exhibited fatigue due to drought stress.     

Cryo-scanning electron microscopy observations of vessel content during transpiration in walnut petioles. Facts or artifacts?

The current controversy about the "cohesion-tension" of water ascent in plants arises from the recent cryo-scanning electron microscopy (cryo-SEM) observations of xylem vessels content by Canny and coworkers (1995). On the basis of these observations it has been claimed that vessels were emptying and refilling during active transpiration in direct contradiction to the previous theory. In this study we compared the cryo-SEM data with the standard hydraulic approach on walnut (Juglans regia) petioles. The results of the two techniques were in clear conflict and could not both be right. Cryo-SEM observations of walnut petioles frozen intact on the tree in a bath of liquid nitrogen (LN(2)) suggested that vessel cavitation was occurring and reversing itself on a diurnal basis. Up to 30% of the vessels were embolized at midday. In contrast, the percentage of loss of hydraulic conductance (PLC) of excised petiole segments remained close to 0% throughout the day. To find out which technique was erroneous we first analyzed the possibility that PLC values were rapidly returned to zero when the xylem pressures were released. We used the centrifugal force to measure the xylem conductance of petiole segments exposed to very negative pressures and established the relevance of this technique. We then analyzed the possibility that vessels were becoming partially air-filled when exposed to LN(2). Cryo-SEM observations of petiole segments frozen shortly after their xylem pressure was returned to atmospheric values agreed entirely with the PLC values. We confirmed, with water-filled capillary tubes exposed to a large centrifugal force, that it was not possible to freeze intact their content with LN(2). We concluded that partially air-filled conduits were artifacts of the cryo-SEM technique in our study. We believe that the cryo-SEM observations published recently should probably be reconsidered in the light of our results before they may be used as arguments against the cohesion-tension theory.     

Interactive effects of water stress and xylem-limited bacterial infection on the water relations of a host vine

Xylella fastidiosa, a xylem-limited bacterial pathogen that causes bacterial leaf scorch in its hosts, has a diverse and extensive host range among plant species worldwide. Previous work has shown that water stress enhances leaf scorch symptom severity and progression along the stem in Parthenocissus quinquefolia infected by X. fastidiosa. The objective here was to investigate the mechanisms underlying the interaction of water stress and infection by X. fastidiosa. Using the eastern deciduous forest vine, P. quinquefolia, infection and water availability were manipulated while measuring leaf water potentials (psi(L)), stomatal conductance (g(s)), whole shoot hydraulic conductance (K(h)), per cent xylem embolism, and xylem vessel dimensions. No significant differences in any of the physiological measurements were found between control and infected plants prior to drought. Drought treatment significantly reduced psi(L) and g(s) at all leaf positions throughout the day in late summer in both years of the study. In addition, infection significantly reduced psi(L) and g(s) in the most basal leaf positions in late summer in both years. Whole shoot hydraulic conductance was reduced by both low water and infection treatments. However, per cent embolized vessels and mean vessel diameter were affected by drought treatment only. These results imply that the major effect of infection by X. fastidiosa occurs due to reduced hydraulic conductance caused by clogging of the vessels, and not increased cavitation and embolism of xylem elements. The reduced K(h) caused by X. fastidiosa infection acts additively with the water limitation imposed by Drought stress.     

Ion-mediated increase in the hydraulic conductivity of Laurel stems: role of pits and consequences for the impact of cavitation on water transport

Changes in hydraulic conductivity (K(h)) were measured in stems of Laurus nobilis L. during perfusion with KCl, NaCl or sucrose solutions. Ionic solutes induced marked increase of K(h) with respect to deionized water but sucrose had no effect. The kinetics of KCl-induced K(h) increase was measured together with changes in [K(+)] of the perfused solution. K(h) increases were paralleled by increases in the [K(+)](out)/[K(+)](in) ratio. Samples of different lengths or with increasing percentage loss of conductivity (PLC) due to xylem cavitation were tested, with the aim of increasing radial flow through intervessel pits. KCl solutions enhanced the K(h) of 12-cm-long samples with a concentration-dependent effect up to 100 mm KCl. DeltaK(h) increased from 3 to 30% in 1.5- and 12-cm-long samples, respectively and remained constant for longer samples. Increasing PLC induced an exponential increase in DeltaK(h). PLC measured with KCl solutions was significantly less than that measured with deionized water, suggesting that measurements of PLC can be affected by the composition of the perfused solution. Experiments support the hypothesis that the 'ionic effect' is mediated by physico-chemical changes of pectins of the pit membranes and raise the possibility that plants might alter the ionic composition of the xylem sap to alleviate the hydraulic impact of cavitation.     

Sensitivity of stem and petiole hydraulic conductance of deciduous trees to xylem sap ion concentration

Hydraulic conductance of stem and petioles increased in response to an increase in xylem sap ion concentration, and decreased in response to a decrease in the ion concentration in six temperate deciduous tree species. The ion sensitivity of hydraulic conductance of stem and petioles was higher than the ion sensitivity of the stem alone. The ion sensitivity was lowest in the earliest developmental stages of the xylem, and had a seasonal maximum in the second half of summer. The ion sensitivity was highest in slow-growing species and lowest in fast-growing species. The ion sensitivity correlated negatively with mean radius of xylem conduits, hydraulic conductance of stem and petioles, hydraulic conductance of leaf laminae, and stomatal conductance, and positively with response of the hydraulic conductance of leaf laminae to HgCl₂, and stomatal response to a decrease in leaf water potential or abscisic acid. It was concluded that the high ion sensitivity of xylem hydraulic conductance is a relevant characteristic of slow growth and a conservative water use strategy.     

Functional coordination between leaf gas exchange and vulnerability to xylem cavitation in temperate forest trees

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both traditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost (psi50) was positively associated (P < 0.001) with specific conductivity (K(S)) and with mean hydraulically weighted xylem conduit diameter (D(h-w)), but was only marginally (P = 0.06) associated with leaf specific conductivity (K(L)). The PIC correlation for each of these relationships, however, was not statistically significant. There was also no relationship between root psi50 and root K(S) in either cross-species or PIC analysis. Photosynthetic rate (A) and stomatal conductance (g(s)) were strongly and positively correlated with root psi50 in the cross-species analysis (P < 0.001), a relationship that was robust to phylogenetic correction (P < 0.01). A and g(s) were also positively correlated with stem psi50 in the cross-species analysis (P = 0.02 and 0.10, respectively). However, only A was associated with stem psi50 in the PIC analysis (P = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g(s) and root psi50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conductance, and highlight the potential physiological and evolutionary significance of root hydraulic properties in controlling interspecific variation in leaf function.     

Stomatal closure during leaf dehydration,correlation with other leaf physiological traits

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.     

Xylem embolism and stomatal regulation in two rubber clones (<Emphasis Type="Italic">Hevea brasiliensis</Emphasis> Muell. Arg.)

Vulnerability to water-stress-induced embolism of stems, petioles, and leaf midribs was evaluated for two rubber clones (RRIM600 and RRIT251). The xylem conduits were relatively vulnerable to cavitation with 50% of embolism measured for xylem pressures between –1 and –2 MPa. This feature can be related to the tropical-humid origin of the species. A distinct basipetal gradient of vulnerability was found, leaf midribs being the least vulnerable. Substantial variation in vulnerability to cavitation was found between the two clones only at the petiole level. A correlation was found between the stomatal behavior and the development of cavitation. Stomata were nearly closed when the xylem pressure reached the point of xylem dysfunction. Stomata may thus contribute to controlling the risk of cavitation. However, for one clone a poor correlation was found between stomatal regulation and petiole vulnerability. This was consistent with a high degree of embolism measured in the petioles after a soil drought event. Therefore, xylem cavitation might represent a promising criterion to evaluate the performance of rubber clones under drought conditions.     

Gas exchange and hydraulic properties in the crowns of two tree species in a Panamanian moist forest

Hydraulic properties and gas exchange were measured in branches of two tropical tree species (Simarouba amara Aubl. and Tapirira guianensis Aubl.) in a moist lowland forest in Panama. Branch-level sapflow, leaf-level stomatal conductance, and water potential measurements, along with measurements of specific hydraulic conductivity of stems in crown tops, were used to relate hydraulic parameters to leaf conductance in two individuals of each species. Branches of the taller trees for each species (28 m, 31 m) showed much higher leaf-specific hydraulic conductance and leaf vapor-phase conductance than those of the smaller trees (18m, 23m). This was probably related to the leaf-to-sapwood area ratio in branches of taller trees, which was less than half that in branches of smaller trees. Dye staining showed evidence of massive cavitation in all trees, indicating that stomata do not control leaf water potential to prevent xylem cavitation in these species. Stomatal conductance of intact leaves also appeared to be insensitive to leaf area removal treatment of nearby foliage. Nevertheless, a simple mass-balance model of water flux combining hydraulic and vapor transport was in close agreement with observed maximal vapor-phase conductance in the four trees (r²=0.98, P=0.006). Our results suggest that the major organismal control over water flux in these species is by structural (leaf area) rather than physiological (stomatal) means.     

Stomatal factors and vulnerability of stem xylem to cavitation in poplars

The relationships between the vulnerability of stem xylem to cavitation, stomatal conductance, stomatal density, and leaf and stem water potential were examined in six hybrid poplar (P38P38, Walker, Okanese, Northwest, Assiniboine and Berlin) and balsam poplar (Populus balsamifera) clones. Stem xylem cavitation resistance was examined with the Cavitron technique in well-watered plants grown in the greenhouse. To investigate stomatal responses to drought, plants were subjected to drought stress by withholding watering for 5 (mild drought) and 7 (severe drought) days and to stress recovery by rewatering severely stressed plants for 30 min and 2 days. The clones varied in stomatal sensitivity to drought and vulnerability to stem xylem cavitation. P38P38 reduced stomatal conductance in response to mild stress while the balsam poplar clone maintained high leaf stomatal conductance under more severe drought stress conditions. Differences between the severely stressed clones were also observed in leaf water potentials with no or relatively small decreases in Assiniboine, P38P38, Okanese and Walker. Vulnerability to drought-induced stem xylem embolism revealed that balsam poplar and Northwest clones reached loss of conductivity at lower stem water potentials compared with the remaining clones. There was a strong link between stem xylem resistance to cavitation and stomatal responsiveness to drought stress in balsam poplar and P38P38. However, the differences in stomatal responsiveness to mild drought suggest that other drought-resistant strategies may also play a key role in some clones of poplars exposed to drought stress.