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Establishing drying-limits for mortality of different tree species and understanding the anatomical and physiological traits involved is crucial to predict forests' responses to climate change. The xylem of Eucalyptus camaldulensis presents a complex of solitary vessels surrounded by different imperforate tracheary elements and parenchyma that influence, in a poorly known way, its hydraulic functioning. We aimed at describing the dynamics of embolism propagation in this type of xylem, seeking any vessel-size pattern, and unraveling the threshold of xylem embolism leading to nonrecovery after drought in E. camaldulensis. We assigned potted saplings to a protracted water-stress for 70 days. We relied on colorimetric and hydraulic methods to test for links between xylem anatomy and embolism propagation in the main stem. On average, the occurrence of embolism was randomly distributed in the stem xylem, but the probability of embolized vessels was higher than predicted by chance in the narrowest vessels of individuals that experienced low to moderate water-stress. The saplings could recover from severe water-stress if their percentage loss of conductance (PLC) was <77%, but not when the PLC was ˃ 85%. We concluded that, contrary to results reported for most species, the narrowest vessels are the most vulnerable to cavitation in E. camaldulensis, suggesting a lack of tradeoff between xylem efficiency and safety (in response to drought) at the tissue level. These results challenge the well-established paradigm of the effect of vessel size on cavitation, which states that the widest conduits are the most vulnerable to both freeze-thaw and drought-induced cavitation., (© 2021 Scandinavian Plant Physiology Society.)

While the xylem hydraulic properties, such as vulnerability to cavitation (VC), are of paramount importance in drought resistance, their genetic determinants remain unexplored. There is evidence that pectins and their methylation pattern are involved, but the detail of their involvement and the corresponding genes need to be clarified. We analyzed the hydraulic properties of the 35S::PME1 transgenic aspen that ectopically under- or over-express a xylem-abundant pectin methyl esterase, PtxtPME1. We also produced and analyzed 4CL1::PGII transgenic poplars expressing a fungal polygalacturonase, AnPGII, under the control of the Ptxa4CL1 promoter that is active in the developing xylem after xylem cell expansion. Both the 35S::PME1 under- and over-expressing aspen lines developed xylem with lower-specific hydraulic conductivity and lower VC, while the 4CL1::PGII plants developed xylem with a higher VC. These xylem hydraulic changes were associated with modifications in xylem structure or in intervessel pit structure that can result in changes in mechanical behavior of the pit membrane. This study shows that homogalacturonans and their methylation pattern influence xylem hydraulic properties, through its effect on xylem cell expansion and on intervessel pit properties and it show a role for PtxtPME1 in the xylem hydraulic properties., (© 2018 Scandinavian Plant Physiology Society.)

Background and Aims: Extreme water stress episodes induce tree mortality, but the physiological mechanisms causing tree death are still poorly understood. This study tests the hypothesis that a potted tree's ability to survive extreme monotonic water stress is determined by the cavitation resistance of its xylem tissue., Methods: Two species were selected with contrasting cavitation resistance (beech and poplar), and potted juvenile trees were exposed to a range of water stresses, causing up to 100 % plant death., Key Results: The lethal dose of water stress, defined as the xylem pressure inducing 50 % mortality, differed sharply across species (1·75 and 4·5 MPa in poplar and beech, respectively). However, the relationships between tree mortality and the degree of cavitation in the stems were similar, with mortality occurring suddenly when >90 % cavitation had occurred., Conclusions: Overall, the results suggest that cavitation resistance is a causal factor of tree mortality under extreme drought conditions.

Understanding drought tolerance mechanisms requires knowledge about the induced weakness that leads to tree death. Bud survival is vital to sustain tree growth across seasons. We hypothesized that the hydraulic connection of the bud to stem xylem structures was critical for its survival. During an artificial drastic water stress, we carried out a census of bud metabolic activity of young Populus nigra L. trees by microcalorimetry. We monitored transcript expression of aquaporins (AQPs; plasma membrane intrinsic proteins (PIPs), X intrinsic proteins (XIPs) and tonoplast membrane intrinsic proteins (TIPs)) and measured local water status within the bud and tissues in the bearer shoot node by nuclear magnetic resonance (NMR) imaging. We found that the bud respiration rate was closely correlated with its water content and decreased concomitantly in buds and their surrounding bearer tissues. At the molecular level, we observed a modulation of AQP pattern expressions (PIP, TIP and XIP subfamilies) linked to water movements in living cells. However, AQP functions remain to be investigated. Both the bud and tree died beyond a threshold water content and respiration rate. Nuclear magnetic resonance images provided relevant local information about the various water reservoirs of the stem, their dynamics and their interconnections. Comparison of pith, xylem and cambium tissues revealed that the hydraulic connection between the bud and saturated parenchyma cells around the pith allowed bud desiccation to be delayed. At the tree death date, NMR images showed that the cambium tissues remained largely hydrated. Overall, the respiration rate (Rco2) and a few AQP isoforms were found to be two suitable, complementary criteria to assess the bud metabolic activity and the ability to survive a severe drought spell. Bud moisture content could be a key factor in determining the capacity of poplar to recover from water stress.

Xylem cavitation resistance is a key physiological trait correlated with species tolerance to extreme drought stresses. Little is known about the genetic variability and phenotypic plasticity of this trait in natural tree populations. Here we measured the cavitation resistance of 17 Fagus sylvatica populations representative of the full range of the species in Europe. The trees were grown in three field trials under contrasting climatic conditions. Our findings suggest that the genotypic variability of cavitation resistance is high between genotypes of a given population. By contrast, no significant differences were found for this trait across populations, the mean population cavitation resistance being remarkably constant in each trial. We found a significant site effect and a significant site × population interaction, suggesting that cavitation resistance has a high phenotypic plasticity and that this plasticity is under genetic control. The implications of our findings for beech forest management in a context of climate change are discussed.

Tests were carried out to determine whether variations in the hydraulic architecture of eight Populus deltoides×Populus nigra genotypes could be related to variations in leaf function and growth performance. Measurements were performed in a coppice plantation on 1-year-old shoots under optimal irrigation. Hydraulic architecture was characterized through estimates of hydraulic efficiency (the ratio of conducting sapwood area to leaf area, A(X):A(L); leaf- and xylem-specific hydraulic conductance of defoliated shoots, k(SL) and k(SS), respectively; apparent whole-plant leaf-specific hydraulic conductance, k(plant)) and xylem safety (water potential inducing 50% loss in hydraulic conductance). The eight genotypes spanned a significant range of k(SL) from 2.63  kg s(-1) m(-2) MPa(-1) to 4.18  kg s(-1) m(-2) MPa(-1), variations being mostly driven by k(SS) rather than A(X):A(L). There was a strong trade-off between hydraulic efficiency and xylem safety. Values of k(SL) correlated positively with k(plant), indicating that high-pressure flowmeter (HPFM) measurements of stem hydraulic efficiency accurately reflected whole-plant water transport efficiency of field-grown plants at maximum transpiration rate. No clear relationship could be found between hydraulic efficiency and either net CO(2) assimilation rates, water-use efficiency estimates (intrinsic water-use efficiency and carbon isotope discrimination against (13)C), or stomatal characteristics (stomatal density and stomatal pore area index). Estimates of hydraulic efficiency were negatively associated with relative growth rate. This unusual pattern, combined with the trade-off observed between hydraulic efficiency and xylem safety, provides the rationale for the positive link already reported between relative growth rate and xylem safety among the same eight P. deltoides×P. nigra genotypes.

Xylem vulnerability to cavitation is a key parameter in understanding drought resistance of trees. We determined the xylem water pressure causing 50% loss of hydraulic conductivity (P(50)), a proxy of vulnerability to cavitation, and we evaluated the variability of this trait at tree and population levels for Fagus sylvatica. We checked for the effects of light on vulnerability to cavitation of stem segments together with a time series variation of P(50). Full sunlight-exposed stem segments were less vulnerable to cavitation than shade-exposed ones. We found no clear seasonal change of P(50), suggesting that this trait was designed for a restricted period. P(50) varied for populations settled along a latitudinal gradient, but not for those sampled along an altitudinal gradient. Moreover, mountainside exposure seemed to play a major role in the vulnerability to cavitation of beech populations, as we observed the differences along north-facing sides but not on south-facing sides. Unexpectedly, both north-facing mountainside and northern populations appeared less vulnerable than those grown on the southern mountainside or in the South of France. These results on beech populations were discussed with respect to the results at within-tree level.

We examined the relationships between xylem resistance to cavitation and 16 structural and functional traits across eight unrelated Populus deltoides x Populus nigra genotypes grown under two contrasting water regimes. The xylem water potential inducing 50% loss of hydraulic conductance (Psi(50)) varied from -1.60 to -2.40 MPa. Drought-acclimated trees displayed a safer xylem, although the extent of the response was largely genotype dependent, with Psi(50) being decreased by as far as 0.60 MPa. At the tissue level, there was no clear relationship between xylem safety and either xylem water transport efficiency or xylem biomechanics; the only structural trait to be strongly associated with Psi(50) was the double vessel wall thickness, genotypes exhibiting a thicker double wall being more resistant. At the leaf level, increased cavitation resistance was associated with decreased stomatal conductance, while no relationship could be identified with traits associated with carbon uptake or bulk leaf carbon isotope discrimination, a surrogate of intrinsic water-use efficiency. At the whole-plant level, increased safety was associated with higher shoot growth potential under well-irrigated regime only. We conclude that common trade-offs between xylem resistance to cavitation and other physiological traits that are observed across species may not necessarily hold true at narrower scales.

Enhanced sapling growth in advance regeneration requires gaps in the canopy, but is often delayed after canopy opening, because acclimation of saplings to the new environment is gradual and may last for several years. Canopy opening is expected to result in an increased transpiration because of a larger climatic demand and a higher stomatal conductance linked to the higher rates of photosynthesis. Therefore, we focused on the changes in water relations and the hydraulic properties of beech (Fagus sylvatica L.) saplings during 2 years after canopy opening. We tested the hypothesis that an increase in leaf-specific hydraulic conductance and a decrease in vulnerability to cavitation occur to sustain an enhanced transpiration. Hydraulic conductance of defoliated shoots, vulnerability to cavitation, size and density of xylem vessels as well as stomatal conductance were recorded on saplings growing in shade (S saplings) or in gaps created by opening the canopy (shade-to-light, SL saplings). Hydraulic conductance per unit cross-sectional area (K(AS)) did not differ in the shoots of S and SL saplings. But a higher ratio stem cross-sectional area/leaf area resulted in a higher leaf-specific hydraulic conductance of the shoots (K(AL)) of SL saplings. Contrary to expectations, vulnerability to cavitation increased transitorily in stems during the first year after canopy opening and no difference was observed between the two treatments in light-saturated stomatal conductance. During the second year, vulnerability to cavitation was similar in the S and SL saplings and light-saturated stomatal conductance increased in SL saplings. These results demonstrate a release of the hydraulic constraints after canopy opening with an adjustment of the ratio stem cross-sectional area/leaf area. But the larger vulnerability to cavitation during the first year could limit stomatal opening and therefore the ability of beech saplings to use the available light for photosynthesis and could therefore partly explain why the growth increase was delayed to the second growing season after canopy opening.

Molecular and physiological studies in walnut (Juglans regia) are combined to establish the putative role of leaf plasma membrane aquaporins in the response of leaf hydraulic conductance (K(leaf)) to irradiance. The effects of light and temperature on K(leaf) are described. Under dark conditions, K(leaf) was low, but increased by 400% upon exposure to light. In contrast to dark conditions, K(leaf) values of light-exposed leaves responded to temperature and 0.1 mm cycloheximide treatments. Furthermore, K(leaf) was not related to stomatal aperture. Data of real-time reverse transcription-polymerase chain reaction showed that K(leaf) dynamics were tightly correlated with the transcript abundance of two walnut aquaporins (JrPIP2,1 and JrPIP2,2). Low K(leaf) in the dark was associated with down-regulation, whereas high K(leaf) in the light was associated with up-regulation of JrPIP2. Light responses of K(leaf) and aquaporin transcripts were reversible and inhibited by cycloheximide, indicating the importance of de novo protein biosynthesis in this process. Our results indicate that walnut leaves can rapidly change their hydraulic conductance and suggest that these changes can be explained by regulation of plasma membrane aquaporins. Model simulation suggests that variable leaf hydraulic conductance in walnut might enhance leaf gas exchanges while buffering leaf water status in response to ambient light fluctuations.

We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2'N, annual precipitation 2200 mm). We assessed leaf δ 15 N as a presumptive indicator of symbiotic N 2 fixation, and leaf and wood cellulose δ 13 C as an indicator of leaf intrinsic water-use efficiency (CO 2 assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N 2 fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ 15 N was spatially quite uniform in the forest (range 3-5‰), whereas average leaf δ 15 N ranged from -0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g -1 . Although nodulated trees displayed significantly lower leaf δ 15 N values than non-nodulated trees, leaf δ 15 N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ 15 N values for nodulated and non-nodulated species at the lower end of the δ 15 N range. Perturbation did not markedly affect the difference δ 15 N soil  -   δ 15 N leaf , and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ 13 C were observed in the forest (average values from -31.4 to -26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO 2 assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ 13 C was positively related to total leaf δ 13 C, the former values being 2-3‰ higher than the latter ones. Leaf δ 13 C was not related to leaf δ 15 N at either intraspecific or interspecific levels. δ 13 C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ 13 C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species.

Canopy CO 2 concentrations in a tropical rainforest in French Guiana were measured continuously for 5 days during the 1994 dry season and the 1995 wet season. Carbon dioxide concentrations ([CO 2 ]) throughout the canopy (0.02-38 m) showed a distinct daily pattern, were well-stratified and decreased with increasing height into the canopy. During both seasons, daytime [CO 2 ] in the upper and middle canopy decreased on average 7-10 μmol mol -1 below tropospheric baseline values measured at Barbados. Within the main part of the canopy (≥ 0.7 m), [CO 2 ] did not differ between the wet and dry seasons. In contrast, [CO 2 ] below 0.7 m were generally higher during the dry season, resulting in larger [CO 2 ] gradients. Supporting this observation, soil CO 2 efflux was on average higher during the dry season than during the wet season, either due to diffusive limitations and/or to oxygen deficiency of root and microbial respiration. Soil respiration rates decreased by 40% after strong rain events, resulting in a rapid decrease in canopy [CO 2 ] immediately above the forest floor of about 50␣μmol mol -1 . Temporal and spatial variations in [CO 2 ] canopy were reflected in changes of δ 13 C canopy and δ 18 O canopy values. Tight relationships were observed between δ 13 C and δ 18 O of canopy CO 2 during both seasons (r 2  > 0.86). The most depleted δ 13 C canopy and δ 18 O canopy values were measured immediately above the forest floor (δ 13 C = -16.4‰; δ 18 O = 39.1‰ SMOW). Gradients in the isotope ratios of CO 2 between the top of the canopy and the forest floor ranged between 2.0‰ and 6.3‰ for δ 13 C, and between 1.0‰ and 3.5‰ for δ 18 O. The δ 13 C leaf and calculated c i /c a of foliage at three different positions were similar for the dry and wet seasons indicating that the canopy maintained a constant ratio of photosynthesis to stomatal conductance. About 20% of the differences in δ 13 C leaf within the canopy was accounted for by source air effects, the remaining 80% must be due to changes in c i /c a . Plotting 1/[CO 2 ] vs. the corresponding δ 13 C ratios resulted in very tight, linear relationships (r 2  = 0.99), with no significant differences between the two seasons, suggesting negligible seasonal variability in turbulent mixing relative to ecosystem gas exchange. The intercepts of these relationships that should be indicative of the δ 13 C of respired sources were close to the measured δ 13 C of soil respired CO 2 and to the δ 13 C of litter and soil organic matter. Estimates of carbon isotope discrimination of the entire ecosystem, Δ e , were calculated as 20.3‰ during the dry season and as 20.5‰ during the wet season.