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1.
The impact of xylem cavitation and embolism on leaf (K leaf) and stem (K stem) hydraulic conductance was measured in current-year shoots of Cercis siliquastrum L. (Judas tree) using the vacuum chamber technique. K stem decreased at leaf water potentials (ΨL) lower than ?1.0 MPa, while K leaf started to decrease only at ΨL L K leaf changes. Field measurements of leaf conductance to water vapour (g L) and ΨL showed that stomata closed when ΨL decreased below the ΨL threshold inducing loss of hydraulic conductance in the leaf. The partitioning of hydraulic resistances within shoots and leaves was measured using the high-pressure flow meter method. The ratio of leaf to shoot hydraulic resistance was about 0.8, suggesting that stem cavitation had a limited impact on whole shoot hydraulic conductance. We suggest that stomatal aperture may be regulated by the cavitation-induced reduction of hydraulic conductance of the soil-to-leaf water pathway which, in turn, strongly depends on the hydraulic architecture of the plant and, in particular, on leaf hydraulics.  相似文献   

2.
The possible link between stomatal conductance (gL), leaf water potential ( Ψ L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of gL in whole plants was the consequence of Ψ L reaching the cavitation threshold ( Ψ CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in gL of about 50%, while Ψ L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between gL and Ψ CAV rather than with Ψ L itself.  相似文献   

3.
Vulnerability to cavitation of leaf minor veins and stems of Laurus nobilis L. was quantified together with that of leaflets, rachides and stems of Ceratonia siliqua L. during air‐dehydration of 3‐year‐old branches. Embolism was estimated by counting ultrasound acoustic emissions (UAE) and relating them to leaf water potential (ΨL). The threshold ΨL for cavitation was less negative in L. nobilis than in C. siliqua according to the known higher drought resistance of the latter species. Leaf minor vein cavitation was also quantified by infiltrating leaves with fluorescein at different dehydration levels and observing them under microscope. Distinct decreases in the functional integrity of minor veins were observed during leaf dehydration, with high correlation between the two variables. The relationship between leaf conductance to water vapour (gL) and ΨL showed that stomata of L. nobilis closed in response to stem and not to leaf cavitation. However, in C. siliqua, gL decreased in coincidence to the leaf cavitation threshold, which was, nevertheless, very close to that of the stem. The hypothesis that stem cavitation acts as a signal for stomatal closure was confirmed, while the same role for leaf cavitation remains an open problem.  相似文献   

4.
This study examined the linkage between xylem vulnerability, stomatal response to leaf water potential (ΨL), and loss of leaf turgor in eight species of seasonally dry tropical forest trees. In order to maximize the potential variation in these traits species that exhibit a range of leaf habits and phenologies were selected. It was found that in all species stomatal conductance was responsive to ΨL over a narrow range of water potentials, and that ΨL inducing 50% stomatal closure was correlated with both the ΨL inducing a 20% loss of xylem hydraulic conductivity and leaf water potential at turgor loss in all species. In contrast, there was no correlation between the water potential causing a 50% loss of conductivity in the stem xylem, and the water potential at stomatal closure (ΨSC) amongst species. It was concluded that although both leaf and xylem characters are correlated with the response of stomata to ΨL, there is considerable flexibility in this linkage. The range of responses is discussed in terms of the differing leaf‐loss strategies exhibited by these species.  相似文献   

5.
The leaf hydraulic conductance (KL) was measured in Prunus laurocerasus L. and Juglans regia L. in which previous measurements had revealed different impacts of dehydration on KL. Leaves of P. laurocerasus lost 8% of their KL at water potentials (ΨL) of ?2.0 MPa. Leaflets of J. regia showed KL losses of 40% at ΨL = ?1.0 MPa. When major veins were blocked using cyanoacrylate to simulate their embolism, the KL of P. laurocerasus was reduced by 57% but that of J. regia leaflets was reduced by 80%. Such differences were hypothesized to be due to different axial‐to‐radial permeabilites of major veins. Infiltration of leaves with Phoxine B revealed that P. laurocerasus major veins were largely leaky in the radial direction whereas those of J. regia leaflets showed prevailing axial water transport. Differences between species in terms of axial‐to‐radial water permeability were confirmed by measurements of changes of hydraulic resistance along the midrib. The two hydraulic models are discussed in terms of leaf vulnerability to embolism and plant adaptation to dry habitats.  相似文献   

6.
Hydraulic architecture was studied in shrub species differing in rooting depth in a cold desert in Southern Argentina. All species exhibited strong hydraulic segmentation between leaves, stems and roots with leaves being the most vulnerable part of the hydraulic pathway. Two types of safety margins describing the degree of conservation of the hydraulic integrity were used: the difference between minimum stem or leaf water potential (Ψ) and the Ψ at which stem or leaf hydraulic function was reduced by 50% (ΨΨ50), and the difference between leaf and stem Ψ50. Leaf Ψ50 – stem Ψ50 increased with decreasing rooting depth. Large diurnal decreases in root‐specific hydraulic conductivity suggested high root vulnerability to embolism across all species. Although stem Ψ50 became more negative with decreasing species‐specific Ψsoil and minimum stem Ψ, leaf Ψ50 was independent of Ψ and minimum leaf Ψ. Species with embolism‐resistant stems also had higher maximum stem hydraulic conductivity. Safety margins for stems were >2.1 MPa, whereas those for leaves were negative or only slightly positive. Leaves acted as safety valves to protect the integrity of the upstream hydraulic pathway, whereas embolism in lateral roots may help to decouple portions of the plant from the impact of drier soil layers.  相似文献   

7.
Hydraulic conductance was measured on leaf and stem segments excised from sugarcane plants at different stages of development. Maximum transpiration rates and leaf water potential (ΨL) associated with maximum transpiration were also measured in intact plants as a function of plant size. Leaf specific hydraulic conductivity (Lsc) and transpiration on a unit leaf area basis (E) were maximal in plants with approximately 0.2 m2 leaf area and decreased with increasing plant size. These changes in Fand Lsc were nearly parallel, which prevented φL in larger plants from decreasing to levels associated with substantial loss in xylem conductivity caused by embolism formation. Coordination of changes in E and leaf hydraulic properties was not mediated by declining leaf water status, since φL increased with plant size. Hydraulic constrictions were present at nodes and in the node-leaf sheath-leaf blade pathway. This pattern of constrictions is in accord with the idea of plant segmentation into regions differing in water transport efficiency and would tend to confine embolisms to the relatively expendable leaves at terminal positions in the pathway, thereby preserving water transport through the stem.  相似文献   

8.
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 (gs) during dehydration were associated with changes in leaf hydraulic conductance (Kleaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of gs was concomitant with declining Kleaf 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 Kleaf 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 Kleaf 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 Kleaf decline during water stress was not a major driver of stomatal closure.  相似文献   

9.
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 (gmax) and maximum leaf photosynthesis (Amax) values than Grenache at lower predawn leaf water potentials, Ψleaf, throughout the season. The Ψleaf 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 Ψleaf did not drop significantly below the minimum Ψleaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, Kl, 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 (Kh) 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.  相似文献   

10.
Pathogens can cause chronic premature needle abscission in coniferous species. To assess the potential impacts on tree productivity, stomatal regulation was investigated in Douglas fir with chronic stomatal occlusion and defoliation from varying levels of the Swiss needle cast (SNC) fungus, Phaeocryptopus gaeumannii. Levels of SNC disease and subsequent defoliation were manipulated by choosing six sites with varying levels of disease and by foliar applications of fungicides on six trees per site. Diurnal measurements of leaf water potential (Ψleaf), stomatal conductance (g s) and vapor pressure deficit (D) were made on six fungicide treated and six control trees per site. In addition, leaf specific hydraulic conductance was calculated on a single branch (K L_B) from three trees per treatment per site. Stomatal conductance at D=1 kPa (g sref) was negatively correlated with fungal colonization (number of fruiting bodies present in needle stomata) and positively correlated with K L_B. Despite reduced needle retention in diseased trees, K L declined due to a reduction in sapwood area and permeability (i.e., increasing presence of latewood in functional sapwood). In general, stomatal sensitivity to D for all foliage was consistent with stomatal regulation based on a simple hydraulic model [g s=K Lsoilleaf)/ D], which assumes strict stomatal regulation of Ψleaf. However, when fungal presence reduced maximum g s below the potential maximum supported by hydraulic architecture, stomatal sensitivity was lower than expected based on the theoretical relationship: dg s/dlnD=0.6·g sref. The results indicate that losses in productivity associated with physical blockage of stomata and defoliation are compounded by additional losses in K L and a reduction in g s in remaining functional stomata.  相似文献   

11.
Solidago canadensis is an invasive species from North America that is spreading across Europe, Australia and temperate Asia. We hypothesized that the species' wide ecological amplitude is also based on its potential in hydraulic acclimation, and analyzed hydraulic and anatomical properties along a transect with decreasing soil humidity. Stem hydraulic conductivity, vulnerability to drought‐induced embolism, stomatal closure during dehydration and xylem‐anatomical parameters were quantified at three sites. At the humid site, specific hydraulic conductivity of stems (1.0 ± 0.2 kg m–1 MPa–1 s–1) was about twofold higher, and leaf‐specific conductivity about 1.5 times higher (3.1 ± 0.5 kg m–1 MPa–1 s–1) than at the dry site. Water potential (Ψ) at 50% loss of conductivity was ?3.7 ± 0.1 MPa at the dry site and ?3.1 ± 0.2 MPa at the humid site (September). Vulnerability to drought‐induced embolism decreased along the transect and over the vegetation period. At drier sites, stomata started closing at lower Ψ while complete stomatal closure was reached at less negative Ψ (12% of maximum stomatal conductance: –2.5 ± 0.0 and ?3.0 ± 0.2 MPa at the dry and humid site). The safety margin between stomatal closure and 50% loss of conductivity was 1.2 and 0.2 MPa at the dry and humid sites. The observed variability indicated an efficient acclimation in hydraulic conductivity and safety: plants at dry sites exhibited lower specific hydraulic conductivity, higher embolism resistance and broader safety margins, signifying a trade‐off between the hydraulic safety and efficiency. The observed intraspecific plasticity in hydraulic and anatomical traits may help to explain the invasive potential of this species.  相似文献   

12.
13.
Two experiments examined simultaneous changes in leaf area (AL), root length (Lr), stomatal conductance (gs), leaf water potential (ΨL), transpiration and hydraulic plant conductance per unit leaf area (G) during the first three shoot cycles of northern red oak (Quercus rubra L.) grown under favourable and controlled conditions. Each shoot cycle consisted of bud swell, stem elongation, leaf expansion and rest; roots grew almost continuously. The gs of all leaves decreased substantially while leaves of the newest flush were expanding and increased modestly when seedling leaf area remained constant. Overall, gs decreased. The ΨL of mature leaves decreased during leaf expansion and increased by an equivalent amount during intervening periods. Possible explanations for the paired changes in gs and ΨL are considered. Changes in G closely paralleled those of canopy gs. These parallel changes during polycyclic seedling growth should act to keep seedling ΨL relatively constant as plant size increases and thereby help prevent ΨL from dropping to levels that would cause runaway embolism.  相似文献   

14.
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 (gs) and leaf water potential (Ψleaf). All clones experienced a substantial decrease in root‐specific root hydraulic conductance (Kroot‐r) in response to the water stress, but leaf‐specific shoot hydraulic conductance (Kshoot‐l) did not change in any of the clones. The reduction in Kroot‐r caused a decrease in leaf‐specific whole‐plant hydraulic conductance (Kplant‐l). Among clones, the larger the decrease in Kplant‐l, the more stomata closed in response to drought. Rewatering resulted in a quick recovery of Kroot‐r and gs. Our results demonstrated that the reduction in Kplant‐l, attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψleaf as water stress started. We concluded that higher Kplant‐l is associated with water stress resistance by sustaining a less negative Ψleaf and delaying stomatal closure.  相似文献   

15.
We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (gs) and xylem cavitation vulnerability. In addition, maximum gs was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum gs increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ50) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ50 values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.  相似文献   

16.
The impact of water deficit on stomatal conductance (g(s)), petiole hydraulic conductance (K(petiole)), and vulnerability to cavitation (PLC, percentage loss of hydraulic conductivity) in leaf petioles has been observed on field-grown vines (Vitis vinifera L. cv. Chasselas). Petioles were highly vulnerable to cavitation, with a 50% loss of hydraulic conductivity at a stem xylem water potential (Ψ(x)) of -0.95?MPa, and up to 90% loss of conductivity at a Ψ(x) of -1.5?MPa. K(petiole) described a daily cycle, decreasing during the day as water stress and evapotranspiration increased, then rising again in the early evening up to the previous morning's K(petiole) levels. In water-stressed vines, PLC increased sharply during the daytime and reached maximum values (70-90%) in the middle of the afternoon. Embolism repair occurred in petioles from the end of the day through the night. Indeed, PLC decreased in darkness in water-stressed vines. PLC variation in irrigated plants showed the same tendency, but with a smaller amplitude. The Chasselas cultivar appears to develop hydraulic segmentation, in which petiole cavitation plays an important role as a 'hydraulic fuse', thereby limiting leaf transpiration and the propagation of embolism and preserving the integrity of other organs (shoots and roots) during water stress. In the present study, progressive stomatal closure responded to a decrease in K(petiole) and an increase in cavitation events. Almost total closure of stomata (90%) was measured when PLC in petioles reached >90%.  相似文献   

17.

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 (Ψpd) or xylem water potential (Ψxylem) in Zserrata. During the gentle increase of PLC in Mazedarach, 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 Ψxylem 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.  相似文献   

18.
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.  相似文献   

19.
A study of a mutant variety of Zea mays (ON8147) revealed that the mutant plants, in contrast with normal maize plants, do not exhibit a light-induced increase in the rate of transpiration, and that the ontogeny of the stomatal complex is abnormal. In later stages of differentiation, the guard cells of mutant plants deteriorate, leaving the mature stomata with only the two subsidiary cells. The subsidiary cells in stomata of mutant leaves are similar to those of normal leaves with respect to their capacity to accumulate K+ in the dark, but they do not lose K+ in the light, as do subsidiary cells of stomata of nonmutant plants. It is suggested that impairment of guard cell function causes death of the mutant plant seedlings primarily by restricting CO2 entry into the leaf.  相似文献   

20.
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 (kL) 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 (ΨL), ranged from 70 to 150 mmol m?2 s?1 MPa?1. Hydraulic efficiency was positively correlated with kL (r2 = 0·86). Minimum annual ΨL ranged from ? 1·5 to ? 4·1 MPa among the six species. Seasonal and diurnal variation in ΨL were associated with differences among species in wood‐saturated water content, wood density and kL. The species with higher wood‐saturated water content were more efficient in terms of long‐distance water transport, exhibited smaller diurnal variation in ΨL and higher maximum photosynthetic rates. Smaller diurnal variation in ΨL in species with higher wood‐saturated water content, kL and hydraulic efficiency was not associated with stomatal restriction of transpiration when soil water deficit was moderate, but avoidance of low minimum seasonal ΨL in these species was associated with a substantial seasonal decline in g. Low seasonal minimum ΨL in species with low kL, 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 kL and stem water storage capacity.  相似文献   

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