Sapwood-specific Conductivity Research Articles

Xylem anatomical structure as a determinant of hydraulic trait variation in C3 plant Reaumuria soongorica and C4 plant Salsola passerina

Moisture variation significantly impacts hydraulic traits and xylem anatomical structure in woody plants, thereby influencing water transport and embolism resistance. However, the precise relationship between these factors within desert shrubs remains unclear, and it may be related to plant functional types. We conducted a water manipulation experiment involving seedlings of the C3 plant Reaumuria soongorica and the C4 plant Salsola passerina within a rain shelter. We applied three water treatments: control, chronic drought, and flash drought. After a period of drought treatment, seedlings were rewatered to control level for 15 days, after which hydraulic traits and anatomical structures were measured. We found that: (1) vessel density was positively correlated with sapwood specific conductivity for R. soongorica, while hydraulic weighted vessel diameter positively correlated with embolism resistance; both hydraulic weighted vessel diameter and vessel internal diameter span was positively correlated with sapwood specific conductivity for S. passerina; (2) R. soongorica had lower edge density yet higher modularity compared to S. passerina; (3) sapwood and leaf specific conductivity emerged as hub traits in R. soongorica, while vessel internal diameter span was identified as a hub trait in S. passerina, thus serving as important predictors of hydraulic function and related attributes. Our study demonstrates distinct hydraulic strategies in the two species. R. soongorica employs a highly modular trait combination to adapt to water fluctuations, maintaining both high hydraulic efficiency and embolism resistance under drought stress. Conversely, S. passerina integrates traits for efficient resource access, ensuring hydraulic efficiency but compromising embolism resistance under drought stress. Integrating hydraulic traits with anatomical structures significantly enhances predictions of desert shrub resilience to drought within the context of global climate change.

Optimising height-growth predicts trait responses to water availability and other environmental drivers.

Future changes in climate, together with rising atmospheric , may reorganise the functional composition of ecosystems. Without long-term historical data, predicting how traits will respond to environmental conditions-in particular, water availability-remains a challenge. While eco-evolutionary optimality theory (EEO) can provide insight into how plants adapt to their environment, EEO approaches to date have been formulated on the assumption that plants maximise carbon gain, which omits the important role of tissue construction and size in determining growth rates and fitness. Here, we show how an expanded optimisation framework, focussed on individual growth rate, enables us to explain shifts in four key traits: leaf mass per area, sapwood area to leaf area ratio (Huber value), wood density and sapwood-specific conductivity in response to soil moisture, atmospheric aridity, and light availability. In particular, we predict that as conditions become increasingly dry, height-growth optimising traits shift from resource-acquisitive strategies to resource-conservative strategies, consistent with empirical responses across current environmental gradients of rainfall. These findings can explain both the shift in traits and turnover of species along existing environmental gradients and changing future conditionsand highlight the importance of both carbon assimilation and tissue construction in shaping the functional composition of vegetation across climates.

Hydraulic Traits and Non-Structural Carbon Responses to Drought Stress in Reaumuria soongorica (Pall.) Maxim. and Salsola passerina Bunge

Drought-induced plant mortality, resulting from either hydraulic failure or carbon starvation, is hypothesized to be modulated by the drought intensity. However, there is a paucity of research investigating the response strategies in desert shrubs under drought stress with different intensities. We transplanted potted Reaumuria soongorica (Pall.) Maxim. and Salsola passerina Bunge seedlings in the rain-out shelter, and implemented three water treatments: a control (well-watered, CK), a chronic drought (gradually less watered, CD), and a flash drought (not watered, FD). We then quantified plant physiological traits associated with water use and carbon assimilation. Both R. soongorica and S. passerina showed similar changes in water use and carbon characteristics under different drought treatments. Water use efficiency was not significantly changed, but embolism resistance was significantly lower in CD, and leaf specific conductivity and embolism resistance were significantly lower in FD compared to CK. Under the drought treatment, both shrubs had significantly lower hydraulic safety margins than CK, with FD being significantly lower than CD. Notably, FD had lower carbon assimilation and a lower leaf non-structural carbon concentration, but higher stem non-structural carbon concentration. The results of a principal component analysis showed that net photosynthetic rate, sapwood specific conductivity, embolism resistance, midday water potential, and leaf and stem soluble sugar concentration were the main axes of variation for R. soongorica traits. CK had the highest water use efficiency, net photosynthetic rate, and gas exchange rate, while FD had the lowest embolism resistance and highest osmoregulation. Midday water potential, leaf and stem soluble sugar concentration were the main axes of variation for S. passerina traits, and individual distribution under three water treatments was associated with drought tolerance traits. The findings suggest that species exhibit different response strategies for resistance to drought stress, with R. soongorica being drought-avoidant and S. passerina being drought-tolerant. These findings highlight the adaptive capacity of desert shrubs to water deficit and provide insights for assessing hydraulic failure and carbon starvation in desert shrubs.

How Climate Shapes the Functioning of Tropical Montane Cloud Forests

Tropical Montane Cloud Forest (TMCF) is a highly vulnerable ecosystem, which occurs at higher elevations in tropical mountains. Many aspects of TMCF vegetation functioning are poorly understood, making it difficult to quantify and project TMCF vulnerability to global change. We compile functional traits data to provide an overview of TMCF functional ecology. We use numerical models to understand the consequences of TMCF functional composition with respect to its responses to climate and link the traits of TMCF to its environmental conditions. TMCF leaves are small and have low SLA but high Rubisco content per leaf area. This implies that TMCF maximum net leaf carbon assimilation (An) is high but often limited by low temperature and leaf wetting. Cloud immersion provides important water and potentially nutrient inputs to TMCF plants. TMCF species possess low sapwood specific conductivity, which is compensated with a lower tree height and higher sapwood to leaf area ratio. These traits associated with a more conservative stomatal regulation results in a higher hydraulic safety margin than nearby forests not affected by clouds. The architecture of TMCF trees including its proportionally thicker trunks and large root systems increases tree mechanical stability. The TMCF functional traits can be conceptually linked to its colder and cloudy environment limiting An, growth, water transport and nutrient availability. A hotter climate would drastically affect the abiotic filters shaping TMCF communities and potentially facilitate the invasion of TMCF by more productive lowland species.

Plant Hydraulic Trait Covariation: A Global Meta-Analysis to Reduce Degrees of Freedom in Trait-Based Hydrologic Models

Current vegetation modeling strategies use broad categorizations of plants to estimate transpiration and biomass functions. A significant source of model error stems from vegetation categorizations that are mostly taxonomical with no basis in plant hydraulic strategy and response to changing environmental conditions. Here, we compile hydraulic traits from 355 species around the world to determine trait covariations in order to represent hydraulic strategies. Simple and stepwise regression analyses demonstrate the interconnectedness of multiple vegetative hydraulic traits, specifically, traits defining hydraulic conductivity and vulnerability to embolism with wood density and isohydricity. Drought sensitivity is strongly (Adjusted R2 = 0.52, p < 0.02) predicted by a stepwise linear model combining rooting depth, wood density, and isohydricity. Drought tolerance increased with increasing wood density and anisohydric response, but with decreasing rooting depth. The unexpected response to rooting depth may be due to other tradeoffs within the hydraulic system. Rooting depth was able to be predicted from sapwood specific conductivity and the water potential at 50% loss of conductivity. Interestingly, the influences of biome or growth form do not increase the accuracy of the drought tolerance model and were able to be omitted. Multiple regression analysis revealed 3D trait spaces and tradeoff axes along which species’ hydraulic strategies can be analyzed. These numerical trait spaces can reduce the necessary input to and parameterization of plant hydraulics modules, while increasing the physical representativeness of such simulations.

Branch age and light conditions determine leaf-area-specific conductivity in current shoots of Scots pine.

Shoot size and other shoot properties more or less follow the availability of light, but there is also evidence that the topological position in a tree crown has an influence on shoot development. Whether the hydraulic properties of new shoots are more regulated by the light or the position affects the shoot acclimation to changing light conditions and thereby to changing evaporative demand. We investigated the leaf-area-specific conductivity (and its components sapwood-specific conductivity and Huber value) of the current-year shoots of Scots pine (Pinus sylvestris L.) in relation to light environment and topological position in three different tree classes. The light environment was quantified in terms of simulated transpiration and the topological position was quantified by parent branch age. Sample shoot measurements included length, basal and tip diameter, hydraulic conductivity of the shoot, tracheid area and density, and specific leaf area. In our results, the leaf-area-specific conductivity of new shoots declined with parent branch age and increased with simulated transpiration rate of the shoot. The relation to transpiration demand seemed more decisive, since it gave higher R(2) values than branch age and explained the differences between the tree classes. The trend of leaf-area-specific conductivity with simulated transpiration was closely related to Huber value, whereas the trend of leaf-area-specific conductivity with parent branch age was related to a similar trend in sapwood-specific conductivity.

Shifts in Leaf and Stem Hydraulic Traits across Aridity Gradients in Eastern Australia

Premise of research. Plants are faced with a challenge across all climates they inhabit—they must transport water to their leaves so that photosynthesis can take place. Although this is simple in concept, it can be achieved by different arrangements of root, stem, and leaf traits. The hydraulic functioning of species across aridity gradients is determined by the coordination of these traits. Nevertheless, we have an imperfect understanding of which trait shifts are favored across aridity gradients as well as the alignment of trait shifts with climate.Methodology. We measured hydraulic traits relating to Darcy’s law for 120 angiosperm species across a broad range of climates in eastern Australia; nearly one-third of all biome space on Earth was represented. We then determined which hydraulic trait shifts have been favored across aridity gradients and which climate characteristics these trait shifts aligned with.Pivotal results. Increasing aridity, from climates with similar precipitation and evaporation to climates where precipitation was only a third of evaporation, was associated with a 4.8-fold decrease in plant height, a 3.0-fold decrease in leaf area–to–sapwood area ratio, and a 3.3-fold decrease in leaf water potential. However, sapwood-specific conductivity decreased by 5.9-fold, more than any other hydraulic trait. Greater sapwood-specific conductivity (decreasing resistance) at wet sites compensated for increasing resistance and hydraulic demand that was associated with taller plants and leafier shoots. All hydraulic traits were strongly correlated with growth season aridity (; ) but were not correlated with maximum aridity. This suggests that plant hydraulic traits are most responsive to water availability and evaporative demand present during the most suitable months for growth rather than the driest months.Conclusions. We suggest that evolution has equipped plants with various mechanisms to avoid desiccation during the dry season while optimizing hydraulic traits for carbon gain during the growth season.

Xylem traits mediate a trade‐off between resistance to freeze–thaw‐induced embolism and photosynthetic capacity in overwintering evergreens

Hydraulic traits were studied in temperate, woody evergreens in a high-elevation heath community to test for trade-offs between the delivery of water to canopies at rates sufficient to sustain photosynthesis and protection against disruption to vascular transport caused by freeze-thaw-induced embolism. Freeze-thaw-induced loss in hydraulic conductivity was studied in relation to xylem anatomy, leaf- and sapwood-specific hydraulic conductivity and gas exchange characteristics of leaves. We found evidence that a trade-off between xylem transport capacity and safety from freeze-thaw-induced embolism affects photosynthetic activity in overwintering evergreens. The mean hydraulically weighted xylem vessel diameter and sapwood-specific conductivity correlated with susceptibility to freeze-thaw-induced embolism. There was also a strong correlation of hydraulic supply and demand across species; interspecific differences in stomatal conductance and CO(2) assimilation rates were correlated linearly with sapwood- and leaf-specific hydraulic conductivity. Xylem vessel anatomy mediated an apparent trade-off between resistance to freeze-thaw-induced embolism and hydraulic and photosynthetic capacity during the winter. These results point to a new role for xylem functional traits in determining the degree to which species can maintain photosynthetic carbon gain despite freezing events and cold winter temperatures.

Plasticity in the Huber value contributes to homeostasis in leaf water relations of a mallee Eucalypt with variation to groundwater depth

Information on how vegetation adapts to differences in water supply is critical for predicting vegetation survival, growth and water use, which, in turn, has important impacts on site hydrology. Many field studies assess adaptation to water stress by comparing between disparate sites, which makes it difficult to distinguish between physiological or morphological changes and long-term genetic adaptation. When planting trees into new environments, the phenotypic adaptations of a species to water stress will be of primary interest. This study examined the response to water availability of Eucalyptus kochii ssp. borealis (C. Gardner) D. Nicolle, commonly integrated with agriculture in south-western Australia for environmental and economic benefits. By choosing a site where the groundwater depth varied but where climate and soil type were the same, we were able to isolate tree response to water supply. Tree growth, leaf area and stand water use were much larger for trees over shallow groundwater than for trees over a deep water table below a silcrete hardpan. However, water use on a leaf area basis was similar in trees over deep and shallow groundwater, as were the minimum leaf water potential observed over different seasons and the turgor loss point. We conclude that homeostasis in leaf water use and water relations was maintained through a combination of stomatal control and adjustment of sapwood-to-leaf area ratios (Huber value). Differences in the Huber value with groundwater depth were associated with different sapwood-specific conductivity and water use on a sapwood area basis. Knowledge of the coordination between water supply, leaf area, sapwood area and leaf transpiration rate for different species will be important when predicting stand water use.

Dynamic variation in sapwood specific conductivity in six woody species

Our goals were to quantify how non-embolism-inducing pressure gradients influence trunk sapwood specific conductivity (k(s)) and to compare the impacts of constant and varying pressure gradients on k(s) with KCl and H2O as the perfusion solutions. We studied six woody species (three conifers and three angiosperms) which varied in pit membrane structure, pit size and frequency of axial water transport across pits (long versus short conduits). Both stepwise ("steady") and nonlinear continuous ("non-steady") decreases in the pressure gradient led to decreased k(s) in all species but white oak (Quercus garryana Dougl. ex Hook), a ring-porous and long-vesseled angiosperm. In one diffuse-porous angiosperm (red alder, Alnus rubra Bong.) and two conifers (western red cedar, Thuja plicata Donn. ex D. Don, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco), k(s) was 10-30% higher under steady pressure gradients than under non-steady pressure gradients, and a decrease in the pressure gradient from 0.15 to 0.01 MPa m(-1) caused a 20-42% decrease in k(s). In another diffuse-porous angiosperm (maple, Acer macrophyllum Pursh) and in a third coniferous species (western hemlock, Tsuga heterophylla (Raf.) Sarg), there was no difference between k(s) measured under steady and non-steady pressure gradients. With the exception of western red cedar, a conifer with simple pit membranes, the differences in k(s) between low and high pressure gradients tended to be lower in the conifers than in the diffuse-porous angiosperms. In Douglas-fir, western red cedar and the diffuse-porous angiosperms, k(s) was higher when measured with KCl than with H2O. In white oak, there were no differences in k(s) whether measured under steady or non-steady pressure gradients, or when xylem was perfused with KCl or H2O. The species differences in the behavior of k(s) suggest that elasticity of the pit membrane was the main factor causing k(s) to be disproportionate to the pressure gradient and to the different pressure regimes. The results imply that, if nonlinearities in pressure-flux relationships are ignored when modeling tree water relations in vivo, large errors will result in the predictions of tree water status and its impact on stomatal control of transpiration and photosynthesis.

Diversity of hydraulic traits in nine Cordia species growing in tropical forests with contrasting precipitation

Inter- and intraspecific variation in hydraulic traits was investigated in nine Cordia (Boraginaceae) species growing in three tropical rainforests differing in mean annual precipitation (MAP). Interspecific variation was examined for the different Cordia species found at each site, and intraspecific variation was studied in populations of the widespread species Cordia alliodora across the three sites. Strong intra- and interspecific variation were observed in vulnerability to drought-induced embolism. Species growing at drier sites were more resistant to embolism than those growing at moister sites; the same pattern was observed for populations of C. alliodora. By contrast, traits related to hydraulic capacity, including stem xylem vessel diameter, sapwood specific conductivity (K(s)) and leaf specific conductivity (K(L)), varied strongly but independently of MAP. For C. alliodora, xylem anatomy, K(s), K(L) and Huber value varied little across sites, with K(s) and K(L) being consistently high relative to other Cordia species. A constitutively high hydraulic capacity coupled with plastic or genotypic adjustment in vulnerability to embolism and leaf water relations would contribute to the ability of C. alliodora to establish and compete across a wide precipitation gradient.

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.

Dynamics of water transport and storage in conifers studied with deuterium and heat tracing techniques

The volume and complexity of their vascular systems make the dynamics of long-distance water transport in large trees difficult to study. We used heat and deuterated water (D2)) as tracers to characterize whole-tree water transport and storage properties in individual trees belonging to the coniferous species Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. The trees used in this study spanned a broad range of height (13.5-58 m) and diameter (0.14-1.43 m). Sap flow was monitored continuously with heat dissipation probes near the base of the trunk prior to, during and following injection of D2O. The transit time for D2O transport from the base of the trunk to the upper crown and the tracer residence time were determined by measuring hydrogen isotope ratios in water extracted from leaves sampled at regular intervals. Transit times for arrival of D2O in the upper crown ranged from 2.5 to 21 d and residence times ranged from 36 to 79 d. Estimates of maximum sap velocity derived from tracer transit times and path length ranged from 2.4 to 5.4 m d(-1). Tracer residence time and half-life increased as tree diameter increased, independent of species. Species-independent scaling of tracer velocity with sapwood-specific conductivity was also observed. When data from this study were combined with similar data from an earlier study of four tropical angiosperm trees, species-independent scaling of tracer velocity and residence time with sapwood hydraulic capacitance was observed. Sapwood capacitance is an intrinsic tissue-level property that appears to govern whole-tree water transport in a similar manner among both tracheid- and vessel-bearing species.

Ecological implications of xylem cavitation for several Pinaceae in the Pacific Northern USA

Abstract1. Xylem hydraulic properties and vulnerability to cavitation (determined using the air‐injection method) were studied in six Pinaceae of the northern Rocky Mountains: Pinus ponderosa, Pseudotsuga menziesii, Larix occidentalis, Pinus contorta, Pinus albicaulis and Abies lasiocarpa. We tested whether species extending into drier habitats exhibited increased resistance to water stress‐induced cavitation, and whether there is a trade‐off between xylem transport capacity and resistance to cavitation.2. At lower elevations the more drought‐tolerant P. ponderosa was much less resistant to cavitation than the codominant P. menziesii. Greater vulnerability to cavitation in P. ponderosa was compensated for, at least in part, by increased stomatal control of water loss (inferred from carbon isotope discrimination) and by increased sapwood to leaf area ratios. Similar differences, but less pronounced, were found in codominant species at higher elevations.3. Leaf specific hydraulic conductivity was negatively correlated with mean cavitation pressure. When species were separated into pines and non‐pines, sapwood specific conductivity and mean cavitation pressure were also negatively correlated within each of the two groups.4. Our results indicate that within the evergreen conifers examined, greater resistance to water stress‐induced cavitation is not required for survival in more xeric habitats, and that there is a trade‐off between xylem conductance and resistance to cavitation.

Ecological implications of xylem cavitation for several Pinaceae in the Pacific Northern USA

1. Xylem hydraulic properties and vulnerability to cavitation (determined using the air-injection method) were studied in six Pinaceae of the northern Rocky Mountains: Pinus ponderosa, Pseudotsuga menziesii, Larix occidentalis, Pinus contorta, Pinus albicaulis and Abies lasiocarpa. We tested whether species extending into drier habitats exhibited increased resistance to water stress-induced cavitation, and whether there is a trade-off between xylem transport capacity and resistance to cavitation. 2. At lower elevations the more drought-tolerant P ponderosa was much less resistant to cavitation than the codominant P menziesii. Greater vulnerability to cavitation in P ponderosa was compensated for, at least in part, by increased stomatal control of water loss (inferred from carbon isotope discrimination) and by increased sapwood to leaf area ratios. Similar differences, but less pronounced, were found in codominant species at higher elevations. 3. Leaf specific hydraulic conductivity was negatively correlated with mean cavitation pressure. When species were separated into pines and non-pines, sapwood specific conductivity and mean cavitation pressure were also negatively correlated within each of the two groups. 4. Our results indicate that within the evergreen conifers examined, greater resistance to water stress-induced cavitation is not required for survival in more xeric habitats, and that there is a trade-off between xylem conductance and resistance to cavitation.