Bark and sapwood water storage and the atypical pattern of recharge and discharge of water reservoirs indicate low vulnerability to drought in Araucaria araucana.

Stored water in inner tissues influences the plant water economy, which might be particularly relevant for trees facing increasing dry conditions due to climate change. We studied the water storage in the inner bark and the sapwood of Araucaria araucana (Molina) K. Koch. This species has an extremely thick inner bark and thus it can be used as a model system to assess the impact of internal water storage on plant water balance. Specifically, we analyzed the water circulation pathways in and out of the elastic water storages by using simultaneously frequency domain moisture sensors and dendrometers inserted in the inner bark and in the sapwood, and sap flow determinations during the dry season. The daily patterns of water content and expansion and contraction of the stem tissues were similar to the sap flow pattern. The whole-stem water content and diameter increased in the morning and decreased in the afternoon, contrary to the typical pattern observed in most tree species. An osmotic gradient favoring the water influx from sapwood to inner bark was observed in the morning. There were no lags in the onset of sap flow between different stem heights at the time that recharge of reservoirs occurred. Sap flow at 6 m height was higher than basal sap flow in the afternoon, when the sapwood water content started to decline followed by the water content of the inner bark. Inner bark and sapwood contributed 5-11% to total daily transpiration, allowing the maintenance of high water potentials in the dry season. Our results suggest that the stored water in the stems, the atypical dynamic of recharge and discharge of water from reservoirs and the high tissue capacitance may make an important contribution to the survival of A. araucana during drought periods by maintaining the water balance.

Leaf surface traits contributing to wettability, water interception and uptake of above-ground water sources in shrubs of Patagonian arid ecosystems.

BACKGROUND AND AIMS: The ecohydrological significance of leaf wetting due to atmospheric water in arid and semiarid ecosystems is not well understood. In these environments, the inputs of precipitation or dew formation resulting in leaf wetting have positive effects on plant functioning. However, its impact on plant water relations may depend on the degree of leaf surface wettability. In this study we evaluated leaf wettability and other leaf traits and its effects on foliar water uptake and canopy interception in plant species of a Patagonian steppe. We also studied how leaf traits affecting wettability vary seasonally from growing to dry season. METHODS: Contact angle of a water droplet with the leaf surface, water adhesion, droplet retention angle, stomatal density, cuticular conductance, canopy interception and maximum foliar water uptake were determined in six dominant shrub species. KEY RESULTS: All species increased leaf wettability during the dry season and most species were considered highly wettable. The leaf surface had very high capacity to store and retain water. We found a negative correlation between foliar water uptake and leaf hydrophilia. CONCLUSIONS: Despite the diversity of life forms, including cushion shrubs and tall shrubs, as well as phenological variability, all species converged in similar seasonal changes in leaf traits that favour wettability. Intercepted water by crowns and the extremely high capacity of retention of droplets on leaf surfaces can have a significant impact on eco-hydrological process in water limited ecosystems where most of water sources during the growing and the dry season may be small rainfall events or dew, which do not always increase soil water availability.

Low-temperature acclimation and legacy effects of summer water deficits in olive freezing resistance.

Low temperatures and drought are the main environmental factors affecting plant growth and productivity across most of the terrestrial biomes. The objective of this study was to analyze the effects of water deficits before the onset of low temperatures in winter to enhance freezing resistance in olive trees. The study was carried out near the coast of Chubut, Argentina. Plants of five olive cultivars were grown outdoor in pots and exposed to different water deficit treatments. We assessed leaf water relations, ice nucleation temperature (INT), cell damage (LT50), plant growth and leaf nitrogen content during summer and winter in all cultivars and across water deficit treatments. Leaf INT and LT50 decreased significantly from summer to winter within each cultivar and between treatments. We observed a trade-off between resources allocation to freezing resistance and vegetative growth, such that an improvement in resistance to sub-zero temperatures was associated with lower growth in tree height. Water deficit applied during summer increased the amount of osmotically active solutes and decreased the leaf water potentials. This type of legacy effect persists during the winter after the water deficit even when treatment was removed by natural rainfalls.

Drought and frost resistance vary between evergreen and deciduous Atlantic Forest canopy trees.

Frost and drought are key stress factors limiting the growth and distribution of tree species. Resistance to stress involves energy costs that may result in trade-offs between different functional traits. Structures or mechanisms that can help to withstand stress imply differences in the carbon economy of the species. Although adaptive responses to frost and drought resistance are usually of a similar nature, they are rarely assessed simultaneously. We investigated these resistance mechanisms in 10 canopy tree species coexisting in the semi-deciduous subtropical forests of northern Argentina. We measured leaf lifespan, anatomical, photosynthetic and water relations traits and performed a thermal analysis in leaves to determined ice nucleation and tissue damage temperatures. Our results showed that evergreen and deciduous species have different adaptive responses to cope with freezing temperatures and water deficits. Evergreen species exhibited cold tolerance, while deciduous species were more resistant to hydraulic dysfunction and showed greater water transport efficiency. Further research is needed to elucidate resistance strategies to stress factors at the whole tree- and stand level, and possible links with hydraulic safety and efficiency among different phenological groups. This will allow us to predict the responses of subtropical forest species to changes in environmental conditions under climate change scenarios.

Foliar water uptake in arid ecosystems: seasonal variability and ecophysiological consequences.

Foliar water uptake (FWU) has been reported for different species across several ecosystems types. However, little attention has been given to arid ecosystems, where FWU during dew formation or small rain events could ameliorate water deficits. FWU and their effects on leaf water potential (ΨLeaf) were evaluated in grasses and shrubs exploring different soil water sources in a Patagonian steppe. Also, seasonal variability in FWU and the role of cell wall elasticity in determining the effects on ΨLeaf were assessed. Eleven small rain events (< 8 mm) and 45 days with dew formation were recorded during the study period. All species exhibited FWU after experimental wetting. There was a large variability in FWU across species, from 0.04 mmol m-2 s-1 in species with deep roots to 0.75 mmol m-2 s-1 in species with shallow roots. Species-specific mean FWU rates were positively correlated with mean transpiration rates. The increase in ΨLeaf after leaf wetting varied between 0.65 MPa and 1.67 MPa across species and seasons. The effects of FWU on ΨLeaf were inversely correlated with cell wall elasticity. FWU integrated over both seasons varied between 28 mol m-2 in species with deep roots to 361 mol m-2 in species with shallow roots. Taking into account the percentage of coverage of each species, accumulated FWU represented 1.6% of the total annual transpiration of grasses and shrubs in this ecosystem. Despite this low FWU integrated over time compared to transpiration, wetting leaves surfaces can help to avoid larger water deficit during the dry season.

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.

Hydraulic traits are coordinated with maximum plant height at the global scale.

Water must be transported long distances in tall plants, resulting in increasing hydraulic resistance, which may place limitations on the maximum plant height (H max) in a given habitat. However, the coordination of hydraulic traits with H max and habitat aridity remains poorly understood. To explore whether H max modifies the trade-off between hydraulic efficiency and safety or how water availability might influence the relationship between H max and other hydraulic traits, we compiled a dataset including H max and 11 hydraulic traits for 1281 woody species from 369 sites worldwide. We found that taller species from wet habitats exhibited greater xylem efficiency and lower hydraulic safety, wider conduits, lower conduit density, and lower sapwood density, which were all associated with habitat water availability. Plant height and hydraulic functioning appear to represent a single, coordinated axis of variation, aligned primarily with water availability, thus suggesting an important role for this axis in species sorting processes.

Correction to: Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees.

The original version of this article unfortunately contained a mistake. The Electronic supplementary material (ESM) was accompanying this article by mistake.

Functional relationships between hydraulic traits and the timing of diurnal depression of photosynthesis.

The hydraulic coordination along the water transport pathway helps trees provide adequate water supply to the canopy, ensuring that water deficits are minimized and that stomata remain open for CO2 uptake. We evaluated the stem and leaf hydraulic coordination and the linkages between hydraulic traits and the timing of diurnal depression of photosynthesis across seven evergreen tree species in the southern Andes. There was a positive correlation between stem hydraulic conductivity (ks ) and leaf hydraulic conductance (KLeaf ) across species. All species had similar maximum photosynthetic rates (Amax ). The species with higher ks and KLeaf attained Amax in the morning, whereas the species with lower ks and KLeaf exhibited their Amax in the early afternoon concurrently with turgor loss. These latter species had very negative leaf water potentials, but far from the pressure at which the 88% of leaf hydraulic conductance is lost. Our results suggest that diurnal gas exchange dynamics may be determined by leaf hydraulic vulnerability such that a species more vulnerable to drought restrict water loss and carbon assimilation earlier than species less vulnerable. However, under stronger drought, species with earlier CO2 uptake depression may increase the risk of hydraulic failure, as their safety margins are relatively narrow.

Leaf phenology and water-use patterns of canopy trees in Northern Argentinean subtropical forests.

Tree physiological processes are affected not only by environmental conditions, but also by phenological leaf stages. During foliar expansion, rapid changes occur, such as the activation of metabolic processes that encompass a hydraulic link between xylem and phloem pathways at a whole-tree level. Daily and seasonal changes in stem diameter may reveal different temporal dynamics of water use and recharge in tree reservoirs. Foliar phenological patterns were studied in relation to stem dimensional changes in 10 canopy tree species with different phenological patterns (three deciduous, three brevideciduous and four evergreen species). Additionally, we assessed (i) daily sap flow fluctuations in branch and main stem, (ii) diurnal changes in sapwood volumetric water content and (iii) stem radius variations during leafless, expanding and mature leaves periods in three of the 10 tree species (two deciduous and one brevideciduous). During the leaf expansion phase, the diameter of trees decreased in all 10 species, with a larger impact on deciduous and brevideciduous species. For the subset of deciduous and brevideciduous species, the movement of long-distance water transport occurred first near the branches and then in the main stem during the leafless stage. Changes in stored water use and a decrease in the volumetric water content and the radius of the main stem during this period suggest that there is a contribution of water from internal stem reservoirs toward the construction of new leaves.

The cost of avoiding freezing in stems: trade-off between xylem resistance to cavitation and supercooling capacity in woody plants.

Stems and leaves of Olea europaea L. (olive) avoid freezing damage by substantial supercooling during the winter season. Physiological changes during acclimation to low temperatures were studied in five olive cultivars. Water relations and hydraulic traits, ice nucleation temperature (INT) and temperatures resulting in 50% damage (LT50) were determined. All cultivars showed a gradual decrease in INT and LT50 from the dry and warm summer to the wet and cold winter in Patagonia, Argentina. During acclimation to low temperatures there was an increase in leaf cell wall rigidity and stomatal conductance (gs), as well as a decrease in leaf apoplastic water content, leaf water potential (Ψ), sap flow and stem hydraulic conductivity (ks). More negative Ψ as a consequence of high gs and detrimental effects of low temperatures on root activity resulted in a substantial loss of ks due to embolism formation. Seasonal stem INT decrease from summer to winter was directly related to the xylem resistance to cavitation, determined by the loss of ks across cultivars. Thus the loss of freezable water in xylem vessels by embolisms increased stem supercooling capacity and delayed ice propagation from stems to the leaves. For the first time, a trade-off between xylem resistance to cavitation and stem and leaf supercooling capacity was observed in plants that avoid extracellular freezing by permanent supercooling. The substantial loss of hydraulic function in olive cultivar stems by embolism formation with their high repair costs are compensated by avoiding plant damage at very low subzero temperatures.

Microenvironment in the canopy rivals the host tree water status in controlling sap flow of a mistletoe species.

Mistletoes absorb water from the vascular system of their hosts and thus the water use of mistletoes can be influenced by the water status of their hosts besides abiotic environmental conditions; however, there is a lack of studies on the dynamics of mistletoe water utilization in relation to both types of controlling factors. By building a canopy platform at 20 m above the ground, we monitored the dynamic changes of sap flow of Viscum coloratum (Kom.) Nakai (Loranthaceae) in combination with continuous measurements of microclimatic variables and volumetric water content (VWC) of its host tree branch xylem. We found that the host tree VWC exhibited substantial fluctuations during sunny days but lower VWC of the host did not negatively affect the sap flow of V. coloratum. Hourly and daily mean transpiration rates (Esap) of V. coloratum calculated from sap flow measurements showed strong positive correlations with photosynthetic photon flux density (PPFD) and vapor pressure deficit (VPD) measured in close vicinity to the point of mistletoe attachment. The mean Esap of V. coloratum was substantially higher than that of their host during clear days (4.55 ± 0.54 vs 2.01 ± 0.15 kg m-2 day-1). Moreover, the mistletoe-to-host transpiration ratio was not constant but became increasingly larger with the increase of PPFD or VPD on both hourly and daily bases, suggesting a weaker control of water loss in the mistletoe in comparison to its host species. The strong dependence of mistletoe Esap on micrometeorological variables and its decoupling from the host tree xylem water status suggests that the development of dense tree canopy functions as a potential mechanism for the host trees in reducing the competitive water use of mistletoes. These findings have important implications for the interactions between mistletoe species and their host trees in temperate forests.

Freezing resistance in Patagonian woody shrubs: the role of cell wall elasticity and stem vessel size.

Freezing resistance through avoidance or tolerance of extracellular ice nucleation is important for plant survival in habitats with frequent subzero temperatures. However, the role of cell walls in leaf freezing resistance and the coordination between leaf and stem physiological processes under subzero temperatures are not well understood. We studied leaf and stem responses to freezing temperatures, leaf and stem supercooling, leaf bulk elastic modulus and stem xylem vessel size of six Patagonian shrub species from two sites (plateau and low elevation sites) with different elevation and minimum temperatures. Ice seeding was initiated in the stem and quickly spread to leaves, but two species from the plateau site had barriers against rapid spread of ice. Shrubs with xylem vessels smaller in diameter had greater stem supercooling capacity, i.e., ice nucleated at lower subzero temperatures. Only one species with the lowest ice nucleation temperature among all species studied exhibited freezing avoidance by substantial supercooling, while the rest were able to tolerate extracellular freezing from -11.3 to -20 °C. Leaves of species with more rigid cell walls (higher bulk elastic modulus) could survive freezing to lower subzero temperatures, suggesting that rigid cell walls potentially reduce the degree of physical injury to cell membranes during the extracellular freezing and/or thaw processes. In conclusion, our results reveal the temporal-spatial ice spreading pattern (from stem to leaves) in Patagonian shrubs, and indicate the role of xylem vessel size in determining supercooling capacity and the role of cell wall elasticity in determining leaf tolerance of extracellular ice formation.

Determinants of water circulation in a woody bamboo species: afternoon use and night-time recharge of culm water storage.

To understand water-use strategies of woody bamboo species, sap flux density (Fd) in the culms of a woody bamboo (Bambusa vulgaris Schrader ex Wendland) was monitored using the thermal dissipation method. The daytime and night-time Fd were analyzed in the dry and rainy seasons. Additionally, diurnal changes in root pressure, culm circumference, and stomatal conductance (gs) were investigated to characterize the mechanisms used to maintain diurnal water balance of woody bamboos. Both in the dry and rainy seasons, daytime Fd responded to vapor pressure deficit (VPD) in an exponential fashion, with a fast initial increase in Fd when VPD increased from 0 to 1 kPa. The Fd and gs started to increase very fast as light intensity and VPD increased in the morning, but they decreased sharply once the maximum value was achieved. The Fd response of this woody bamboo to VPD was much faster than that of representative trees and palms growing in the same study site, suggesting its fast sap flow and stomatal responses to changes in ambient environmental factors. The Fd in the lower and higher culm positions started to increase at the same time in the morning, but the Fd in the higher culm position was higher than that of the lower culm in the afternoon. Consistently, distinct decreases in its culm circumference in the afternoon were detected. Therefore, unlike trees, water storage of bamboo culms was not used for its transpiration in the morning but in the afternoon. Nocturnal sap flow of this woody bamboo was also detected and related to root pressure. We conclude that this bamboo has fast sap flow/stomatal responses to irradiance and evaporative demands, and it uses substantial water storage for transpiration in the afternoon, while root pressure appears to be a mechanism resulting in culm water storage recharge during the night.

Freezing avoidance by supercooling in Olea europaea cultivars: the role of apoplastic water, solute content and cell wall rigidity.

Plants can avoid freezing damage by preventing extracellular ice formation below the equilibrium freezing temperature (supercooling). We used Olea europaea cultivars to assess which traits contribute to avoid ice nucleation at sub-zero temperatures. Seasonal leaf water relations, non-structural carbohydrates, nitrogen and tissue damage and ice nucleation temperatures in different plant parts were determined in five cultivars growing in the Patagonian cold desert. Ice seeding in roots occurred at higher temperatures than in stems and leaves. Leaves of cold acclimated cultivars supercooled down to -13 °C, substantially lower than the minimum air temperatures observed in the study site. During winter, leaf ice nucleation and leaf freezing damage (LT50 ) occurred at similar temperatures, typical of plant tissues that supercool. Higher leaf density and cell wall rigidity were observed during winter, consistent with a substantial acclimation to sub-zero temperatures. Larger supercooling capacity and lower LT50 were observed in cold-acclimated cultivars with higher osmotically active solute content, higher tissue elastic adjustments and lower apoplastic water. Irreversible leaf damage was only observed in laboratory experiments at very low temperatures, but not in the field. A comparative analysis of closely related plants avoids phylogenetic independence bias in a comparative study of adaptations to survive low temperatures.

Extending the generality of leaf economic design principles in the cycads, an ancient lineage.

Cycads are the most ancient lineage of living seed plants, but the design of their leaves has received little study. We tested whether cycad leaves are governed by the same fundamental design principles previously established for ferns, conifers and angiosperms, and characterized the uniqueness of this relict lineage in foliar trait relationships. Leaf structure, photosynthesis, hydraulics and nutrient composition were studied in 33 cycad species from nine genera and three families growing in two botanical gardens. Cycads varied greatly in leaf structure and physiology. Similarly to other lineages, light-saturated photosynthetic rate per mass (Am ) was related negatively to leaf mass per area and positively to foliar concentrations of chlorophyll, nitrogen (N), phosphorus and iron, but unlike angiosperms, leaf photosynthetic rate was not associated with leaf hydraulic conductance. Cycads had lower photosynthetic N use efficiency and higher photosynthetic performance relative to hydraulic capacity compared with other lineages. These findings extend the relationships shown for foliar traits in angiosperms to the cycads. This functional convergence supports the modern synthetic understanding of leaf design, with common constraints operating across lineages, even as they highlight exceptional aspects of the biology of this key relict lineage.

Water storage dynamics in the main stem of subtropical tree species differing in wood density, growth rate and life history traits.

Wood biophysical properties and the dynamics of water storage discharge and refilling were studied in the trunk of canopy tree species with diverse life history and functional traits in subtropical forests of northeast Argentina. Multiple techniques assessing capacitance and storage capacity were used simultaneously to improve our understanding of the functional significance of internal water sources in trunks of large trees. Sapwood capacitances of 10 tree species were characterized using pressure-volume relationships of sapwood samples obtained from the trunk. Frequency domain reflectometry was used to continuously monitor the volumetric water content in the main stems. Simultaneous sap flow measurements on branches and at the base of the tree trunk, as well as diurnal variations in trunk contraction and expansion, were used as additional measures of stem water storage use and refilling dynamics. All evidence indicates that tree trunk internal water storage contributes from 6 to 28% of the daily water budget of large trees depending on the species. The contribution of stored water in stems of trees to total daily transpiration was greater for deciduous species, which exhibited higher capacitance and lower sapwood density. A linear relationship across species was observed between wood density and growth rates with the higher wood density species (mostly evergreen) associated with lower growth rates and the lower wood density species (mostly deciduous) associated with higher growth rates. The large sapwood capacitance in deciduous species may help to avoid catastrophic embolism in xylem conduits. This may be a low-cost adaptation to avoid water deficits during peak water use at midday and under temporary drought periods and will contribute to higher growth rates in deciduous tree species compared with evergreen ones. Large capacitance appears to have a central role in the rapid growth patterns of deciduous species facilitating rapid canopy access as these species are less shade tolerant than evergreen species.

Hydraulic architecture and photoinhibition influence spatial distribution of the arborescent palm Euterpe edulis in subtropical forests.

Physiological characteristics of saplings can be considered one of the most basic constraints on species distribution. The shade-tolerant arborescent palm Euterpe edulis Mart. is endemic to the Atlantic Forest of Argentina, Brazil and Paraguay. At a local scale, saplings of this species growing in native forests are absent in gaps. We tested the hypothesis whether sensitivity to photoinhibition or hydraulic architecture constrains the distribution of E. edulis saplings in sun-exposed forest environments. Using shade houses and field studies, we evaluated growth, survival, hydraulic traits and the susceptibility of Photosystem II to photoinhibition in E. edulis saplings under different growth irradiances. Survival rates in exposed sites in the field were very low (a median of 7%). All saplings exhibited photoinhibition when exposed to high radiation levels, but acclimation to a high radiation environment increased the rate of recovery. Petiole hydraulic conductivity was similar across treatments regardless of whether it was expressed per petiole cross-sectional area or per leaf area. At the plant level, investment in conductive tissues relative to leaf area (Huber values) increased with increasing irradiance. Under high irradiance conditions, plants experienced leaf water potentials close to the turgor-loss point, and leaf hydraulic conductance decreased by 79% relative to its maximum value. Euterpe edulis saplings were able to adjust their photosynthetic traits to different irradiance conditions, whereas hydraulic characteristics at the leaf level did not change across irradiance treatments. Our results indicate that uncoupling between water demand and supply to leaves apparently associated with high resistances to water flow at leaf insertion points, in addition to small stems with low water storage capacity, weak stomatal control and high vulnerability of leaves to hydraulic dysfunction, are the main ecophysiological constraints that prevent the growth and survival of E. edulis saplings in gaps in the native forest where native lianas and bamboos show aggressive growth.

Functional relationships between leaf hydraulics and leaf economic traits in response to nutrient addition in subtropical tree species.

Leaves can be both a hydraulic bottleneck and a safety valve against hydraulic catastrophic dysfunctions, and thus changes in traits related to water movement in leaves and associated costs may be critical for the success of plant growth. A 4-year fertilization experiment with nitrogen (N) and phosphorus (P) addition was done in a semideciduous Atlantic forest in northeastern Argentina. Saplings of five dominant canopy species were grown in similar gaps inside the forests (five control and five N + P addition plots). Leaf lifespan (LL), leaf mass per unit area (LMA), leaf and stem vulnerability to cavitation, leaf hydraulic conductance (K(leaf_area) and K(leaf_mass)) and leaf turgor loss point (TLP) were measured in the five species and in both treatments. Leaf lifespan tended to decrease with the addition of fertilizers, and LMA was significantly higher in plants with nutrient addition compared with individuals in control plots. The vulnerability to cavitation of leaves (P50(leaf)) either increased or decreased with the nutrient treatment depending on the species, but the average P50(leaf) did not change with nutrient addition. The P50(leaf) decreased linearly with increasing LMA and LL across species and treatments. These trade-offs have an important functional significance because more expensive (higher LMA) and less vulnerable leaves (lower P50(leaf)) are retained for a longer period of time. Osmotic potentials at TLP and at full turgor became more negative with decreasing P50(leaf) regardless of nutrient treatment. The K(leaf) on a mass basis was negatively correlated with LMA and LL, indicating that there is a carbon cost associated with increased water transport that is compensated by a longer LL. The vulnerability to cavitation of stems and leaves were similar, particularly in fertilized plants. Leaves in the species studied may not function as safety valves at low water potentials to protect the hydraulic pathway from water stress-induced cavitation. The lack of rainfall seasonality in the subtropical forest studied probably does not act as a selective pressure to enhance hydraulic segmentation between leaves and stems.

Extended leaf senescence promotes carbon gain and nutrient resorption: importance of maintaining winter photosynthesis in subtropical forests.

The relative advantages of being deciduous or evergreen in subtropical forests and the relationship between leaf phenology and nutrient resorption efficiency are not well understood. The most successful deciduous species (Lyonia ovalifolia) in an evergreen-dominated subtropical montane cloud forest in southwest (SW) China maintains red senescing leaves throughout much of the winter. The aim of this study was to investigate whether red senescing leaves of this species were able to assimilate carbon in winter, to infer the importance of maintaining a positive winter carbon balance in subtropical forests, and to test whether an extended leaf life span is associated with enhanced nutrient resorption and yearly carbon gain. The red senescing leaves of L. ovalifolia assimilated considerable carbon during part of the winter, resulting in a higher yearly carbon gain than co-occurring deciduous species. Its leaf N and P resorption efficiency was higher than for co-occurring non-anthocyanic deciduous species that dropped leaves in autumn, supporting the hypothesis that anthocyanin accumulation and/or extended leaf senescence help in nutrient resorption. Substantial winter carbon gain and efficient nutrient resorption may partially explain the success of L. ovalifolia versus that of the other deciduous species in this subtropical forest. The importance of maintaining a positive carbon balance for ecological success in this forest also provides indirect evidence for the dominance of evergreen species in the subtropical forests of SW China.