Sapwood-specific Hydraulic Conductivity Research Articles

Water storage capacity is inversely associated with xylem embolism resistance in tropical karst tree species.

Tropical karst habitats are characterized by limited and patchy soil, large rocky outcrops and porous substrates, resulting in high habitat heterogeneity and soil moisture fluctuations. Xylem hydraulic efficiency and safety can determine the drought adaptation and spatial distribution of woody plants growing in karst environments. In this study, we measured sapwood-specific hydraulic conductivity (Ks), vulnerability to embolism, wood density, saturated water content, and vessel and pit anatomical characteristics in the branch stems of 12 evergreen tree species in a tropical karst seasonal rainforest in southwestern China. We aimed to characterize the effects of structural characteristics on hydraulic efficiency and safety. Our results showed that there was no significant correlation between Ks and hydraulic safety across the tropical karst woody species. Ks was correlated with hydraulic vessel diameter (r=0.80, P<0.05) and vessel density (r=-0.60, P<0.05), while the stem water potential at 50 and 88% loss of hydraulic conductivity (P50 and P88) were both significantly correlated with wood density (P<0.05) and saturated water content (P=0.052 and P<0.05, respectively). High stem water storage capacity was associated with low cavitation resistance possibly because of its buffering the moisture fluctuations in karst environments. However, both Ks and P50/P88 were decoupled from the anatomical traits of pit and pit membranes. This may explain the lack of tradeoff between hydraulic safety and efficiency in tropical karst evergreen tree species. Our results suggest that diverse hydraulic trait combination may facilitate species coexistence in karst environments with high spatial heterogeneity.

Climate-driven sapwood-specific hydraulic conductivity and the Huber value but not leaf-specific hydraulic conductivity on a global scale.

Efficient water transport is crucial for plant growth and survival. Plant hydraulic conductivity varies between functional groups and biomes and is strongly influenced by changing environmental conditions. However, correlations of conductivity-related hydraulic traits with climatic variables are not fully understood, preventing clarification of plant form and function under climate change scenarios. By compiling leaf-specific hydraulic conductivity (KL), sapwood-specific hydraulic conductivity (Ks), and Huber values (Hv, sapwood area to leaf area ratio) along with climatic variables including mean annual temperature (MAT), mean annual precipitation (MAP) and aridity index (AI) for 428 species across a wide range of plant functional types (PFTs) and biomes at a global scale, we found greater variability of KL within PFTs and biomes than across PFTs and biomes. Interaction effects between PFTs and biomes on KL and Ks were found. The interaction between MAT and MAP played a significant role in Ks and Hv (t = 3.89, P < 0.001 for Ks and t = -5.77, P < 0.001 for Hv). With increasing AI, Ks increased and Hv decreased. KL was not influenced by the investigated climatic variables. Our study provides a better understanding of the dynamics of hydraulic structure and function across functional groups and biomes and of the abiotic drivers of their large-scale variations.

Stem hydraulic conductivity and embolism resistance of Quercus species are associated with their climatic niche.

The hydraulic traits of a plant species may reflect its climate adaptations. Southwest China is considered as a biodiversity hotpot of the genus Quercus (oak). However, the hydraulic adaptations of Asian oaks to their climate niches remain unclear. Ten common garden-grown oak species with distinct natural distributions in eastern Asia were used to determine their stem xylem embolism resistance (water potential at 50% loss of hydraulic conductivity, P50), stem hydraulic efficiency (vessel anatomy and sapwood specific hydraulic conductivity (Ks)) and leaf anatomical traits. We also compiled four key functional traits: wood density, hydraulic-weighted vessel diameter, Ks and P50 data for 31 oak species from previous literature. We analyzed the relationship between hydraulic traits and climatic factors over the native ranges of 41 oak species. Our results revealed that the 10 Asian oak species, which are mainly distributed in humid subtropical habitats, possessed a stem xylem with low embolism resistance and moderate hydraulic efficiency. The deciduous and evergreen species of the 10 Asian oaks differed in the stem and leaf traits related to hydraulic efficiency. Ks differed significantly between the two phenological groups (deciduous and evergreens) in the 41-oak dataset. No significant difference in P50 between the two groups was found for the 10 Asian oaks or the 41-oak dataset. The oak species that can distribute in arid habitats possessed a stem xylem with high embolism resistance. Ks negatively related to the humidity of the native range of the 10 Asian oaks, but showed no trend when assessing the entire global oak dataset. Our study suggests that stem hydraulic conductivity and embolism resistance in Quercus species are shaped by their climate niche. Our findings assist predictions of oak drought resistance with future climate changes for oak forest management.

Trade-offs between xylem hydraulic efficiency and mechanical strength in Chinese evergreen and deciduous savanna species.

Evergreen and deciduous species coexist in tropical dry forests and savannas, but differ in physiological mechanisms and life-history strategies. Hydraulic conductivity and mechanical support are two major functions of the xylems of woody plant species related to plant growth and survival. In this study, we measured sapwood-specific hydraulic conductivity (Ks), leaf-specific hydraulic conductivity (KL), modulus of rupture (MOR) and elasticity (MOE), xylem anatomical traits and fiber contents in the xylems of 20 woody species with contrasting leaf phenology (evergreen vs deciduous) in a Chinese savanna. Our results showed that deciduous species had significantly higher Ks and KL but lower MOR and MOE than evergreen species. Evergreen species experienced more negative seasonal minimum water potential (Pmin) than deciduous species during the dry season. Furthermore, we found trade-offs between xylem hydraulic efficiency and mechanical strength across species and within the evergreen and deciduous groups, and these trade-offs were modulated by structural and chemical traits. Both Ks and KL were significantly related to hydraulic weighted vessel diameter (Dh) across all species and within the deciduous group. Both MOR and MOE were significantly related to wood density, neutral detergent fiber and acid detergent fiber across species and within evergreen and deciduous groups. Our findings demonstrated that Chinese evergreen and deciduous savanna species diverged in xylem hydraulic and mechanical functions, reflecting conservative and acquisitive life-history strategies for evergreen and deciduous species, respectively. This study provides new information with which to understand the hydraulic and biomechanical properties and ecological strategies of savanna species in long-term dry-hot environments.

Urbanization intensifies tree sap flux but divergently for different tree species groups in China.

In recent years, positive and negative effects of urbanization on forest ecosystem have been reported by many studies, while some uncertainties about the impact of urbanization-induced spatial heterogeneity of environmental factors on forest systems still remain unclear. In this study, we analyzed the urbanization effects on sap flux of a common subtropical evergreen tree species Schima superba along an urban-rural gradient in Guangdong Province, South China, and identified the consistency of these results among different groups (evergreen, deciduous, and coniferous species) using data from 83 previously published studies in China. The mean sap flux density (Fd) of S. superba in Xiaoqingshan (XQS), Heshan (HS), Dinghushan (DHS), and Shimentai (SMT), along the urban-rural gradient was 40.9gm-2s-1, 32.1gm-2s-1, 17.0gm-2s-1, and 17.5gm-2s-1, respectively, presenting a decreasing trend with the diminishing urbanization. This pattern in Fd tended to enlarge with tree size and was well confirmed by the enhanced leaf transpiration rate (by 119%) and photosynthetic rate (by 8.8%) for the S. superba in another urbanization gradient from the urban (Hangzhou, denoted as "HZ") to rural sites (Jiande, denoted as "JD") in Zhejiang Province, East China, which has similar climatic condition and urbanization with Guangdong Province. We attributed such positive effects to the decreased sapwood density and specific leaf area (SLA), as well as the increased Huber value (sap wood area/leaf area) and the sap wood specific hydraulic conductivity (KS). We also found that pollutant emission exerted more impact on Fd than climatic factors change, since the variation of the latter was not large enough to cause significant change of Fd under the same climatic zone. In addition, we conducted a principal component analysis (PCA) based on the published 83 studies. Results showed Fd of evergreen tree species was related positively to principle 1 and negatively to principle 2, respectively, whereas the Fd of deciduous broadleaf and coniferous tree species was positively and negatively related to both principles, respectively. This study demonstrated the potential impact of urbanization-related pollutant emission changes on water use of forest trees and the growth among different groups.

Altitudinal variations of hydraulic traits in Faxon fir (Abies fargesii var. faxoniana): Mechanistic controls and environmental adaptability

Global climate change has been seen to result in marked impacts on forest ecosystems such as accelerated tree mortality worldwide due to incidental hydraulic failure caused by intensified and more frequent occurrence of extreme drought and heat-waves. However, it is well understood how the tree hydrological strategies would adjust to environmental variability brough about by climate changes. Here we investigated the hydraulic adjustment as a mechanism of acclimation to different climate conditions along an altitudinal gradient in Faxon fir ( Abies fargesii var. faxoniana ) ― a tree species that plays a key role in conservation of wildlife and maintenance of ecosystem services in subalpine forests. The hydraulic traits and selective morphological and physiological variables were measured seasonally along an altitudinal gradient from 2,800 to 3,600 ​m a.s.l. We found that the native percentage loss of conductivity (PLC) increased with altitude across the seasonal measurements . Both the native sapwood-specific hydraulic conductivity ( K s ) and native leaf-specific hydraulic conductivity ( K l ) significantly decreased with altitude for measurements in July and October, coinciding with the timing for peak growth and pre-dormancy, respectively. The morphological traits varied toward more conservative tree hydrological strategies with increases in altitude, exhibiting trade-offs with hydraulic traits. The total non-structural carbohydrates in both needle (NSC Needle ) and branch (NSC Branch ) as well as photosynthetic capacity of current-year leaves played variable roles in maintaining the integrity of the hydraulic functioning and shaping the hydraulic adjustment under prevailing environmental conditions. Our findings indicate that Faxon fir possesses some degree of hydraulic adaptability to water limitation imposed by climate fluctuations in subalpine region through morphological and physiological modifications.

Variation in Xylem Hydraulic Structure and Function of Two Mangrove Species across a Latitudinal Gradient in Eastern Australia

Mangroves growing in tropical and subtropical intertidal zones face challenges from warming and altered rainfall patterns associated with global climate change. Intraspecific variation in hydraulic traits may allow a mangrove species to acclimate to novel climatic conditions, yet little is known regarding the potential for adaptive plasticity in these traits. In this study, we aimed to quantify the variation in plant hydraulic traits of two widespread mangrove species growing across a latitudinal gradient. We investigated the xylem hydraulic structure and function of Avicennia marina and Aegiceras corniculatum, across three sites spanning a latitudinal gradient of 17.45° in eastern Australia. We found that both species were highly resistant to xylem embolism and that there was significant intraspecific variation in hydraulic traits between sites. The highest embolism resistance and sapwood-specific hydraulic conductivity (KS) were found at the lowest latitude site that had the highest mean annual temperature and precipitation. A. marina showed no differences in vessel size and density among sites. It has other special features such as successive cambia enhancing its ability to adapt to a large environmental gradient. In contrast, A. corniculatum showed higher vessel densities at lower latitudes. There was a significant and positive correlation (R2 = 0.72, p &lt; 0.05) between KS and embolism resistance across species and sites, suggesting the absence of a tradeoff between hydraulic efficiency and safety. Both embolism resistance and KS were negatively correlated with wood density but positively with vessel wall reinforcement. This study reveals that these two widespread mangrove species were adapted to warmer climates by enhancing both hydraulic efficiency and safety.

Stem hydraulic architecture of Malus sieversii in degraded wild fruit forest in Ili valley, China

To reveal mechanisms underlying the dieback of Malus sieversii in degraded wild fruit forest in Ili valley of China, we compared the differences in stem hydraulic architecture, water transport effectiveness and safety, as well as their influencing factors among three growth classes of dead branches ratios, including Class Ⅰ (<20%), Class Ⅱ (40%-60%) and Class Ⅲ (>80%), respectively. With the increases in degradation degree of Xinjiang wild fruit forest, sapwood-specific hydraulic conductivity and leaf-specific hydraulic conductivity decreased, without significant differences among the three growth classes. Branch embolism at dusk and hydraulic safety significantly increased. The xylem water potential at 50% loss of hydraulic conductivity was -1.87, -1.35 and -0.53 MPa for Class Ⅰ, Ⅱ and Ⅲ, respectively. Predawn and midday leaf water potential and the hydraulic safety margin exhibited an order of Ⅰ>Ⅱ>Ⅲ. Xylem anatomical cha-racteristics and branch and leaf traits related to hydraulics were significantly different among the three growth classes. Results from correlation analysis revealed a weak tradeoff between xylem-specific hydraulic efficiency and xylem safety of M. sieversii. Stem hydraulic architecture of M. sieversii altered with the decline of Xinjiang wild apple forest. With increasing degrees of degradation, the severity of xylem embolisms aggravated, resistance to cavitation embolisms reduced, and the risk of water imbalance increased.

Shade tree species affect gas exchange and hydraulic conductivity of cacao cultivars in an agroforestry system.

Shade tolerance is a widespread strategy of rainforest understory plants. Many understory species have green young stems that may assimilate CO2 and contribute to whole-plant carbon balance. Cacao commonly grows in the shaded understory and recent emphasis has been placed on diversifying the types of trees used to shade cacao plants to achieve additional ecosystem services. We studied three agricultural cacao cultivars growing in the shade of four timber species (Cedrela odorata L., Cordia thaisiana Agostini, Swietenia macrophylla King and Tabebuia rosea (Bertol) A.D.C.) in an agroforestry system to (i) evaluate the timber species for their effect on the physiological performance of three cacao cultivars; (ii) assess the role of green stems on the carbon economy of cacao; and (iii) examine coordination between stem hydraulic conductivity and stem photosynthesis in cacao. Green young stem photosynthetic CO2 assimilation rate was positive and double leaf CO2 assimilation rate, indicating a positive contribution of green stems to the carbon economy of cacao; however, green stem area is smaller than leaf area and its relative contribution is low. Timber species showed a significant effect on leaf gas exchange traits and on stomatal conductance of cacao, and stem water-use efficiency varied among cultivars. There were no significant differences in leaf-specific hydraulic conductivity among cacao cultivars, but sapwood-specific hydraulic conductivity varied significantly among cultivars and there was an interactive effect of cacao cultivar × timber species. Hydraulic efficiency was coordinated with stem-stomatal conductance, but not with leaf-stomatal conductance or any measure of photosynthesis. We conclude that different shade regimes determined by timber species and the interaction with cacao cultivar had an important effect on most of the physiological traits and growth variables of three cacao cultivars growing in an agroforestry system. Results suggested that C. odorata is the best timber species to provide partial shade for cacao cultivars in the Barlovento region in Venezuela, regardless of cultivar origin.

The Accumulated Response of Deciduous Liquidambar formosana Hance and Evergreen Cyclobalanopsis glauca Thunb. Seedlings to Simulated Nitrogen Additions.

Nitrogen depositions in the Yangtze River Delta have is thought to shift the coexistence of mixed evergreen and deciduous species. In this study, the seedlings of the dominant evergreen species Cyclobalanopsis glauca Thunb. and the deciduous species Liquidambar formosana Hance from the Yangtze River Delta were chosen to test their responses to simulated N additions using an ecophysiological approach. N was added to the tree canopy at rates of 0 (CK), 25 kg N ha−1 year−1 (N25), and 50 kg N ha−1 year−1 (N50). The leaf N content per mass (N m, by 44.03 and 49.46%) and total leaf chlorophyll content (Chl, by 72.15 and 63.63%) were enhanced for both species, and C. glauca but not L. formosana tended to allocate more N to Chl per leaf area (with a higher slope). The enhanced N availability and Chl promoted the apparent quantum yield (AQY) significantly by 15.38 and 43.90% for L. formosana and C. glauca, respectively. Hydraulically, the increase in sapwood density (ρ) for L. formosana was almost double that of C. glauca. Synchronous improved sapwood specific hydraulic conductivity (K S, by 37.5%) for C. glauca induced a significant reduction in stomatal conductance (g s) (p < 0.05) in the N50 treatments, which is in contrast to the weak varied g s accompanied by a 59.49% increase in K S for L. formosana. As a result, the elevated maximum photosynthesis (A max) of 12.19% for L. formosana in combination with the increase in the total leaf area (indicated by a 37.82% increase in the leaf area ratio-leaf area divided by total aboveground biomass) ultimately yielded a 34.34% enhancement of total biomass. In contrast, the A max and total biomass were weakly promoted for C. glauca. The reason for these distinct responses may be attributed to the lower water potential at 50% of conductivity lost (P 50) for C. glauca, which enables higher hydraulic safety at the cost of a weak increase in Amax due to the stomatal limitation in response to elevated N availability. Altogether, our results indicate that the deciduous L. formosana would be more susceptible to elevated N availability even if both species received similar N allocation.

The adjustment of Prosopis tamarugo hydraulic architecture traits has a homeostatic effect over its performance under descent of phreatic level in the Atacama Desert

Prosopis tamarugo undergoes several modifications of its hydraulic architecture, which has a homeostatic effect on its performance under descent of phreatic level. Groundwater extraction for mining has generated a large area of descent of phreatic level in the Llamara Salt Flat, Atacama Desert, where Prosopis tamarugo lives as a strict phreatophyte. We study different hydraulic architecture traits of P. tamarugo after 12 years of exposure to a descent of phreatic level of ~ 11 m to assess its acclimation capacity. Under this condition, tree size was reduced making the structure of trees shrubbier. At branch level, leaf shedding increased Huber Value and theoretical leaf-specific hydraulic conductivity. At vascular tissue level, vessel composition decreased significantly due to an increase in vessel density and a slight decrease in mean vessel area. This meant that there were no differences of theoretical sapwood-specific hydraulic conductivity and wood density between conditions of descent of phreatic level. Although the theoretical hydraulic efficiency is high under descent of phreatic level, a lower midday water potential indicates other possible factors that may affect its performance (e.g., microclimate, daily loss of conductivity and a high root resistances). The above is consistent with a higher leaf mass per area of the remaining leaves, but no differences were observed in stomatal conductance, leaf 13C isotopic composition and predawn water potential between descent condition. While the adjustment of hydraulic architecture traits has a homeostatic effect on P. tamarugo performance, it is necessary to evaluate the limits of its resistance to contribute to its conservation in a context of intervention of its habit, mainly due to demographic growth and non-metallic mining.

Water transport from stem to stomata: the coordination of hydraulic and gas exchange traits across 33 subtropical woody species.

Coordination between sapwood-specific hydraulic conductivity (Ks) and stomatal conductance (gs) has been identified in previous studies; however, coordination between leaf hydraulic conductance (Kleaf) and gs, as well as between Kleaf and Ks is not always consistent. This suggests that there is a need to improve our understanding of the coordination among hydraulic and gas exchange traits. In this study, hydraulic traits (e.g., Ks and Kleaf) and gas exchange traits, including gs, transpiration (E) and net CO2 assimilation (Aarea), were measured across 33 co-occurring subtropical woody species. Kleaf was divided into two components: leaf hydraulic conductance inside the xylem (Kleaf-x) and outside the xylem (Kleaf-ox). We found that both Kleaf-x and Kleaf-ox were coordinated with gs and E, but the correlations between Kleaf-ox and gs (or E) were substantially weaker, and that Ks was coordinated with Kleaf-x, but not with Kleaf-ox. In addition, we found that Ks, Kleaf-x and Kleaf-ox together explained 63% of the variation in gs and 42% of the variation in Aarea across species, with Ks contributing the largest proportion of explanatory power, whereas Kleaf-ox contributed the least explanatory power. Our results demonstrate that the coordination between leaf water transport and gas exchange, as well as the hydraulic linkage between leaf and stem, were weakened by Kleaf-ox. This highlights the possibility that water transport efficiencies of stem and leaf xylem, rather than that of leaf tissues outside the xylem, are important determinants of stomatal conductance and photosynthetic capacity across species.

The effects of intervessel pit characteristics on xylem hydraulic efficiency and photosynthesis in hemiepiphytic and non-hemiepiphytic Ficus species.

Xylem vulnerability to cavitation and hydraulic efficiency are directly linked to fine-scale bordered pit features in water-conducting cells of vascular plants. However, it is unclear how pit characteristics influence water transport and carbon economy in tropical species. The primary aim of this study was to evaluate functional implications of changes in pit characteristics for water relations and photosynthetic traits in tropical Ficus species with different growth forms (i.e. hemiepiphytic and non-hemiepiphytic) grown under common conditions. Intervessel pit characteristics were measured using scanning electron microscopy in five hemiepiphytic and five non-hemiepiphytic Ficus species to determine whether these traits were related to hydraulics, leaf photosynthesis, stomatal conductance and wood density. Ficus species varied greatly in intervessel pit structure, hydraulic conductivity and leaf physiology, and clear differences were observed between the two growth forms. The area and diameter of pit aperture were negatively correlated with sapwood-specific hydraulic conductivity, mass-based net assimilation rate, stomatal conductance (gs ), intercellular CO2 concentration (Ci ) and the petiole vessel lumen diameters (Dv ), but positively correlated with wood density. Pit morphology was only negatively correlated with sapwood- and leaf-specific hydraulic conductivity and Dv . Pit density was positively correlated with gs , Ci and Dv , but negatively with intrinsic leaf water-use efficiency. Pit and pit aperture shape were not significantly correlated with any of the physiological traits. These findings indicate a significant role of pit characteristics in xylem water transport, carbon assimilation and ecophysiological adaptation of Ficus species in tropical rain forests.

Conifers but not angiosperms exhibit vulnerability segmentation between leaves and branches in a temperate forest

Vulnerability segmentation (VS), an important mechanism for protecting plants from drought, hypothesizes that the distal organs of a plant should be more susceptible to embolism than the basal organs. However, experimental studies testing the VS hypothesis for trees are limited and have reached inconsistent conclusions. Here, we tested the VS hypothesis with three angiosperms and four conifers co-existing in a temperate forest in northeastern China. The results showed that the difference in vulnerability to cavitation between leaves and branches (P50leaf-branch) was positive for the conifers but negative for the angiosperms, implying that the conifers rather than the angiosperms exhibited VS. The conifers had lower leaf hydraulic safety margins and more embolism-resistant branches than the angiosperms. Although the angiosperms did not display VS, they took a hydraulic compensatory strategy (e.g., great leaf and branch hydraulic conductivities) to maintain the water supply of their leaves. In addition, we found a significant trade-off between the sapwood-specific hydraulic conductivity (KSS) and xylem pressure inducing 50% loss of hydraulic conductivity (P50branch) across all species. Both KSS and P50branch increased with the area-based light-saturated photosynthetic rate (Aarea), suggesting that increased embolism resistance of branches comes at the cost of reduced hydraulic efficiency, which in turn constrains the photosynthesis. Aarea was negatively correlated with P50leaf-branch, further indicating that the conifers had strong VS and were associated with a conservative strategy. Conversely, the angiosperms displayed an acquisitive strategy, tending to have higher Aarea, leaf and branch hydraulic conductivities, but lower embolism resistance. These differentiations in the functional traits between the angiosperms and conifers provide potential mechanisms for their co-existence in this temperate forest community.

Changes in structure and physiological functioning due to experimentally enhanced precipitation seasonality in a widespread shrub species

Semi-arid shrub steppe occupies a vast geographic range that is characterized in part by distinct seasonal patterns in precipitation. Few studies have evaluated how variability in both the amount and timing of precipitation affect the structure and physiology of shrubs in these systems. We quantified changes in foliar crown parameters, xylem anatomy, gas exchange, and hydraulic transport capacity in deep-rooted Artemisia tridentata shrubs following 20 years of experimental manipulations in amount and seasonal timing of precipitation. We hypothesized that shrub growth (total leaf area per shrub and cover of shrub community), hydraulic transport efficiency, and gas exchange would increase in shrubs in irrigated plots compared to non-irrigated control plots, especially for irrigation applied in winter rather than summer. We also predicted similar changes in xylem anatomy (ring width, vessel size and frequency). Most treatment responses entailed changes in plant structure, and were generally consistent with our hypotheses: total-shrub leaf area, shrub basal area, canopy cover, and maximum sapwood-specific branch hydraulic conductivity were more than 2× greater in shrubs in winter-irrigated compared to control plots, while summer irrigation had few effects on these variables. Irrigation in either season did not affect xylem vessel size, but did increase xylem ring width by ~ 2 × and decreased xylem vessel frequency by about half. Anatomical, morphological, and stand-level abundance of A. tridentata appeared much more responsive to irrigation than state changes in gas exchange, particularly when the extra water is received during winter. Thus, it appears for sagebrush that seasonal timing is at least as important as the amount of precipitation, and that responses to changes in precipitation timing occur through changes in carbon allocation more so than changes in leaf-level carbon gain.

Divergent Hydraulic Safety Strategies in Three Co-occurring Anacardiaceae Tree Species in a Chinese Savanna.

Vulnerability segmentation, the condition under which plant leaves are more vulnerable to drought-induced cavitation than stems, may act as a “safety valve” to protect stems from hydraulic failure. Evergreen, winter-deciduous, and drought-deciduous tree species co-occur in tropical savannas, but there have been no direct studies on the role of vulnerability segmentation and stomatal regulation in maintaining hydraulic safety in trees with these three leaf phenologies. To this end, we selected three Anacardiaceae tree species co-occurring in a Chinese savanna, evergreen Pistacia weinmanniifolia, drought-deciduous Terminthia paniculata, and winter-deciduous Lannea coromandelica, to study inter-species differentiation in leaf and stem hydraulic safety. We found that the two deciduous species had significantly higher sapwood-specific hydraulic conductivity and leaf-specific hydraulic conductance than the evergreen species. Moreover, two deciduous species were more vulnerable to stem cavitation than the evergreen species, although both drought-deciduous species and evergreen species had drought-resistance leaves. The evergreen species maintained a wide hydraulic safety margin (HSM) in stems and leaves; which was achieved by embolism resistance of both stems and leaves and isohydric stomatal control. Both deciduous species had limited HSMs in stems and leaves, being isohydric in the winter-deciduous species and anisohydric in drought-deciduous species. The difference in water potential at 50% loss of hydraulic conductivity between the leaves and the terminal stems (P50leaf−stem) was positive in P. weinmanniifolia and L. coromandelica, whereas, T. paniculata exhibited a lack of vulnerability segmentation. In addition, differences in hydraulic architecture were found to be closely related to other structural traits, i.e., leaf mass per area, wood density, and sapwood anatomy. Overall, the winter-deciduous species exhibits a drought-avoidance strategy that maintains the hydraulic safety of the more carbon-costly stems by sacrificing cheaper and more vulnerable leaves, while the evergreen species exhibits a hydraulic strategy of drought tolerance with strong stomatal regulation. In contrast, the drought-deciduous species lacks vulnerability segmentation and sheds leaves at the expense of top shoots during peak drought. This study demonstrates that even sympatric tree species that differ in leaf phenology can exhibit divergent adaptive hydraulic safety strategies.

Interspecific variation in branch and leaf traits among three Syzygium tree species from different successional tropical forests.

Plant functional traits are closely associated with plant habitats. In this study, we investigated the interspecific variations in stem and leaf hydraulics, xylem and leaf anatomy, gas-exchange rates and leaf pressure-volume relationships among three Syzygium tree species in early, mid- and late successional tropical forests. The objective was to understand the response and adaptation of congeneric species, in terms of branch and leaf functional traits, to different environments. A consistent pattern of decline with succession was evident in leaf and sapwood specific hydraulic conductivity (ks), maximum leaf hydraulic conductance (Kleaf), and photosynthetic rates for the three Syzygium species. Variations of ks and Kleaf were correlated with changes in vessel anatomy (i.e. vessel density and diameter) and leaf flux-related structure (i.e. stomatal pore index and vein density) respectively. However, specific leaf area and leaf to sapwood area ratio did not significantly differ among the three species. In addition, the mid-successional species had the lowest values of leaf water potential at full turgor and turgor loss point and 50% loss of Kleaf, but the greatest value of xylem water potential at 50% loss of ks. Our results demonstrate that leaf and branch traits associated with photosynthesis and/or hydraulic conductance, rather than those associated with drought tolerance, are the key factors underlying the response and adaptation of the three Syzygium tree species along the tropical forest succession.

Functional coordination between branch hydraulic properties and leaf functional traits in miombo woodlands: implications for water stress management and species habitat preference

We investigated functional coordination between branch hydraulic properties and leaf functional traits among nine miombo woodlands canopy tree species differing in habitat preference and phenology. Specifically, we were seeking to answer the question: are branch hydraulic properties coordinated with leaf functional traits linked to plant drought tolerance in seasonally dry tropical forests and what are the implications for species habitat preference? The hydraulic properties investigated in this study were stem area specific hydraulic conductivity (KS), Huber value (Hv), and xylem cavitation vulnerability (Ψ50). The leaf functional traits measured were specific leaf area (SLA), leaf dry matter content (LDMC), and mean leaf area (MLA). Generalists displayed significantly (P < 0.05) higher cavitation resistance (Ψ50) and SLA, but lower sapwood specific hydraulic conductivity (KS), leaf specific conductivity (KL), MLA, and LDMC than mesic specialists. Although MLA was uncorrelated with Ψ50, we found significant (P < 0.05) positive and negative correlations between plant hydraulic properties and leaf functional traits linked to plant drought tolerance ability, indicating that the interactions between branch hydraulics and leaf functional traits related to plant drought tolerance ability may influence tree species habitat preference in water-limited ecosystems.

Change in hydraulic properties and leaf traits in a tall rainforest tree species subjected to long-term throughfall exclusion in the perhumid tropics

Abstract. A large-scale replicated throughfall exclusion experiment was conducted in a pre-montane perhumid rainforest in Sulawesi (Indonesia) exposing the trees for two years to pronounced soil desiccation. The lack of regularly occurring dry periods and shallow rooting patterns distinguish this experiment from similar experiments conducted in the Amazonian rainforest. We tested the hypotheses that a tree's sun canopy is more affected by soil drought than its shade crown, making tall trees particularly vulnerable even under a perhumid climate, and that extended drought periods stimulate an acclimation in the hydraulic system of the sun canopy. In the abundant and tall tree species Castanopsis acuminatissima (Fagaceae), we compared 31 morphological, anatomical, hydraulic and chemical variables of leaves, branches and the stem together with stem diameter growth between drought and control plots. There was no evidence of canopy dieback. However, the drought treatment led to a 30 % reduction in sapwood-specific hydraulic conductivity of sun canopy branches, possibly caused by the formation of smaller vessels and/or vessel filling by tyloses. Drought caused an increase in leaf size, but a decrease in leaf number, and a reduction in foliar calcium content. The δ13C and δ18O signatures of sun canopy leaves gave no indication of a permanent down-regulation of stomatal conductance during the drought, indicating that pre-senescent leaf shedding may have improved the water status of the remaining leaves. Annual stem diameter growth decreased during the drought, while the density of wood in the recently produced xylem increased in both the stem and sun canopy branches (marginally significant). The sun canopy showed a more pronounced drought response than the shade crown indicating that tall trees with a large sun canopy are more vulnerable to drought stress. We conclude that the extended drought prompted a number of medium- to long-term responses in the leaves, branches and the trunk, which may have reduced drought susceptibility. However, unlike a natural drought, our drought simulation experiment was carried out under conditions of high humidity, which may have dampened drought induced damages.

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.