Long-term acclimatization of hydraulic properties, xylem conduit size, wall strength and cavitation resistance in Phaseolus vulgaris in response to different environmental effects

Phaseolus vulgaris grown under various environmental conditions was used to assess long-term acclimatization of xylem structural characteristics and hydraulic properties. Conduit diameter tended to be reduced and 'wood' density (of 'woody' stems) increased under low moisture ('dry'), increased soil porosity ('porous soil') and low phosphorus ('low P') treatments. Dry and low P had the largest percentage of small vessels. Dry, low light ('shade') and porous soil treatments decreased P50 (50% loss in conductivity) by 0.15-0.25 MPa (greater cavitation resistance) compared with 'controls'. By contrast, low P increased P50 by 0.30 MPa (less cavitation resistance) compared with porous soil (the control for low P). Changes in cavitation resistance were independent of conduit diameter. By contrast, changes in cavitation resistance were correlated with wood density for the control, dry and porous soil treatments, but did not appear to be a function of wood density for the shade and low P treatments. In a separate experiment comparing control and porous soil plants, stem hydraulic conductivity (kh), specific conductivity (ks), leaf specific conductivity (LSC), total pot water loss, plant biomass and leaf area were all greater for control plants compared to porous soil plants. Porous soil plants, however, demonstrated higher midday stomatal conductance to water vapour (gs), apparently because they experienced proportionally less midday xylem cavitation.     

Light environment,sapling architecture,and leaf display in six rain forest tree species

Architecture and leaf display were compared in saplings of six rain forest tree species differing in shade tolerance. Saplings were selected along the whole light range encountered in a forest environment. Species differed largely in realized height and crown expansion per unit support biomass, but this could not be related to differences in shade tolerance. The results demonstrate that there exist various solutions to an effective expansion of plant height and crown area. It is argued that choice of the study species and the ontogenetic trajectory regarded determine to a large extent the outcome of interspecific comparisons. No evidence was found that pioneers were characterized by a multilayered and shade tolerants by a monolayered leaf distribution. Yet, sun plants had a similar crown area, a deeper crown, and a higher leaf area index compared to shade plants and their leaves were more evenly distributed along the stem. This suggests that differences in leaf layering are found between plants growing in different light environments, rather than between species differing in shade tolerance.     

Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees

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

Leaf traits and herbivory rates of tropical tree species differing in successional status

We evaluated leaf characteristics and herbivory intensities for saplings of fifteen tropical tree species differing in their successional position. Eight leaf traits were selected, related to the costs of leaf display (specific leaf area [SLA], water content), photosynthesis (N and P concentration per unit mass), and herbivory defence (lignin concentration, C:N ratio). We hypothesised that species traits are shaped by variation in abiotic and biotic (herbivory) selection pressures along the successional gradient. All leaf traits varied with the successional position of the species. The SLA, water content and nutrient concentration decreased, and lignin concentration increased with the successional position. Herbivory damage (defined as the percentage of damage found at one moment in time) varied from 0.9-8.5% among the species, but was not related to their successional position. Herbivory damage appeared to be a poor estimator of the herbivory rate experienced by species, due to the confounding effect of leaf lifespan. Herbivory rate (defined as percentage leaf area removal per unit time) declined with the successional position of the species. Herbivory rate was only positively correlated to water content, and negatively correlated to lignin concentration, suggesting that herbivores select leaves based upon their digestibility rather than upon their nutritive value. Surprisingly, most species traits change linearly with succession, while resource availability (light, nutrients) declines exponentially with succession.     

The stem xylem of Patagonian shrubs operates far from the point of catastrophic dysfunction and is additionally protected from drought‐induced embolism by leaves and roots

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

Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

Functional groups of woody species in semi‐arid regions at low latitudes

In seasonally dry environments, woody species have different survival strategies. However, little is known about how environmental variables affect the phenology and water dynamics of these species. We aim to understand which variables initiate the vegetative phenophases of species in a tropical semiarid climate at 3°S latitude, where variation in photoperiod is minimal and rainfall is seasonal. We hypothesize that groups of species with similar vegetative phenologies, under similar conditions, are functionally similar in terms of water storage and use. We analyse the relationship between functional characteristics related to the acquisition and utilization of water, such as wood density, water storage capacity, water potential and vegetative phenology. The attributes were ordered by multidimensional scaling, and a multiple response permutation procedure was used to test consistency of the groups. Canonical correspondence analysis and Mantel tests were used to evaluate the phenophase response to environmental variables. We found four functional groups: (i) deciduous low wood density, which lose 75% of their leaves one month before the end of the rains; (ii) evergreen high wood density; (iii) early deciduous high wood density, which lose 75% of their leaves one month after the end of the rains; and (iv) late deciduous high wood density, which lose 75% of their leaves two months after the end of the rains. As expected, the vegetative phenodynamics of the deciduous high wood density group were mainly influenced by water availability. The evergreens did not show a correlation with rainfall. Only leaf shedding of the late deciduous, and the vegetative phenophases of the evergreens, responded to an increase in temperature and photoperiod. Bud‐break responded to increased photoperiod and soil humidity in the deciduous low wood density group. The foliar periodicity groups can be explained by the presence of species that differ mainly in their mechanisms of water acquisition and use.     

Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae)

Recent studies have demonstrated significant correlations between stem and leaf hydraulic properties when comparing across species within ecological communities. This implies that these traits are co-evolving, but there have been few studies addressing plant water relations within an explicitly evolutionary framework. This study tests for correlated evolution among a suite of plant water-use traits and environmental parameters in seven species of Pereskia (Cactaceae), using phylogenetically independent contrasts. There were significant evolutionary correlations between leaf-specific xylem hydraulic conductivity, Huber Value, leaf stomatal pore index, leaf venation density and leaf size, but none of these traits appeared to be correlated with environmental water availability; only two water relations traits - mid-day leaf water potentials and photosynthetic water use efficiency - correlated with estimates of moisture regime. In Pereskia, it appears that many stem and leaf hydraulic properties thought to be critical to whole-plant water use have not evolved in response to habitat shifts in water availability. This may be because of the extremely conservative stomatal behavior and particular rooting strategy demonstrated by all Pereskia species investigated. These results highlight the need for a lineage-based approach to understand the relative roles of functional traits in ecological adaptation.     

Morphological and physiological differentiation of seedlings between dry and wet habitats in a tropical dry forest

A common observation in tropical dry forests is the habitat preference of tree species along spatial soil water gradients. This pattern of habitat partitioning might be a result of species differentiation in their strategy for using water, along with competing functions such as maximizing water exploitation and tolerating soil water stress. We tested whether species from drier soil conditions exhibited a tolerance strategy compared with that of wet-habitat species. In a comparison of 12 morphophysiological traits in seedlings of 10 closely related dry and wet-habitat species pairs, we explored what trade-offs guide differentiation between habitats and species. Contrary to our expectations, dry-habitat species showed mostly traits associated with an exploitation strategy (higher carbon assimilation capacity, specific leaf area and leaf-specific conductivity and lower water-use efficiency). Strikingly, dry-habitat species tended to retain their leaves longer during drought. Additionally, we detected multiple strategies to live within each habitat, in part due to variation of strategies among lineages, as well as functional differentiation along the water storage capacity-stem density (xylem safety) trade-off. Our results suggest that fundamental trade-offs guide functional niche differentiation among tree species expressed both within and between soil water habitats in a tropical dry forest.     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

Wood structure and function change with maturity: Age of the vascular cambium is associated with xylem changes in current‐year growth

Xylem vessel structure changes as trees grow and mature. Age‐ and development‐related changes in xylem structure are likely related to changes in hydraulic function. We examined whether hydraulic function, including hydraulic conductivity and vulnerability to water‐stress‐induced xylem embolism, changed over the course of cambial development in the stems of 17 tree species. We compared current‐year growth of young (1–4 years), intermediate (2–7 years), and older (3–10 years) stems occurring in series along branches. Diffuse and ring porous species were examined, but nearly all species produced only diffuse porous xylem in the distal branches that were examined irrespective of their mature xylem porosity type. Vessel diameter and length increased with cambial age. Xylem became both more conductive and more cavitation resistant with cambial age. Ring porous species had longer and wider vessels and xylem that had higher conductivity and was more vulnerable to cavitation; however, these differences between porosity types were not present in young stem samples. Understanding plant hydraulic function and architecture requires the sampling of multiple‐aged tissues because plants may vary considerably in their xylem structural and functional traits throughout the plant body, even over relatively short distances and closely aged tissues.     

Co-ordination between xylem anatomy,plant architecture and leaf functional traits in response to abiotic and biotic drivers in a nurse cushion plant

Background and AimsPlants in dry Mediterranean mountains experience a double climatic stress: at low elevations, high temperatures coincide with water shortage during summer, while at high elevations temperature decreases and water availability increases. Cushion plants often act as nurses by improving the microclimate underneath their canopies, hosting beneficiary species that may reciprocally modify their benefactors’ microenvironment. We assess how the nurse cushion plant Arenaria tetraquetra subsp. amabilis adjusts its hydraulic system to face these complex abiotic and biotic constraints.MethodsWe evaluated intra-specific variation and co-ordination of stem xylem anatomy, leaf functional traits and plant architecture in response to elevation, aspect and the presence of beneficiary species in four A. tetraquetra subsp. amabilis populations in the Sierra Nevada mountains, southern Spain.Key ResultsXylem anatomical and plant architectural traits were the most responsive to environmental conditions, showing the highest mutual co-ordination. Cushions were more compact and had smaller, more isolated conductive vessels in the southern than in the northern aspect, which allow minimization of the negative impacts of more intense drought. Only vessel size, leaf mass per area and terminal branch length varied with elevation. Nurse cushions co-ordinated plant architecture and xylem traits, having higher canopy compactness, fewer leaves per branch and fewer, more isolated vessels than non-nurse cushions, which reflects the negative effects of beneficiary plants on nurse water status. In non-nurse cushions, plant architecture co-ordinated with leaf traits instead. The interacting effects of aspect and elevation on xylem traits showed that stress due to frost at high elevation constrained xylem anatomy in the north, whereas stress due to drought had a parallel effect in the south.ConclusionsTrait co-ordination was weaker under more demanding environmental conditions, which agrees with the hypothesis that trait independence allows plants to better optimize different functions, probably entailing higher adjustment potential against future environmental changes.     

Midday stomatal conductance is more related to stem rather than leaf water status in subtropical deciduous and evergreen broadleaf trees

Midday depressions in stomatal conductance (g_s) and photosynthesis are common in plants. The aim of this study was to understand the hydraulic determinants of midday g_s, the coordination between leaf and stem hydraulics and whether regulation of midday g_s differed between deciduous and evergreen broadleaf tree species in a subtropical cloud forest of Southwest (SW) China. We investigated leaf and stem hydraulics, midday leaf and stem water potentials, as well as midday g_s of co‐occurring deciduous and evergreen tree species. Midday g_s was correlated positively with midday stem water potential across both groups of species, but not with midday leaf water potential. Species with higher stem hydraulic conductivity and greater daily reliance on stem hydraulic capacitance were able to maintain higher stem water potential and higher g_s at midday. Deciduous species exhibited significantly higher stem hydraulic conductivity, greater reliance on stem capacitance, higher stem water potential and g_s at midday than evergreen species. Our results suggest that midday g_s is more associated with midday stem than with leaf water status, and that the functional significance of stomatal regulation in these broadleaf tree species is probably for preventing stem xylem dysfunction.