The complexity of biological disturbance agents, fuels heterogeneity, and fire in coniferous forests of the western United States.

Forest biological disturbance agents (BDAs) are insects, pathogens, and parasitic plants that affect tree decline, mortality, and forest ecosystems processes. BDAs are commonly thought to increase the likelihood and severity of fire by converting live standing trees to more flammable, dead and downed fuel. However, recent research indicates that BDAs do not necessarily increase, and can reduce, the likelihood or severity of fire. This has led to confusion regarding the role of BDAs in influencing fuels and fire in fire-prone western United States forests. Here, we review the existing literature on BDAs and their effects on fuels and fire in the western US and develop a conceptual framework to better understand the complex relationships between BDAs, fuels and fire. We ask: 1) What are the major BDA groups in western US forests that affect fuels? and 2) How do BDA-affected fuels influence fire risk and outcomes? The conceptual framework is rooted in the spatiotemporal aspects of BDA life histories, which drive forest impacts, fuel characteristics and if ignited, fire outcomes. Life histories vary among BDAs from episodic, landscape-scale outbreaks (bark beetles, defoliators), to chronic, localized disturbance effects (dwarf mistletoes, root rots). Generally, BDAs convert aboveground live biomass to dead biomass, decreasing canopy fuels and increasing surface fuels. However, the rate of conversion varies with time-since-event and among BDAs and forest types, resulting in a wide range of effects on the amount of dead fuels at any given time and place, which interacts with the structure and composition of the stand before and subsequent to BDA events. A major influence on fuels may be that BDAs have emerged as dominant agents of forest heterogeneity creation. Because BDAs play complex roles in fuels and fire heterogeneity across the western US which are further complicated by interactions with climate change, drought, and forest management (fire suppression), their impacts on fuels, fire and ecological consequences cannot be categorized simply as positive or negative but need to be evaluated within the context of BDA life histories and ecosystem dynamics.

An Ecological Perspective on Living with Fire in Ponderosa Pine Forests of Oregon and Washington: Resistance, Gone but not Forgotten.

Wildland fires (WLF) have become more frequent, larger, and severe with greater impacts to society and ecosystems and dramatic increases in firefighting costs. Forests throughout the range of ponderosa pine in Oregon and Washington are jeopardized by the interaction of anomalously dense forest structure, a warming and drying climate, and an expanding human population. These forests evolved with frequent interacting disturbances including low-severity surface fires, droughts, and biological disturbance agents (BDAs). Chronic low-severity disturbances were, and still are, critical to maintaining disturbance resistance, the property of an ecosystem to withstand disturbance while maintaining its structure and ecological function. Restoration of that historical resistance offers multiple social and ecological benefits. Moving forward, we need a shared understanding of the ecology of ponderosa pine forests to appreciate how restoring resistance can reduce the impacts of disturbances. Given contemporary forest conditions, a warming climate, and growing human populations, we predict continued elevation of tree mortality from drought, BDAs, and the large high-severity WLFs that threaten lives and property as well as ecosystem functions and services. We recommend more comprehensive planning to promote greater use of prescribed fire and management of reported fires for ecological benefits, plus increased responsibility and preparedness of local agencies, communities and individual homeowners for WLF and smoke events. Ultimately, by more effectively preparing for fire in the wildland urban interface, and by increasing the resistance of ponderosa pine forests, we can greatly enhance our ability to live with fire and other disturbances.

Water potential control of turgor-driven tracheid enlargement in Scots pine at its xeric distribution edge.

The extent to which water availability can be used to predict the enlargement and final dimensions of xylem conduits remains an open issue. We reconstructed the time course of tracheid enlargement in Pinus sylvestris trees in central Spain by repeated measurements of tracheid diameter on microcores sampled weekly during a 2 yr period. We analyzed the role of water availability in these dynamics empirically through time-series correlation analysis and mechanistically by building a model that simulates daily tracheid enlargement rate and duration based on Lockhart's equation and water potential as the sole input. Tracheid enlargement followed a sigmoid-like time course, which varied intra- and interannually. Our empirical analysis showed that final tracheid diameter was strongly related to water availability during tracheid enlargement. The mechanistic model was calibrated and successfully validated (R2  = 0.92) against the observed tracheid enlargement time course. The model was also able to reproduce the seasonal variations of tracheid enlargement rate, duration and final diameter (R2  = 0.84-0.99). Our results support the hypothesis that tracheid enlargement and final dimensions can be modeled based on the direct effect of water potential on turgor-driven cell expansion. We argue that such a mechanism is consistent with other reported patterns of tracheid dimension variation.

Coordination and trade-offs between leaf and stem hydraulic traits and stomatal regulation along a spectrum of isohydry to anisohydry.

The degree of plant iso/anisohydry, a widely used framework for classifying species-specific hydraulic strategies, integrates multiple components of the whole-plant hydraulic pathway. However, little is known about how it associates with coordination of functional and structural traits within and across different organs. We examined stem and leaf hydraulic capacitance and conductivity/conductance, stem xylem anatomical features, stomatal regulation of daily minimum leaf and stem water potential (Ψ), and the kinetics of stomatal responses to vapour pressure deficit (VPD) in six diverse woody species differing markedly in their degree of iso/anisohydry. At the stem level, more anisohydric species had higher wood density and lower native capacitance and conductivity. Like stems, leaves of more anisohydric species had lower hydraulic conductance; however, unlike stems, their leaves had higher native capacitance at their daily minimum values of leaf Ψ. Moreover, rates of VPD-induced stomatal closure were related to intrinsic rather than native leaf capacitance and were not associated with species' degree of iso/anisohydry. Our results suggest a trade-off between hydraulic storage and efficiency in the leaf, but a coordination between hydraulic storage and efficiency in the stem along a spectrum of plant iso/anisohydry.

Co-occurring woody species have diverse hydraulic strategies and mortality rates during an extreme drought.

From 2011 to 2013, Texas experienced its worst drought in recorded history. This event provided a unique natural experiment to assess species-specific responses to extreme drought and mortality of four co-occurring woody species: Quercus fusiformis, Diospyros texana, Prosopis glandulosa, and Juniperus ashei. We examined hypothesized mechanisms that could promote these species' diverse mortality patterns using postdrought measurements on surviving trees coupled to retrospective process modelling. The species exhibited a wide range of gas exchange responses, hydraulic strategies, and mortality rates. Multiple proposed indices of mortality mechanisms were inconsistent with the observed mortality patterns across species, including measures of the degree of iso/anisohydry, photosynthesis, carbohydrate depletion, and hydraulic safety margins. Large losses of spring and summer whole-tree conductance (driven by belowground losses of conductance) and shallower rooting depths were associated with species that exhibited greater mortality. Based on this retrospective analysis, we suggest that species more vulnerable to drought were more likely to have succumbed to hydraulic failure belowground.

Interactions of predominant insects and diseases with climate change in Douglas-fir forests of western Oregon and Washington, U.S.A.

Forest disturbance regimes are beginning to show evidence of climate-mediated changes, such as increasing severity of droughts and insect outbreaks. We review the major insects and pathogens affecting the disturbance regime for coastal Douglas-fir forests in western Oregon and Washington State, USA, and ask how future climate changes may influence their role in disturbance ecology. Although the physiological constraints of light, temperature, and moisture largely control tree growth, episodic and chronic disturbances interacting with biological factors have substantial impacts on the structure and functioning of forest ecosystems in this region. Understanding insect and disease interactions is critical to predicting forest response to climate change and the consequences for ecosystem services, such as timber, clean water, fish and wildlife. We focused on future predictions for warmer wetter winters, hotter drier summers, and elevated atmospheric CO2 to hypothesize the response of Douglas-fir forests to the major insects and diseases influencing this forest type: Douglas-fir beetle, Swiss needle cast, black stain root disease, and laminated root rot. We hypothesize that 1) Douglas-fir beetle and black stain root disease could become more prevalent with increasing, fire, temperature stress, and moisture stress, 2) future impacts of Swiss needle cast are difficult to predict due to uncertainties in May-July leaf wetness, but warmer winters could contribute to intensification at higher elevations, and 3) laminated root rot will be influenced primarily by forest management, rather than climatic change. Furthermore, these biotic disturbance agents interact in complex ways that are poorly understood. Consequently, to inform management decisions, insect and disease influences on disturbance regimes must be characterized specifically by forest type and region in order to accurately capture these interactions in light of future climate-mediated changes.

Stomatal kinetics and photosynthetic gas exchange along a continuum of isohydric to anisohydric regulation of plant water status.

Species' differences in the stringency of stomatal control of plant water potential represent a continuum of isohydric to anisohydric behaviours. However, little is known about how quasi-steady-state stomatal regulation of water potential may relate to dynamic behaviour of stomata and photosynthetic gas exchange in species operating at different positions along this continuum. Here, we evaluated kinetics of light-induced stomatal opening, activation of photosynthesis and features of quasi-steady-state photosynthetic gas exchange in 10 woody species selected to represent different degrees of anisohydry. Based on a previously developed proxy for the degree of anisohydry, species' leaf water potentials at turgor loss, we found consistent trends in photosynthetic gas exchange traits across a spectrum of isohydry to anisohydry. More anisohydric species had faster kinetics of stomatal opening and activation of photosynthesis, and these kinetics were closely coordinated within species. Quasi-steady-state stomatal conductance and measures of photosynthetic capacity and performance were also greater in more anisohydric species. Intrinsic water-use efficiency estimated from leaf gas exchange and stable carbon isotope ratios was lowest in the most anisohydric species. In comparisons between gas exchange traits, species rankings were highly consistent, leading to species-independent scaling relationships over the range of isohydry to anisohydry observed.

Thermotolerance and heat stress responses of Douglas-fir and ponderosa pine seedling populations from contrasting climates.

Temperature and the frequency and intensity of heat waves are predicted to increase throughout the 21st century. Germinant seedlings are expected to be particularly vulnerable to heat stress because they are in the boundary layer close to the soil surface where intense heating occurs in open habitats. We quantified leaf thermotolerance and whole-plant physiological responses to heat stress in first-year germinant seedlings in two populations each of Pinus ponderosa P. and C. Lawson (PIPO) and Pseudotsuga menziesii (Mirb.) Franco (PSME) from climates with contrasting precipitation and temperature regimes. Thermotolerance of detached needles was evaluated using chlorophyll fluorescence (FV/FM, FO) and electrolyte leakage. PSME was more heat tolerant than PIPO according to both independent assessments of thermotolerance. Following exposure of whole seedlings to a simulated heat wave at 45 °C for 1 h in a growth chamber, we monitored FV/FM, photosynthesis, stomatal conductance, non-structural carbohydrates (NSCs) and carbon isotope ratios (δ13C) for 14 days. Heat treatment induced significant reductions in FV/FM in both species and a transient reduction in photosynthetic gas exchange only in PIPO 1 day after treatment. Heat treatment induced an increase in glucose + fructose concurrent with a decrease in starch in both species, whereas total NSC and sucrose were not affected by heat treatment. The negative relationship between glucose + fructose and starch observed in treated plants may be due to the conversion of starch to glucose + fructose to aid recovery from heat-induced damage. Populations from drier sites displayed greater δ13C values than those from wetter sites, consistent with higher intrinsic water-use efficiency and drought resistance of populations from drier climates. Thermotolerance and heat stress responses appeared to be phenotypically plastic and representative of the environment in which plants were grown, whereas intrinsic water-use efficiency appeared to reflect ecotypic differentiation and the climate of origin.

Mapping 'hydroscapes' along the iso- to anisohydric continuum of stomatal regulation of plant water status.

The concept of iso- vs. anisohydry has been used to describe the stringency of stomatal regulation of plant water potential (ψ). However, metrics that accurately and consistently quantify species' operating ranges along a continuum of iso- to anisohydry have been elusive. Additionally, most approaches to quantifying iso/anisohydry require labour-intensive measurements during prolonged drought. We evaluated new and previously developed metrics of stringency of stomatal regulation of ψ during soil drying in eight woody species and determined whether easily-determined leaf pressure-volume traits could serve as proxies for their degree of iso- vs. anisohydry. Two metrics of stringency of stomatal control of ψ, (1) a 'hydroscape' incorporating the landscape of ψ over which stomata control ψ, and (2) the slope of the daily range of ψ as pre-dawn ψ declined, were strongly correlated with each other and with the leaf osmotic potential at full and zero turgor derived from pressure-volume curves.

Reliance on shallow soil water in a mixed-hardwood forest in central Pennsylvania.

We investigated depth of water uptake of trees on shale-derived soils in order to assess the importance of roots over a meter deep as a driver of water use in a central Pennsylvania catchment. This information is not only needed to improve basic understanding of water use in these forests but also to improve descriptions of root function at depth in hydrologic process models. The study took place at the Susquehanna Shale Hills Critical Zone Observatory in central Pennsylvania. We asked two main questions: (i) Do trees in a mixed-hardwood, humid temperate forest in a central Pennsylvania catchment rely on deep roots for water during dry portions of the growing season? (ii) What is the role of tree genus, size, soil depth and hillslope position on the depth of water extraction by trees? Based on multiple lines of evidence, including stable isotope natural abundance, sap flux and soil moisture depletion patterns with depth, the majority of water uptake during the dry part of the growing season occurred, on average, at less than ∼60 cm soil depth throughout the catchment. While there were some trends in depth of water uptake related to genus, tree size and soil depth, water uptake was more uniformly shallow than we expected. Our results suggest that these types of forests may rely considerably on water sources that are quite shallow, even in the drier parts of the growing season.

Expression of functional traits during seedling establishment in two populations of Pinus ponderosa from contrasting climates.

First-year tree seedlings represent a particularly vulnerable life stage and successful seedling establishment is crucial for forest regeneration. We investigated the extent to which Pinus ponderosa P. & C. Lawson populations from different climate zones exhibit differential expression of functional traits that may facilitate their establishment. Seeds from two populations from sites with contrasting precipitation and temperature regimes east (PIPO dry) and west (PIPO mesic) of the Oregon Cascade mountains were sown in a common garden experiment and grown under two water availability treatments (control and drought). Aboveground biomass accumulation, vegetative phenology, xylem anatomy, plant hydraulic architecture, foliar stable carbon isotope ratios (δ(13)C), gas exchange and leaf water relations characteristics were measured. No treatment or population-related differences in leaf water potential were detected. At the end of the first growing season, aboveground biomass was 74 and 44% greater in PIPO mesic in the control and drought treatments, respectively. By early October, 73% of PIPO dry seedlings had formed dormant buds compared with only 15% of PIPO mesic seedlings. Stem theoretical specific conductivity, calculated from tracheid dimensions and packing density, declined from June through September and was nearly twice as high in PIPO mesic during most of the growing season, consistent with measured values of specific conductivity. Intrinsic water-use efficiency based on δ(13)C values was higher in PIPO dry seedlings for both treatments across all sampling dates. There was a negative relationship between values of δ(13)C and leaf-specific hydraulic conductivity across populations and treatments, consistent with greater stomatal constraints on gas exchange with declining seedling hydraulic capacity. Integrated growing season assimilation and stomatal conductance estimated from foliar δ(13)C values and photosynthetic CO2-response curves were 6 and 28% lower, respectively, in PIPO dry seedlings. Leaf water potential at the turgor loss point was 0.33 MPa more negative in PIPO dry, independent of treatment. Overall, PIPO dry seedlings exhibited more conservative behavior, suggesting reduced growth is traded off for increased resistance to drought and extreme temperatures.

Linking nonstructural carbohydrate dynamics to gas exchange and leaf hydraulic behavior in Pinus edulis and Juniperus monosperma.

Leaf hydraulics, gas exchange and carbon storage in Pinus edulis and Juniperus monosperma, two tree species on opposite ends of the isohydry-anisohydry spectrum, were analyzed to examine relationships between hydraulic function and carbohydrate dynamics. Leaf hydraulic vulnerability, leaf water potential (Ψl ), leaf hydraulic conductance (Kleaf ), photosynthesis (A), stomatal conductance (gs) and nonstructural carbohydrate (NSC) content were analyzed throughout the growing season. Leaf hydraulic vulnerability was significantly lower in the relatively anisohydric J. monosperma than in the more isohydric P. edulis. In P. edulis, Ψl dropped and stayed below 50% loss of leaf hydraulic conductance (P50) early in the day during May, August and around midday in September, leading to sustained reductions in Kleaf . In J. monosperma, Ψl dropped below P50 only during August, resulting in the maintenance of Kleaf during much of the growing season. Mean A and gs during September were significantly lower in P. edulis than in J. monosperma. Foliar total NSC was two to three times greater in J. monosperma than in P. edulis in June, August and September. Consistently lower levels of total NSC in P. edulis suggest that its isohydric strategy pushes it towards the exhaustion of carbon reserves during much of the growing season.

Impacts of dwarf mistletoe on the physiology of host Tsuga heterophylla trees as recorded in tree-ring C and O stable isotopes.

Dwarf mistletoes, obligate, parasitic plants with diminutive aerial shoots, have long-term effects on host tree water relations, hydraulic architecture and photosynthetic gas exchange and can eventually induce tree death. To investigate the long-term (1886-2010) impacts of dwarf mistletoe on the growth and gas exchange characteristics of host western hemlock, we compared the diameter growth and tree-ring cellulose stable carbon (C) and oxygen (O) isotope ratios (δ(13)Ccell, δ(18)Ocell) of heavily infected and uninfected trees. The relative basal area growth of infected trees was significantly greater than that of uninfected trees in 1886-90, but declined more rapidly in infected than uninfected trees through time and became significantly lower in infected than uninfected trees in 2006-10. Infected trees had significantly lower δ(13)Ccell and δ(18)Ocell than uninfected trees. Differences in δ(18)Ocell between infected and uninfected trees were unexpected given that stomatal conductance and environmental variables that were expected to influence the δ(18)O values of leaf water were similar for both groups. However, estimates of mesophyll conductance (gm) were significantly lower and estimates of effective path length for water movement (L) were significantly higher in leaves of infected trees, consistent with their lower values of δ(18)Ocell. This study reconstructs the long-term physiological responses of western hemlock to dwarf mistletoe infection. The long-term diameter growth and δ(13)Ccell trajectories suggested that infected trees were growing faster than uninfected trees prior to becoming infected and subsequently declined in growth and leaf-level photosynthetic capacity compared with uninfected trees as the dwarf mistletoe infection became severe. This study further points to limitations of the dual-isotope approach for identifying sources of variation in δ(13)Ccell and indicates that changes in leaf internal properties such as gm and L that affect δ(18)Ocell must be considered.

Dynamics of leaf water relations components in co-occurring iso- and anisohydric conifer species.

Because iso- and anisohydric species differ in stomatal regulation of the rate and magnitude of fluctuations in shoot water potential, they may be expected to show differences in the plasticity of their shoot water relations components, but explicit comparisons of this nature have rarely been made. We subjected excised shoots of co-occurring anisohydric Juniperus monosperma and isohydric Pinus edulis to pressure-volume analysis with and without prior artificial rehydration. In J. monosperma, the shoot water potential at turgor loss (Ψ(TLP)) ranged from -3.4 MPa in artificially rehydrated shoots to -6.6 MPa in shoots with an initial Ψ of -5.5 MPa, whereas in P. edulis mean Ψ(TLP) remained at ∼ -3.0 MPa over a range of initial Ψ from -0.1 to -2.3 MPa. The shoot osmotic potential at full turgor and the bulk modulus of elasticity also declined sharply with shoot Ψ in J. monosperma, but not in P. edulis. The contrasting behaviour of J. monosperma and P. edulis reflects differences in their capacity for homeostatic regulation of turgor that may be representative of aniso- and isohydric species in general, and may also be associated with the greater capacity of J. monosperma to withstand severe drought.

Seasonal carbohydrate dynamics and growth in Douglas-fir trees experiencing chronic, fungal-mediated reduction in functional leaf area.

Stored non-structural carbohydrates (NSCs) could play an important role in tree survival in the face of a changing climate and associated stress-related mortality. We explored the effects of the stomata-blocking and defoliating fungal disease called Swiss needle cast on Douglas-fir carbohydrate reserves and growth to evaluate the extent to which NSCs can be mobilized under natural conditions of low water stress and restricted carbon supply in relation to potential demands for growth. We analyzed the concentrations of starch, sucrose, glucose and fructose in foliage, twig wood and trunk sapwood of 15 co-occurring Douglas-fir trees expressing a gradient of Swiss needle cast symptom severity quantified as previous-year functional foliage mass. Growth (mean basal area increment, BAI) decreased by ∼80% and trunk NSC concentration decreased by 60% with decreasing functional foliage mass. The ratio of relative changes in NSC concentration and BAI, an index of the relative priority of storage versus growth, more than doubled with increasing disease severity. In contrast, twig and foliage NSC concentrations remained nearly constant with decreasing functional foliage mass. These results suggest that under disease-induced reductions in carbon supply, Douglas-fir trees retain NSCs (either actively or due to sequestration) at the expense of trunk radial growth. The crown retains the highest concentrations of NSC, presumably to maintain foliage growth and shoot extension in the spring, partially compensating for rapid foliage loss in the summer and fall.

The impacts of water stress on phloem transport in Douglas-fir trees.

Despite the critical role that phloem plays in a number of plant functional processes and the potential impact of water stress on phloem structural and phloem sap compositional characteristics, little research has been done to examine how water stress influences phloem transport. The objectives of this study were to develop a more accurate understanding of how water stress affects phloem transport in trees, both in terms of the short-term impacts of water stress on phloem sap composition and the longer-term impacts on sieve cell anatomical characteristics. Phloem sieve cell conductivity (kp) was evaluated along a gradient of tree height and xylem water potential in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees in order to evaluate the influence of water stress on phloem transport capacity. The Hagen-Poiseuille equation was used with measurements of sieve cell anatomical characteristics, water content of phloem sap, non-structural carbohydrate content of phloem sap and shoot water potential (Ψl) to evaluate impacts of water stress on kp. Based on regression analysis, for each 1 MPa decrease in mean midday Ψl, sieve cell lumen radius decreased by 2.63 µm MPa(-1). Although there was no significant trend in sucrose content with decreasing Ψl, glucose and fructose content increased significantly with water stress and sieve cell relative water content decreased by 13.5% MPa(-1), leading to a significant increase in sugar molar concentration of 0.46 mol l(-1) MPa(-1) and a significant increase in viscosity of 0.27 mPa s MPa(-1). Modeled kp was significantly influenced both by trends in viscosity as well as by water stress-related trends in sieve cell anatomy.

The dynamic pipeline: hydraulic capacitance and xylem hydraulic safety in four tall conifer species.

Recent work has suggested that plants differ in their relative reliance on structural avoidance of embolism versus maintenance of the xylem water column through dynamic traits such as capacitance, but we still know little about how and why species differ along this continuum. It is even less clear how or if different parts of a plant vary along this spectrum. Here we examined how traits such as hydraulic conductivity or conductance, xylem vulnerability curves, and capacitance differ in trunks, large- and small-diameter branches, and foliated shoots of four species of co-occurring conifers. We found striking similarities among species in most traits, but large differences among plant parts. Vulnerability to embolism was high in shoots, low in small- and large-diameter branches, and high again in the trunks. Safety margins, defined as the pressure causing 50% loss of hydraulic conductivity or conductance minus the midday water potential, were large in small-diameter branches, small in trunks and negative in shoots. Sapwood capacitance increased with stem diameter, and was correlated with stem vulnerability, wood density and latewood proportion. Capacitive release of water is a dynamic aspect of plant hydraulics that is integral to maintenance of long-distance water transport.

Above- and belowground controls on water use by trees of different wood types in an eastern US deciduous forest.

Stomata control tree transpiration by sensing and integrating environmental signals originating in the atmosphere and soil, and co-occurring species may differ in inherent stomatal sensitivity to these above- and belowground signals and in the types of signals to which they respond. Stomatal responsiveness to environmental signals is likely to differ across species having different types of wood (e.g., ring-porous, diffuse-porous and coniferous) because each wood type differs in the structure, size and spatial distribution of its xylem conduits as well as in the scaling of hydraulic properties with stem diameter. The objective of this study was to evaluate the impact of variation in soil water availability and atmospheric evaporative demand on stomatal regulation of transpiration in seven co-occurring temperate deciduous forest species representing three wood types. We measured whole-tree sap flux and soil and atmospheric variables in a mixed deciduous forest in central Pennsylvania over the course of a growing season characterized by severe drought and large fluctuations in atmospheric vapor pressure deficit (D). The relative sensitivity of sap flux to soil drying was ∼2.2-2.3 times greater in the diffuse-porous and coniferous species than in the ring-porous species. Stomata of the ring-porous oaks were only about half as responsive to increased D as those of trees of the other two wood types. These differences in responsiveness to changes in the below- and aboveground environment implied that regulation of leaf water potential in the ring-porous oaks was less stringent than that in the diffuse-porous angiosperms or the conifers. The results suggest that increases in the frequency or intensity of summer droughts in the study region could have multiple consequences for forest function, including altered successional time courses or climax species composition and cumulative effects on whole-tree architecture, resulting in a structural and physiological legacy that would restrict the ability of trees to respond rapidly to more favorable growth conditions.

Contrasting hydraulic strategies in two tropical lianas and their host trees.

PREMISE OF THE STUDY: Tropical liana abundance has been increasing over the past 40 yr, which has been associated with reduced rainfall. The proposed mechanism allowing lianas to thrive in dry conditions is deeper root systems than co-occurring trees, although we know very little about the fundamental hydraulic physiology of lianas. METHODS: To test the hypothesis that two abundant liana species would physiologically outperform their host tree under reduced water availability, we measured rooting depth, hydraulic properties, plant water status, and leaf gas exchange during the dry season in a seasonally dry tropical forest. We also used a model to compare water use by one of the liana species and the host tree during drought. KEY RESULTS: All species measured were shallowly rooted. The liana species were more vulnerable to embolism than host trees and experienced water potentials that were predicted to result in substantial hydraulic losses in both leaves and stems. Water potentials measured in host trees were not negative enough to result in significant hydraulic losses. Model results predicted the liana to have greater gas exchange than its host tree during drought and nondrought conditions. CONCLUSIONS: The host tree species had a more conservative strategy for maintenance of the soil-to-leaf hydraulic pathway than the lianas it supported. The two liana species experienced embolism in stems and leaves, based on vulnerability curves and water potentials. These emboli were presumably repaired before the next morning. However, in the host tree species, reduced stomatal conductance prevented leaf or stem embolism.

Hydraulic safety margins and embolism reversal in stems and leaves: why are conifers and angiosperms so different?

Angiosperm and coniferous tree species utilize a continuum of hydraulic strategies. Hydraulic safety margins (defined as differences between naturally occurring xylem pressures and pressures that would cause hydraulic dysfunction, or differences between pressures resulting in loss of hydraulic function in adjacent organs (e.g., stems vs. leaves) tend to be much greater in conifers than angiosperms and serve to prevent stem embolism. However, conifers tend to experience embolism more frequently in leaves and roots than angiosperms. Embolism repair is thought to occur by active transport of sugars into empty conduits followed by passive water movement. The most likely source of sugar for refilling is from nonstructural carbohydrate depolymerization in nearby parenchyma cells. Compared to angiosperms, conifers tend to have little parenchyma or nonstructural carbohydrates in their wood. The ability to rapidly repair embolisms may rely on having nearby parenchyma cells, which could explain the need for greater safety margins in conifer wood as compared to angiosperms. The frequent embolisms that occur in the distal portions of conifers are readily repaired, perhaps due to the abundant parenchyma in leaves and roots, and these distal tissues may act as hydraulic circuit breakers that prevent tension-induced embolisms in the attached stems. Frequent embolisms in conifer leaves may also be due to weaker stomatal response to changes in ambient humidity. Although there is a continuum of hydraulic strategies among woody plants, there appear to be two distinct 'behaviors' at the extremes: (1) embolism prevention and (2) embolism occurrence and subsequent repair.