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1. Occult precipitation events (fog, dew, light-rain) can alter plant water and nutritional status, both directly through the aerial uptake of surface water and nutrients, and indirectly via redistribution of atmospheric resources to the soil. However, current frameworks that explain niche segregation, species interactions, and coexistence still consider that ground-rooted plants obtain resources almost exclusively via root absorption from soil.2. Here, we expand the plant hydrological niches model to incorporate both soil and atmospheric resource-axes, thus providing a more complete picture of how ground-rooted terrestrial plants obtain, remobilize, share, and compete for water and soluble nutrients.3. First, we describe how plants with different water acquisition strategies access directly or indirectly atmospheric resources. Then, we discuss how the use of such resources may promote spatiotemporal niche segregation, contributing to shape species distribution and abundance within plant communities. We illustrate this argument with examples from arid, mesic, and wet vegetation types. Finally, we examine how climate and land-use changes may influence plant hydrological niches, potentially altering community structure.4. Synthesis: Understanding how available atmospheric resources influences niche segregation in plant communities is a crucial step towards better predictions of species responses (e.g., changes in distribution, abundance and interactions) to climate change.

Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species-specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white-sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

Vulnerability to embolism varies between con-generic species distributed along aridity gradients, yet little is known about intraspecific variation and its drivers. Even less is known about intraspecific variation in tissues other than stems, despite results suggesting that roots, stems and leaves can differ in vulnerability. We hypothesized that intraspecific variation in vulnerability in leaves and stems is adaptive and driven by aridity. We quantified leaf and stem vulnerability of Quercus douglasii using the optical technique. To assess contributions of genetic variation and phenotypic plasticity to within-species variation, we quantified the vulnerability of individuals growing in a common garden, but originating from populations along an aridity gradient, as well as individuals from the same wild populations. Intraspecific variation in water potential at which 50% of total embolism in a tissue is observed (P50 ) was explained mostly by differences between individuals (>66% of total variance) and tissues (16%). There was little between-population variation in leaf/stem P50 in the garden, which was not related to site of origin aridity. Unexpectedly, we observed a positive relationship between wild individual stem P50 and aridity. Although there is no local adaptation and only minor phenotypic plasticity in leaf/stem vulnerability in Q. douglasii, high levels of potentially heritable variation within populations or strong environmental selection could contribute to adaptive responses under future climate change.

In the Earth’s tallest tree species, axial variation in conduit size matches theoretical predictions minimizing the accumulation of hydraulic resistance with height growth, within a constraint of maximum conduit diameter. Hydraulic limitations to tree height can be mitigated by widening the conducting elements toward a tree’s base. However, size limits of tracheid and vessel dimensions may constrain this compensation mechanism as the water transport pathway elongates. Moreover, variation in conduit size is poorly described in tall trees even though their long transport paths have high potential for hydraulic resistance. Here, we evaluated whether axial variation in conduit diameter was uniquely structured, or matched theoretical predictions in Sequoia sempervirens, Sequoiadendron giganteum, and Eucalyptus regnans that were 86–105 m tall and exceeded 85% of the maximum height for each species. Across Sequoia and Sequoiadendron, tree top tracheids maintained constant width, whereas tree base tracheids in the outermost ring were 20% wider in taller trees, suggesting maintenance of basipetal conduit widening with height growth. In all trees, the observed widening decreased at a rate per unit path length that fitted well to a power function with an exponent consistent with hydraulic compensation. However, below about 60 m from the tree tops, conduit diameters approached an asymptote beneath the power function, indicating a limit to maximum conduit size. Quantifying the distribution of base-to-top hydraulic resistance suggested that the minimal hydraulic benefit gained with increasingly wider conduits near the tree base may trade off with other factors such as maintaining mechanical strength or reducing fluid volume. We summarize these results into an anatomical model of height growth that includes limits to axial variation in conduit diameter and is supported by many physiological and anatomical observations.

Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012–2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a “common garden” thought experiment, how existing trait-based or response-based isohydricity metrics can be confounded by these environmental variations, leading to Type-1 (false positive) and Type-2 (false negative) errors; and (d) advocate for the use of model-based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale. This is the peer reviewed version of the following article: Feng, X, Ackerly, DD, Dawson, TE, et al. Beyond isohydricity: The role of environmental variability in determining plant drought responses. Plant Cell Environ. 2019; 42: 1104– 1111, which has been published in final form at https://doi.org/10.1111/pce.13486. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. This article may not be enhanced, enriched or otherwise transformed into a derivative work, without express permission from Wiley or by statutory rights under applicable legislation. Copyright notices must not be removed, obscured or modified. The article must be linked to Wiley’s version of record on Wiley Online Library and any embedding, framing or otherwise making available the article or pages thereof by third parties from platforms, services and websites other than Wiley Online Library must be prohibited. &nbsp

Water stored in tree stems (i.e., trunks and branches) is an important contributor to transpiration that can improve photosynthetic carbon gain and reduce the probability of cavitation. However, in tall trees, the capacity to store water may decline with height because of chronically low water potentials associated with the gravitational potential gradient. We quantified the importance of elastic stem water storage in the top 5–6 m of large (4.2–5.0 m diameter at breast height, 82.1–86.3 m tall) Sequoiadendron giganteum (Lindley) J. Buchholz (giant sequoia) trees using a combination of architectural measurements and automated sensors that monitored summertime diel rhythms in sap flow, stem diameter and water potential. Stem water storage contributed 1.5–1.8% of water transpired at the tree tops, and hydraulic capacitance ranged from 2.6 to 4.1 l MPa−1 m−3. These values, which are considerably smaller than reported for shorter trees, may be associated with persistently low water potentials imposed by gravity and could indicate a trend of decreasing water storage dynamics with height in tree. Branch diameter contraction and expansion consistently and substantially lagged behind fluxes in water potential and sap flow, which occurred in sync. This lag suggests that the inner bark, which consists mostly of live secondary phloem tissue, was an important hydraulic capacitor, and that hydraulic resistance between xylem and phloem retards water transfer between these tissues. We also measured tree-base sap flux, which lagged behind that measured in trunks near the tree tops, indicating additional storage in the large trunks between these measurement positions. Whole-tree sap flow ranged from 2227 to 3752 l day−1, corroborating previous records for similar-sized giant sequoia and representing the largest yet reported for any individual tree. Despite such extraordinarily high daily water use, we estimate that water stored in tree-top stems contributes minimally to transpiration on typical summer days.

The emergence of critical zone (CZ) science has provided an integrative platform for investigating plant ecophysiology in the context of landscape evolution, weathering and hydrology. The CZ lies between the top of the vegetation canopy and fresh, chemically unaltered bedrock and plays a pivotal role in sustaining life. We consider what the CZ perspective has recently brought to the study of plant ecophysiology. We specifically highlight novel research demonstrating the importance of the deeper subsurface for plant water and nutrient relations. We also point to knowledge gaps and research opportunities, emphasising, in particular, greater focus on the roles of deep, nonsoil resources and how those resources influence and coevolve with plants as a frontier of plant ecophysiological research.

Contents Summary 1156 I. Introduction 1156 II. How often are leaves wet? 1157 III. The costs of leaf wetting 1157 IV. The real and potential benefits of leaf wetting 1161 V. Wet leaves: costs, benefits and tradeoffs in a changing world 1165 Acknowledgements 1166 References 1166 SUMMARY: An often-overlooked feature of all plants is that their leaf surfaces are wet for significant periods over their lifetimes. Leaf wetting has a number of direct and indirect effects on plant function from the scale of the leaf to that of the ecosystem. The costs of leaf wetting for plant function, such as the growth of pathogens and the leaching of nutrients, have long been recognized. However, an emerging body of research has also begun to demonstrate some very clear benefits. For instance, leaf wetting can improve plant-water relations and lead to increased photosynthesis. Leaf wetting may also lead to synergistic effects on plant function, such as when leaf water potential improvements lead to enhanced growth that does not occur when plant leaves are dry. We identify important reasons why leaf wetting can be critical for plant sciences to not only acknowledge, but also directly address, in future research. To do so, we provide a framework for the consideration of the relative balance of the various costs and benefits resulting from leaf wetting, as well as how this balance may be expected to change given projected scenarios of global climate change in the future.

Desiccation tolerance (DT) is the ability of plants to tolerate low leaf water content from which they can recover when water becomes available. DT species mostly occur in seasonally-dry environments, tropical rocky outcrops such as the campos rupestres. Velloziaceae, a prominent family in the campos rupestres, comprises both DT and non-DT species. We tested if DT Vellozia nivea was associated with low productivity under mild drought conditions and survival under extreme drought. We measured traits related to drought resistance (turgor loss point – ΨTLP), plant functioning and productivity (e.g., gas exchange, biomass), non-structural carbohydrate dynamics and mortality in two species, one DT and one non-DT Vellozia species in two water-stress experiments simulating mild and extreme drought. Both species had similar survival rates under mild drought conditions, but the DT species had a lower productivity. Under extreme drought, the dynamics of non-structural carbohydrates was similar in both species but only the non-DT species died, and the cause of mortality was likely hydraulic dysfunction, as they died after reaching ΨTLP. All individuals of the DT species desiccated when they reached ΨTLP but rehydrated after re-watering. We show that the DT strategy in V. nivea implies a costs in terms of productivity, specially under mild drought conditions, but it allows survival under extreme drought, indicating a trade-off between productivity and survival in the DT species. Climate change predictions with more frequent and severe droughts will likely affect the abundance and distribution of DT and non-DT taxa.

Epiphytes are common in tropical montane cloud forests (TMCFs) and play many important ecological roles, but the degree to which these unique plants will be affected by changes in climate is unknown. We investigated the drought responses of three vascular epiphyte communities bracketing the cloud base during a severe, El Nino-impacted dry season. Epiphytes were instrumented with sap flow probes in each site. Leaf water potential and pressure-volume curve parameters were also measured before and during the drought. We monitored the canopy microclimate in each site to determine the drivers of sap velocity across the sites. All plants greatly reduced their water use during the drought, but recovery occurred more quickly for plants in the lower and drier sites. Plants in drier sites also exhibited the greatest shifts in the osmotic potential at full saturation and the turgor loss point. Although all individuals survived this intense drought, epiphytes in the cloud forest experienced the slowest recovery, suggesting that plants in the TMCF are particularly sensitive to severe drought. Although vapor pressure deficit was an important driver of sap velocity in the highest elevation site, other factors, such as the volumetric water content of the canopy soil, were more important at lower (and warmer) sites.

The ratio of leaf internal (ci ) to ambient (ca ) partial pressure of CO2 , defined here as χ, is an index of adjustments in both leaf stomatal conductance and photosynthetic rate to environmental conditions. Measurements and proxies of this ratio can be used to constrain vegetation model uncertainties for predicting terrestrial carbon uptake and water use. We test a theory based on the least-cost optimality hypothesis for modelling historical changes in χ over the 1951-2014 period, across different tree species and environmental conditions, as reconstructed from stable carbon isotopic measurements across a global network of 103 absolutely dated tree-ring chronologies. The theory predicts optimal χ as a function of air temperature, vapour pressure deficit, ca and atmospheric pressure. The theoretical model predicts 39% of the variance in χ values across sites and years, but underestimates the intersite variability in the reconstructed χ trends, resulting in only 8% of the variance in χ trends across years explained by the model. Overall, our results support theoretical predictions that variations in χ are tightly regulated by the four environmental drivers. They also suggest that explicitly accounting for the effects of plant-available soil water and other site-specific characteristics might improve the predictions.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015–2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season (“control scenario”). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf. © 2019 Gimenez, Jardine, Higuchi, Negrón-Juárez, Sampaio-Filho, Cobello, Fontes, Dawson, Varadharajan, Christianson, Spanner, Araújo, Warren, Newman, Holm, Koven, McDowell and Chambers.

Understanding how climate alters plant-soil water dynamics, and its impact on physiological functions, is critical to improved predictions of vegetation responses to climate change. Here we analyzed how belowground interactions for water shift under warming and drought, and associated impacts on plant functions. In a semi-arid woodland, adult trees (pinon and juniper) and perennial grasses (blue grama) were exposed to warming and precipitation reduction. After 6 years of continuous treatment exposure, soil and plant water isotopic composition was measured to assess plant water uptake depths and community-level water source partitioning. Warming and drought modified plant water uptake depths. Under warming, contrasting changes in water sources between grasses and trees reduced belowground water source partitioning, resulting in higher interspecific competition for water. Under drought, shifts in trees and grass water sources to deeper soil layers resulted in the maintenance of the naturally occurring water source partitioning among species. Trees showed higher water stress, and reduced water use and photosynthesis in response to warming and drought. This case study demonstrates that neighboring plants shift their competitive interactions for water under prolonged warming and drought, but regardless of whether changes in moisture sources will result in increased competition among species or maintained partitioning of water resources, these competitive adaptations may easily be overridden by climate extremes.

Flowers face desiccating conditions, yet little is known about their ability to transport water. We quantified variability in floral hydraulic conductance (Kflower ) for 20 species from 10 families and related it to traits hypothesized to be associated with liquid and vapour phase water transport. Basal angiosperm flowers had trait values associated with higher water and carbon costs than monocot and eudicot flowers. Kflower was coordinated with water supply (vein length per area, VLA) and loss (minimum epidermal conductance, gmin ) traits among the magnoliids, but was insensitive to variation in these traits among the monocots and eudicots. Phylogenetic independent contrast (PIC) correlations revealed that few traits had undergone coordinated evolution. However, VLA and the desiccation time (Tdes ), the quotient of water content and gmin , had significant trait and PIC correlations. The near absence of stomata from monocot and eudicot flowers may have been critical in minimizing water loss rates among these clades. Early divergent, basal angiosperm flowers maintain higher Kflower because of traits associated with high rates water loss and water supply, while monocot and eudicot flowers employ a more conservative strategy of limiting water loss and may rely on stored water to maintain turgor and delay desiccation.

© 2018 American Society of Plant Biologists. All rights reserved. Although recent findings suggest that xylem embolism represents a significant, drought-induced damaging process in land plants, substantial debate surrounds the capacity of long-vesseled, ring-porous species to resist embolism. We investigated whether recent methodological developments could help resolve this controversy within Quercus, a long-vesseled, ring-porous temperate angiosperm genus, and shed further light on the importance of xylem vulnerability to embolism as an indicator of drought tolerance. We used the optical technique to quantify leaf and stem xylem vulnerability to embolism of eight Quercus species from the Mediterranean-type climate region of California to examine absolute measures of resistance to embolism as well as any potential hydraulic segmentation between tissue types. We demonstrated that our optical assessment reflected flow impairment for a subset of our sample species by quantifying changes in leaf hydraulic conductance in dehydrating branches. Air-entry water potential varied 2-fold in leaves, ranging from −1.7 ± 0.25 MPa to −3.74 ± 0.23 MPa, and 4-fold in stems, ranging from −1.17 ± 0.04 MPa to −4.91 ± 0.3 MPa. Embolism occurred earlier in leaves than in stems in only one out of eight sample species, and plants always lost turgor before experiencing stem embolism. Our results show that long-vesseled North American Quercus species are more resistant to embolism than previously thought and support the hypothesis that avoiding stem embolism is a critical component of drought tolerance in woody trees. Accurately quantifying xylem vulnerability to embolism is essential for understanding species distributions along aridity gradients and predicting plant mortality during drought.

Author(s): Templer, PH; Weathers, KC; Ewing, HA; Dawson, TE; Mambelli, S; Lindsey, AM; Webb, J; Boukili, VK; Firestone, MK | Abstract: A defining feature of the redwood forest in coastal California is the presence of fog in the summer months, a time when there is typically little rainfall. Our goal was to determine the role of summer fog in canopy transformation of nitrogen, nitrogen uptake by trees and photosynthesis within a coastal redwood forest ecosystem. We measured horizontal and vertical inputs of nitrogen, the isotopic composition of nitrogen in a variety of atmospheric sources (summer fog, winter rain and throughfall throughout the year), nitrogen pools (soil solution) and plant tissue (roots and foliage), as well as rates of photosynthesis and nitrogen uptake by trees. Throughfall nitrogen fluxes were greater at the forest edge compared to the interior both within the canopy (sampled 10m above-ground) and onto the forest floor (sampled 1m above-ground; Pl0.05). Similarly, soil solution NO3- and total inorganic nitrogen were greater at the forest edge compared to the interior (P=0.0014 and 0.009, respectively). Whereas natural abundance δ15NO3 values were not significantly different between winter rain (measured as bulk precipitation) and summer fog water (average δ15N=-1.2±0.680/00), δ15NH4 values were significantly greater in fog water (11.4±2.70/00) compared to rain (1.2±0.90/00). We found no difference in δ15N in roots from forest edge trees compared to interior trees. In contrast, nitrogen concentrations and δ15N in foliage from forest edge trees were significantly greater compared to interior trees (Pl0.0001), suggesting that the leaves of forest edge trees may be obtaining a greater proportion of their nitrogen from fog compared to those of the interior trees. Natural abundance 13C of leaf sugars and rates of photosynthesis were significantly higher at the forest edge compared to the interior during the fog season (Pl0.05), but not different between locations in the rain season (Pg0.05). Nitrification in the forest floor, rather than the canopy, is the primary source of NO3- in these soils throughout the year. Synthesis. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees.

© The Author 2015. Published by Oxford University Press. The California Mediterranean savanna has harsh summer conditions with minimal soil moisture, high temperature, high incoming solar radiation and little or no precipitation. Deciduous blue oaks, Quercus douglasii Hook. and Arn., are winter-deciduous obligate phreatophytes, transpiring mostly groundwater throughout the summer drought. The objective of this work is to fully characterize the seasonal trends of photosynthesis in blue oaks as well as the mechanistic relationships between leaf structure and function. We estimate radiative load of the leaves via the FLiES model and perform in situ measurements of leaf water potential, leaf nitrogen content, an index of chlorophyll content (SPAD readings), photosynthetic and electron transport capacity, and instantaneous rates of CO2 assimilation and electron transport. We measured multiple trees over 3 years providing data from a range of conditions. Our study included one individual that demonstrated strong drought stress as indicated by changes in SPAD readings, leaf nitrogen and all measures of leaf functioning. In the year following severe environmental stress, one individual altered foliation patterns on the crown but did not die. In all other individuals, we found that net carbon assimilation and photosynthetic capacity decreased during the summer drought. SPAD values, electron transport rate (ETR) and quantum yield of photosystem II (PSII) did not show a strong decrease during the summer drought. In most individuals, PSII activity and SPAD readings did not indicate leaf structural or functional damage throughout the season. While net carbon assimilation was tightly coupled to stomatal conductance, the coupling was not as tight with ETR possibly due to contributions from photorespiration or other protective processes. Our work demonstrates that the blue oaks avoid structural damage by maintaining the capacity to convert and dissipate incoming solar radiation during the hot summer drought and are effective at fixing carbon by maximizing rates during the mild spring conditions.

Key message Highly variable nighttime transpiration, with higher rates generally observed after preceding fog, is prevalent in dominant tree species of the nutri- ent-poor tropical montane cloud forest environment of central Veracruz, Mexico. Abstract Although nighttime transpiration (En) is pre- valent in a wide range of species from cloud-affected for- ests, its magnitude relative to total daily transpiration (Ed) as reported in the literature is generally small (En/Ed is 0.12 on average). In the present study, we observed high dry- season En/Ed ratios with large night-to-night variation in dominant species from the tropical montane cloud forest (TMCF) zone of central Veracruz, Mexico: 0.22 ± 0.18 for Quercus lancifolia (old-growth TMCF); 0.26 ± 0.14 and 0.16 ± 0.16 for Alchornea latifolia and Alnus jorull- ensis, respectively (regenerating post-fire TMCF); and 0.30 ± 0.20 to 0.12 ± 0.21 for Pinus patula (young and mature pine plantations). En was determined as the dif- ference between observed nocturnal sap flow and estimated refilling of stem water storage, the latter of which was on average: 21-25 % of nocturnal sap flow for Q. lancifolia ;6 and 5 % for A. latifolia and A. jorullensis, respectively; and 21-23 % for P. patula. Night-to-night variation in En was mostly due to large variation in vapor pressure deficit (VPD) related in turn to the alternation of cold fronts (producing fog events) and high pressure weather (pro- ducing nights with VPD up to 2 kPa). Moreover, in the hours following fog events without concurring rainfall, En was often higher as compared to fog-free nights with similar VPD across all species examined. Low-nutrient availability and high water content of the soils in the study area suggest a nutrient uptake benefit associated with the relatively high En rates observed.

For most angiosperms, producing and maintaining flowers is critical to sexual reproduction, yet little is known about the physiological processes involved in maintaining flowers throughout anthesis. Among extant species, flowers of the genus Calycanthus have the highest hydraulic conductance and vein densities of species measured to date, yet they can wilt by late morning under hot conditions. Here, we combine diurnal measurements of gas exchange and water potential, pressure-volume relations, functional responses of gas exchange, and characterization of embolism formation using high resolution X-ray computed microtomography to determine drought responses of Calycanthus flowers. Transpiration from flowers frequently exceeded transpiration from leaves, and flowers were unable to limit transpiration under conditions of high vapour pressure deficit. As a result, they rely heavily on hydraulic capacitance to prevent water potential declines. Despite having high water potentials at turgor loss, flowers were very resistant to embolism formation, with no embolism apparent until tepal water potentials had declined to -2 MPa. Although Calycanthus flowers remain connected to the stem xylem and have high hydraulic capacitance, their inability to curtail transpiration leads to turgor loss. These results suggest that extreme climate events may cause flower failure, potentially preventing successful reproduction.

The goal of this study was to determine the effects of atmospheric demand on both plant water relations and daily wholetree water balance across the upper limit of a cloud forest at the mean base height of the trade wind inversion in the tropical trade wind belt. We measured the microclimate and water relations (sap flow, water potential, stomatal conductance, pres sure–volume relations) of Metrosideros polymorpha Gaudich. var. polymorpha in three habitats bracketing the cloud forest’s upper limit in Hawai‘i to understand the role of water relations in determining ecotone position. The subalpine shrubland site, located 100 m a bove the cloud forest boundary, had the highest vapor pressure deficit, the least amount of rainfall and the highest levels of nighttime transpiration (EN) of all three sites. In the shrubland site, on average, 29% of daily whole-tree transpiration occurred at night, while on the driest day of the study 50% of total daily transpiration occurred at night. While EN occurred in the cloud forest habitat, the proportion of total daily transpiration that occurred at night was much lower (4%). The average leaf water potential (Ψleaf) was above the water potential at the turgor loss point (ΨTLP) on both sides of the ecotone due to strong stomatal regulation. While stomatal closure maintained a high Ψleaf, the minimum leaf water potential (Ψleafmin) was close to ΨTLP, indicating that drier conditions may cause drought stress in these habitats and may be an important driver of current landscape patterns in stand density.

In tall trees, the mechanisms by which foliage maintains sufficient turgor pressure and water content against height-related constraints remain poorly understood. Pressure–volume curves generated from leafy shoots collected crown-wide from 12 large Sequoiadendron giganteum (Lindley) J. Buchholz (giant sequoia) trees provided mechanistic insights into how the components of water potential vary with height in tree and over time. The turgor loss point (TLP) decreased with height at a rate indistinguishable from the gravitational potential gradient and was controlled by changes in tissue osmotica. For all measured shoots, total relative water content at the TLP remained above 75%. This high value has been suggested to help leaves avoid precipitous declines in leaf-level physiological function, and in giant sequoia was controlled by both tissue elasticity and the balance of water between apoplasm and symplasm. Hydraulic capacitance decreased only slightly with height, but importantly this parameter was nearly double in value to that reported for other tree species. Total water storage capacity also decreased with height, but this trend essentially disappeared when considering only water available within the typical range of water potentials experienced by giant sequoia. From summer to fall measurement periods we did not observe osmotic adjustment that would depress the TLP. Instead we observed a proportional shift of water into less mobile apoplastic compartments leading to a reduction in hydraulic capacitance. This collection of foliar traits allows giant sequoia to routinely, but safely, operate close to its TLP, and suggests that gravity plays a major role in the water relations of Earth’s largest tree species.

The ecophysiology of tropical montane cloud forest (TMCF) trees is influenced by crown-level microclimate factors including regular mist/fog water inputs, and large variations in evaporative demand, which in turn can significantly impact water balance. We investigated the effect of such microclimatic factors on canopy ecophysiology and branch-level water balance in the dry season of a seasonal TMCF in Veracruz, Mexico, by quantifying both water inputs (via foliar uptake, FU) and outputs (day- and night-time transpiration, NT). Measurements of sap flow, stomatal conductance, leaf water potential and pressure-volume relations were obtained in Quercus lanceifolia, a canopy-dominant tree species. Our results indicate that FU occurred 34% of the time and led to the recovery of 9% (24 ± 9.1 L) of all the dry-season water transpired from individual branches. Capacity for FU was independently verified for seven additional common tree species. NT accounted for approximately 17% (46 L) of dry-season water loss. There was a strong correlation between FU and the duration of leaf wetness events (fog and/or rain), as well as between NT and the night-time vapour pressure deficit. Our results show the clear importance of fog and NT for the canopy water relations of Q. lanceifolia.

During daylight hours, the isotope composition of leaf water generally approximates steady-state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady-state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of 18O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non-steady state. We show that non–steady-state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.

It is commonly assumed that transpiration does not occur at night because leaf stomata are closed in the dark. We tested this assumption across a diversity of ecosystems and woody plant species by various methods to explore the circumstances when this assumption is false. Our primary goals were: (1) to evaluate the nature and magnitude of nighttime transpiration, E(n), or stomatal conductance, g(n); and (2) to seek potential generalizations about where and when it occurs. Sap-flow, porometry and stable isotope tracer measurements were made on 18 tree and eight shrub species from seven ecosystem types. Coupled with environmental data, our findings revealed that most of these species transpired at night. For some species and circumstances, nighttime leaf water loss constituted a significant fraction of total daily water use. Our evidence shows that E(n) or g(n) can occur in all but one shrub species across the systems we investigated. However, under conditions of high nighttime evaporative demand or low soil water availability, stomata were closed and E(n) or g(n) approached zero in eleven tree and seven shrub species. When soil water was available, E(n) or g(n) was measurable in these same species demonstrating plasticity for E(n) or g(n). We detected E(n) or g(n) in both trees and shrubs, and values were highest in plants from sites with higher soil water contents and in plants from ecosystems that were less prone to atmospheric or soil water deficits. Irrespective of plant or ecosystem type, many species showed E(n) or g(n) when soil water deficits were slight or non-existent, or immediately after rainfall events that followed a period of soil water deficit. The strongest relationship was between E(n) or g(n) and warm, low humidity and (or) windy (> 0.8 m s(-1)) nights when the vapor pressure deficit remained high (> 0.2 kPa in wet sites, > 0.7 kPa in dry sites). Why E(n) or g(n) occurs likely varies with species and ecosystem type; however, our data support four plausible explanations: (1) it may facilitate carbon fixation earlier in the day because stomata are already open; (2) it may enhance nutrient supply to distal parts of the crown when these nutrients are most available (in wet soils) and transport is rapid; (3) it may allow for the delivery of dissolved O(2) via the parenchyma to woody tissue sinks; or (4) it may occur simply because of leaky cuticles in older leaves or when stomata cannot close fully because of obstructions from stomatal (waxy) plugs, leaf endophytes or asymmetrical guard cells (all non-adaptive reasons). We discuss the methodological, ecophysiological, and theoretical implications of the occurrence of E(n) or g(n) for investigations at a variety of scales.

Across land plants there is a general pattern of xylem conduit diameters widening towards the stem base thus reducing the accumulation of hydraulic resistance as plants grow taller. In conifers, xylem conduits consist of cells with closed end-walls and water must flow through bordered pits imbedded in the side walls. As a consequence both cell size, which determines the numbers of walls that the conductive stream of water must cross, as well as the characteristics of the pits themselves, crucially affect total hydraulic resistance. Because both conduit size and pit features influence hydraulic resistance in tandem, we hypothesized that features of both should vary predictably with one another. To test this prediction we sampled a single tall (94.8 m) Sequoiadendron giganteum tree (giant sequoia), collecting wood samples from the most recent annual ring progressively downwards from the tree top to the base. We measured tracheid diameter and length, number of pits per tracheid, and the areas of pit apertures, tori, and margos. Tracheid diameter widened from treetop to base following a power law with an exponent (tracheid diameterstem length slope) of approximately 0.20. A similar scaling exponent was found between tracheid length and distance from tree top. Additionally, pit aperture, torus, and margo areas all increased (again with a power of ~0.20) with distance from tree top, paralleling the observed variation in tracheid diameter and length. Pit density scaled isometrically with tracheid length. Within individual tracheids, total permeable area of pits, measured as the sum of the margo areas, scaled isometrically with lumen area. Given that pores of the margo membrane are believed to increase in parallel with membrane area, from a strictly anatomical perspective, our results support the interpretation that pit resistance remains a relatively constant fraction of total resistance along the hydraulic pathway.

Previous research suggests that the lifetime carbon gain of a leaf is constrained by a tradeoff between metabolism and longevity. The biophysical reasons underlying this tradeoff are not fully understood. We used a photosynthesis-leaf water balance model to evaluate biophysical constraints on carbon gain. Leaf hydraulic conductance (K(Leaf)), carbon isotope discrimination (Δ(13)C), leaf mass per unit area (LMA) and the driving force for water transport from stem to leaf (ΔΨ(Stem-Leaf)) were characterized for leaves spanning three orders of magnitude in surface area and two orders of magnitude in lifespan. We observed positive isometric scaling between K(Leaf) and leaf area but no relationship between Δ(13)C and leaf area. Leaf lifespan and LMA had minimal effect on K(Leaf) per unit leaf area, but a negative correlation exists among LMA, lifespan, and K(Leaf) per unit dry mass. During periods of leaf water loss, ΔΨ(Stem-Leaf) was relatively constant. We show for the first time that K(Leaf, mass), an index of the carbon cost associated with water use, is negatively correlated with lifespan. This highlights the importance of characterizing K(Leaf, mass) and suggests a tradeoff between resource investment in liquid phase processes and structural rigidity.

Background and Aims Cerradao (Brazilian woodland savannas) and seasonally dry forests (SDF) from southeastern Brazil occur under the same climate but are remarkably distinct in species composition. The objective of this study was to evaluate the role of soil origin in the initial growth and distribution of SDF and Cerradao species. Methods We conducted a greenhouse experiment growing Cerradao and SDF tree seedlings over their soil and the soil of the contrasting vegetation type. We evaluated soil nutrient availability and seedling survivorship, growth and leaf functional traits. Results Despite the higher nutrient availability in SDF soils,soilorigindidnotaffectseedlingsurvivorship.The three SDF species demonstrated home-soil advantage, enhanced growth with increasing soil nutrient availability and had higher growth rates than Cerradao species, even on Cerradao soils. Growth of Cerradao seedlings was not higher on Cerradao soil and, overall, was not positively correlated with soil nutrient availability. Conclusions SDF species are fast-growing species while Cerradao trees tend to be slow-growing species. Although savanna soil reduces growth of forest species, our findings suggest that soil chemical attributes, alone, does not exclude the occurrence of SDF seedlings in Cerradao and vice-versa.

The stable hydrogen isotope ratios (δD) of leaf wax n-alkanes record valuable information on plant and ecosystem water relations. It remains, however, unknown if leaf wax n-alkane δD values record only environmental variation during the brief period of time of leaf growth or if leaf wax n-alkane δD values are affected by environmental variability throughout the entire lifespan of a leaf. To resolve these uncertainties, we irrigated Populus trichocarpa trees with a pulse of deuterium-enriched water and used compound-specific stable hydrogen isotope analyses to test if the applied tracer could be recovered from leaf wax n-alkanes of leaves that were at different stages of their development during the tracer application. Our experiment revealed that only leaf wax n-alkanes from leaves that had developed during the time of the tracer application were affected, while leaves that were already fully matured at the time of the tracer application were not. We conclude from our study that under controlled environmental conditions, leaf wax n-alkanes are synthesized only early in the ontogeny of a leaf. Our experiment has implications for the interpretation of leaf wax n-alkane δD values in an environmental context, as it suggests that these compounds record only a brief period of the environmental variability that a leaf experiences throughout its life.

Stem photosynthesis can contribute significantly to woody plant carbon balance, particularly in times when leaves are absent or in ‘open’ crowns with sufficient light penetration. We explored the significance of woody tissue (stem) photosynthesis for the carbon income in three California native plant species via measurements of chlorophyll concentrations, radial stem growth, bud biomass and stable carbon isotope composition of sugars in different plant organs. Young plants of Prunus ilicifolia, Umbellularia californica and Arctostaphylos manzanita were measured and subjected to manipulations at two levels: trunk light exclusion (100 and 50%) and complete defoliation. We found that long-term light exclusion resulted in a reduction in chlorophyll concentration and radial growth, demonstrating that trunk assimilates contributed to trunk carbon income. In addition, bud biomass was lower in covered plants compared to uncovered plants. Excluding 100% of the ambient light from trunks on defoliated plants led to an enrichment in 13C of trunk phloem sugars. We attributed this effect to a reduction in photosynthetic carbon isotope discrimination against 13C that in turn resulted in an enrichment in 13C of bud sugars. Taken together our results reveal that stem photosynthesis contributes to the total carbon income of all species including the buds in defoliated plants.

Water transport in conifers occurs through single-celled tracheids that are connected to one another via intertracheid pit membranes. These membranes have two components: the porous margo, which allows water to pass through the membrane, and the impermeable torus, which functions to isolate gas-filled tracheids. During drought, tracheids can become air filled and thus hydraulically dysfunctional, a result of air entering through the pit membrane and nucleating cavitation in the water column. What are the hydraulic tradeoffs associated with cavitation resistance at the pit level, and how do they vary within the structural components of the intertracheid pit? To address these questions, we examined pit structure in 15 species of Cupressaceae exhibiting a broad range of cavitation resistances. Across species, cavitation resistance was most closely correlated to the ratio of the torus to pit aperture diameter but did not vary systematically with margo porosity. Furthermore, our data indicate that constraints on pit hydraulic efficiency are shared: the pit aperture limits pit conductivity in more drought-resistant taxa, while increased margo resistance is more likely to control pit conductivity in species that are more vulnerable to cavitation. These results are coupled with additional data concerning pit membrane structure and function and are discussed in the context of the evolutionary biogeography of the Cupressaceae.

The stable oxygen isotope ratio (delta(18)O) of plant material has been shown to contain essential information on water and carbon fluxes at the plant and ecosystem scales. However, the effective path length (L(m)), a parameter introduced to leaf-water models still requires a comprehensive biological characterization to allow interpretation of delta(18)O values in plant material with confidence. Here, we tested the variability of L(m) across and within three species that developed leaves in environments with different relative humidity. We also tested whether the L(m) of fully developed leaves is affected by short-term fluctuations in relative humidity. We determined that significant differences in L(m) exist among Phaseolus vulgaris, Rizinus communis and Helianthus annuus. Within a given species, however, L(m) values did not differ significantly among individuals. These findings indicate that L(m) is species specific and a relatively constant parameter and that L(m) will not obscure the interpretation of delta(18)O values in plant material of a given species. We urge caution, however, because values for L(m) are derived from fitting leaf-water models to measured values of delta(18)O, so care must be taken in assigning a 'cause' to values of L(m) as they likely capture a combination of different biological leaf properties.

Although crown wetting events can increase plant water status, leaf wetting is thought to negatively affect plant carbon balance by depressing photosynthesis and growth. We investigated the influence of crown fog interception on the water and carbon relations of juvenile and mature Sequoia sempervirens trees. Field observations of mature trees indicated that fog interception increased leaf water potential above that of leaves sheltered from fog. Furthermore, observed increases in leaf water potential exceeded the maximum water potential predicted if soil water was the only available water source. Because field observations were limited to two mature trees, we conducted a greenhouse experiment to investigate how fog interception influences plant water status and photosynthesis. Pre-dawn and midday branchlet water potential, leaf gas exchange and chlorophyll fluorescence were measured on S. sempervirens saplings exposed to increasing soil water deficit, with and without overnight canopy fog interception. Sapling fog interception increased leaf water potential and photosynthesis above the control and soil water deficit treatments despite similar dark-acclimated leaf chlorophyll fluorescence. The field observations and greenhouse experiment show that fog interception represents an overlooked flux into the soil-plant-atmosphere continuum that temporarily, but significantly, decouples leaf-level water and carbon relations from soil water availability.

Summary • Access to deeper soil water and water-conserving traits should reduce water stress for ectomycorrhizal fungi, permitting function during drought. Here, we explored whether epigeous fruiting of ectomycorrhizal fungi during drought was facilitated by access to deep soil water, how much water was lost from sporocarps, and how sporocarp surface to volume ratios affected water-loss rates.  We used oxygen stable isotope analysis of water combined with modeling of water sources used by ectomycorrhizal fungi; measured sporocarp water loss using a transient porometer, and related water loss to vapor pressure deficit (VPD) and sporocarp morphology.  In drier soils sporocarps likely derived a significant portion (25–80%) of their water from deep (> 30 cm) or hydraulically lifted water. Amanita muscaria had water-loss rates over twice those of Suillus sp., Boletus edulis, Tricholoma spp. and Russula albonigra. Vapor pressure deficit was an excellent predictor of water-loss rates for individual mushrooms. Sporocarp surface to volume ratios explained much of the variation among mushrooms in the slope of VPD–water loss relationships.  Access to deeper soil water might be a significant driver of ectomycorrhizal symbiotic function, sporocarp distribution, fruiting habit and morphology. Sporocarp morphology can affect water-loss rates and hence influences fungal ability to fruit during summer drought.

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.

Thermometric sap flow sensors are widely used to measure water flow in roots, stems and branches of plants. Comparison of the timing of flow in branches and stems has been used to estimate water capacitance of large trees. We review studies of sap flow in branches and present our own data to show that there is wide variation in the patterns and timing of sap flow of branches in different parts of the crown, owing to the course of daily solar illuminance. In contiguous forest, east-facing and upper branches are illuminated earlier than west-facing and lower branches and most capacitance studies do not include adequate information about branch sampling regimes relative to the overall pattern of crown illuminance, raising questions about the accuracy of capacitance estimates. Measuring only upper branches and normalising these results to represent the entire crown is dangerous because flows at the stem base likely peak in response to maximum crown illuminance (and transpiration) and this will differ compared to the timing of peak flows in upper branches. We suggest that the magnitude of flow lags between branches and stems needs further study, with careful attention to branch position and method application before a robust understanding of capacitance, particularly in woody tissues of large trees, can be formed. We did not detect flow lags in the world’s tallest and largest tree species Sequoia sempervirens and Sequoiadendron giganteum, despite measurement along large pathlengths (∼57 and 85 m), which raises questions as to why large flow lags are often recorded for much smaller species. One conspicuous possibility is the different methods used among studies. Constant-heating methods such as the thermal dissipation probe (and also heat balance methods) include heat capacitance behaviour due to warming of wood tissues, which delays the response of the sensors to changing sap flow conditions. We argue that methods with intrinsic heat-capacitance present dangers when trying to measure water-capacitance in trees. In this respect heat pulse methods hold an advantage.

Summary • Leaf morphology and physiological functioning demonstrate considerable plasticity within tree crowns, with various leaf traits often exhibiting pronounced vertical gradients in very tall trees. It has been proposed that the trajectory of these gradients, as determined by regression methods, could be used in conjunction with theoretical biophysical limits to estimate the maximum height to which trees can grow. • Here, we examined this approach using published and new experimental data from tall conifer and angiosperm species. • We showed that height predictions were sensitive to tree-to-tree variation in the shape of the regression and to the biophysical endpoints selected. We examined the suitability of proposed end-points and their theoretical validity. We also noted that site and environment influenced height predictions considerably. • Use of leaf mass per unit area or leaf water potential coupled with vulnerability of twigs to cavitation poses a number of difficulties for predicting tree height. Photosynthetic rate and carbon isotope discrimination show more promise, but in the second case, the complex relationship between light, water availability, photosynthetic capacity and internal conductance to CO2 must first be characterized.

Woody plant encroachment into semiarid ecosystems has become a global trend in recent decades. Due to stream channel incision, the semiarid riparian montane meadows of the southern Sierra Nevada Mountains, USA are experiencing long-term declines in soil moisture. A woody shrub, Artemisia rothrockii A. Gray (Rothrock sagebrush, Asteraceae) is invading these herbaceous meadows. We used an analysis of the stable oxygen isotope ratios of plant and soil water to measure the depth of plant water acquisition during the early stages of this woody plant encroachment. Sagebrush used deeper water on average than most herbs, but it also acquired 10–30% of its water from shallow (

Summary Under well-watered conditions leaf hydraulic conductance increases with transpiration rate. This reduces the water potential gradient in leaves and potentially improves productivity under daily variation in evaporative demand., Leaf hydraulic conductance (k leaf) is a central element in the regulation of leaf water balance but the properties of k leaf remain uncertain. Here, the evidence for the following two models for k leaf in well-hydrated plants is evaluated: (i) k leaf is constant or (ii) k leaf increases as transpiration rate (E) increases. The difference between stem and leaf water potential (ΔΨstem–leaf), stomatal conductance (g s), k leaf, and E over a diurnal cycle for three angiosperm and gymnosperm tree species growing in a common garden, and for Helianthus annuus plants grown under sub-ambient, ambient, and elevated atmospheric CO2 concentration were evaluated. Results show that for well-watered plants k leaf is positively dependent on E. Here, this property is termed the dynamic conductance, k leaf(E), which incorporates the inherent k leaf at zero E, which is distinguished as the static conductance, k leaf(0). Growth under different CO2 concentrations maintained the same relationship between k leaf and E, resulting in similar k leaf(0), while operating along different regions of the curve owing to the influence of CO2 on g s. The positive relationship between k leaf and E minimized variation in ΔΨstem–leaf. This enables leaves to minimize variation in Ψleaf and maximize g s and CO2 assimilation rate over the diurnal course of evaporative demand.

Water limitation is one of the most important factors limiting crop productivity world-wide and has likely been an important selective regime influencing the evolution of plant physiology. Understanding the genetic and physiological basis of drought adaptation is therefore important for improving crops as well as for understanding the evolution of wild species. Here, results are presented from quantitative trait loci (QTL) mapping of flowering time (a drought escape mechanism) and carbon stable isotope ratio (¿13C) (a drought-avoidance mechanism) in Arabidopsis thaliana. Whole-genome scans were performed using multiple-QTL models for both additive and epistatic QTL effects. We mapped five QTL affecting flowering time and five QTL affecting ¿13C, but two genomic regions contained QTL with effects on both traits, suggesting a potential pleiotropic relationship. In addition, we observed QTL¿QTL interaction for both traits. Two ¿13C QTL were captured in near-isogenic lines to further characterize their physiological basis. These experiments revealed allelic effects on ¿13C through the upstream trait of stomatal conductance with subsequent consequences for whole plant transpiration efficiency and water loss. Our findings document considerable natural genetic variation in whole-plant, drought resistance physiology of Arabidopsis and highlight the value of quantitative genetic approaches for exploring functional relationships regulating physiology.

Watersheds of the Catskill Mountains, New York have marked differences in nitrogen (N) dynamics among dominant tree species stands. Our objectives were to study how tree species vary in N uptake to better understand the basis for the observed variation in these forested watersheds. We conducted a 15N tracer greenhouse study to determine NH4 + and NO3 − uptake of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga Canadensis L.), red oak (Quercus rubra L.) and sugar maple (Acer saccharum Marsh.) seedlings. Seedlings and their native soil were collected in November 1997, over-wintered and allowed to break dormancy in spring 1998. Half of the seedlings of each tree species received 15NH4-NO3 to examine NH4 + uptake and the other half received NH4-15NO3 to examine NO3 − uptake. Plants were harvested 4 days following 15N addition. Tree species varied in their preference for NH4 + and NO3 −. Sugar maple and eastern hemlock seedlings took up more NH4 + than NO3 − per unit plant biomass, while beech was the only species to take up more NO3 − than NH4 +. Red oak took up more NH4 + than NO3 − into roots, stems and leaves, but the difference between the two forms of N was not statistically significant. These results demonstrate that tree species of the Catskill Mountains vary in their capacity to take up NH4 + and NO3 −. Coupled with stand-level studies of N dynamics, this variation can help explain some of the patterns of forested watershed N retention and loss in the Catskill Mountains shown in our field investigations (Templer, 2001; Templer et al., in press Ecosystems).

The early ecophysiological trajectories of angiosperm evolution remain uncertain. This hinders resolution of the possible ecological and physiological catalysts that set the stage for the emergence of our modern flora. One approach for generating hypotheses on the early ecology of angiosperms is to consider the distribution of functional traits among lineages that diverge near the root of the extant angiosperm phylogenetic tree. Using recent phylogenetic results to choose lineages for comparative work, we review and expand on previous studies that examine the ecology and physiology of basal angiosperm lineages. Phylogenetic mapping of functional traits reveals that the basal lineages, Amborella, Austrobaileyales, and some Chloranthaceae, share ecological and physiological traits linked to shady, disturbed, and possibly wet habitats. Traits include low and easily light‐saturated photosynthetic rates, absence of leaf palisade cells, guttation resulting from root pressure, as well as small seed sizes. Some C...

Many species have the ability to resprout vegetatively after a substantial loss of biomass induced by environmental stress, including drought. Many of the regions characterised by ecosystems where resprouting is common are projected to experience more frequent and intense drought during the 21st Century. However, in assessments of ecosystem response to drought disturbance there has been scant consideration of the resilience and post-drought recovery of resprouting species. Systematic differences in hydraulic and allocation traits suggest that resprouting species are more resilient to drought-stress than nonresprouting species. Evidence suggests that ecosystems dominated by resprouters recover from disturbance more quickly than ecosystems dominated by nonresprouters. The ability of resprouters to avoid mortality and withstand drought, coupled with their ability to recover rapidly, suggests that the impact of increased drought stress in ecosystems dominated by these species may be small. The strategy of resprouting needs to be modelled explicitly to improve estimates of future climate-change impacts on the carbon cycle, but this will require several important knowledge gaps to be filled before resprouting can be properly implemented.

We compared the physiology and growth of seedlings originating from different Sequoia sempervirens (D. Don.) Endl. (coast redwood) and Sequoiadendron giganteum (Lindl.) Buchh. (giant sequoia) populations subjected to progressive drought followed by a recovery period in a controlled greenhouse experiment. Our objective was to examine how multiple plant traits interact to influence the response of seedlings of each species and seed population to a single drought and recovery cycle. We measured soil and plant water status, leaf gas exchange, stem embolism and growth of control (well-watered) and drought-stressed (water withheld) seedlings from each population at the beginning, middle and end of a 6-week drought period and again 2 weeks after re-watering. The drought had a significant effect on many aspects of seedling performance, but water-stressed seedlings regained most physiological functioning by the end of the recovery period. Sequoiadendron seedlings exhibited a greater degree of isohydry (water status regulation), lower levels of stem embolism, higher biomass allocation to roots and lower sensitivity of growth to drought compared with Sequoia. Only minor intra-specific differences were observed among populations. Our results show that seedlings of the two redwood species exhibit contrasting drought-response strategies that align with the environmental conditions these trees experience in their native habitats, and demonstrate trade-offs and coordination among traits affecting plant water use, carbon gain and growth under drought.

Sap flows in the xylem of plant roots in response to gradients in water potential, either between soil and atmosphere (transpiration) or soil layers of different moisture content (termed hydraulic redistribution). The latter has the potential to influence water budgets and species interactions, but we lack information for all but a few plant communities. We combined heat pulse measurements of sap flow with dye and isotope tracing techniques to gauge the movement of xylem sap within, and exudation from, roots of Banksia prionotes (Lindley). We demonstrated ‘ hydraulic lift’ during the dry season and provide some evidence that extremely dry soils limit hydraulic lift. In addition we report difficulties posed by spiralled xylem tissue in roots for the application of heat pulse techniques.

The purpose of this study was to use split-root chambers to determine whether sunflowers (Helianthus annuus), which possess high transpirational demand, can preferentially extract macropore water when they are given the option of using either micropore or macropore water. Sunflower plants were grown in split-root chambers with half of their roots in a macroporous soil and half in a microporous soil of identical mineralogy. The two chambers were irrigated with water of different stable hydrogen isotope compositions (δD) (tap water and melted snow water). By measuring the δD of the sunflower xylem sap, it was possible to determine from which soil type the sunflowers were extracting water. It was found that sunflowers did not preferentially extract macropore water. Since sunflower plants possess very high transpiration rates on a whole-plant basis, and because transpiration remains high even under soil drying, we believe sunflower roots must have a consistently very low water potential compared with the soil water potentials of either the macroporous or microporous soils. If root water potentials are sufficiently low, then root water uptake cannot discriminate between macropores and micropores.

Early angiosperm evolution, beginning approximately 140 million years ago, saw many innovations that enabled flowering plants to alter ecosystems globally. These included the development of novel, flower-based pollinator attraction mechanisms and the development of increased water transport capacity in stems and leaves. Vein length per area (VLA) of leaves increased nearly threefold in the first 30–40 million years of angiosperm evolution, increasing the capacity for transpiration and photosynthesis. In contrast to leaves, high water transport capacities in flowers may not be an advantage because flowers do not typically contribute to plant carbon gain. Although flowers of extant basal angiosperms are hydrated by the xylem, flowers of more recently derived lineages may be hydrated predominantly by the phloem. In the present study, we measured leaf and flower VLA for a phylogenetically diverse sample of 132 species from 52 angiosperm families to ask (i) whether flowers have lower VLA than leaves, (ii) whether flowers of basal angiosperm lineages have higher VLA than more recently derived lineages because of differences between xylem and phloem hydration, and (iii) whether flower and leaf VLA evolved independently. It was found that floral structures had lower VLA than leaves, but basal angiosperm flowers did not have higher VLA than more derived lineages. Furthermore, the independent evolution of leaf and petal VLA suggested that these organs may be developmentally modular. Unlike leaves, which have experienced strong selection for increased water transport capacity, flowers may have been shielded from such selective pressures by different developmental processes controlling VLA throughout the plant bauplan.

The hypothesis tested in this paper is that, because the freshest water occurs in the largest soil pores (macropores), plants of low to moderate transpiration rate can survive in salinized soil because they preferentially extract water from macropores. The hypothesis predicts that a plant growing in a macroporous soil should have greater growth under a given salinity treatment than a similar plant growing in a soil with the same mineralogy but without macropores. This hypothesis was tested by growing bell pepper (Capsicum annuum) in the greenhouse in pots filled with either a commercial fritted clay (a highly macroporous soil) or the same clay ground to a finer texture and sieved to remove macropores and produce a microporous soil. The pots sat in pans filled with salt water. Half of the pots were irrigated once a day with fresh water and the other half received no fresh water. Plants growing in the macroporous soil had greater growth for a given salinity treatment than the plants growing in the microporous soil under both the irrigated and non-irrigated conditions. Under the irrigated condition for the highest salinity treatment, the non-reproductive fresh weight per plant, total dry weight per plant and fruit fresh weight per plant was 114 g, 12 g and 50 g, respectively, for the macroporous soil and 47 g, 4.5 g and 5 g, respectively, for the microporous soil. The results of this study provide evidence to suggest that a better understanding of what constitutes a ‘good’ structure in a saline soil may aid us in our efforts to improve the management of saline soils. We suggest that it may be possible to increase the agricultural production on salinized land by no-tillage agriculture which preserves macroporosity. Possible obstacles could be the tendency of field saline-sodic soils to swell and the unavailability of relatively fresh irrigation water in areas with saline soils.

Use of soil water and groundwater by open-grown Acer saccharum Marsh. (sugar maple) tree canopies and forests was estimated by measuring transpiration (E) rates using porometry, sap flow methods, and the Bowen ratio method. The Bowen ratio and sap flow methods showed the best agreement; porometer measurements scaled to whole canopies always underestimated E by 15-50%. Trees of different sizes showed very different rates of E. I hypothesized that these differences were due to the differential access of large and small trees to groundwater and soil water, respectively. Transpirational flux was partitioned between soil water and groundwater by tracing the water sources based on their hydrogen stable isotopic composition (deltaD). Soil water deltaD varied between -41 and -16 per thousand seasonally (May to September), whereas groundwater deltaD was -79 +/- 5 per thousand during the entire growing season. Daily transpiration rates of large (9-14 m tall) trees were significantly higher than those of small (3-5 m tall) trees (2.46-6.99 +/- 1.02-2.50 versus 0.69-1.80 +/- 0.39-0.67 mm day(-1)). Small trees also showed greater variation in E during the growing season than large trees. In addition, compared to the large trees, small trees demonstrated greater sensitivity to environmental factors that influence E, such as soil water deficits and increased evaporative demand. Over the entire growing season, large trees and forest stands composed of trees > 10 m tall transpired only groundwater. The high rates of water loss from large trees and older forests were likely a result of the influence of an enhanced "pool" of transpirational water in the upper soil layers caused by hydraulic lift (see Dawson 1993b). The hydraulically lifted water reservoir enabled large trees to use more potential transpirational water during daylight hours than small trees, leading to a greater total water flux. In contrast, small trees and forest stands composed of younger trees almost exclusively used soil water, except during two dry periods when their transpirational water was composed of between 7 and 17% groundwater. Thus groundwater discharge from sugar maple trees and forest stands of different sizes (ages) differs significantly, and large trees and older forest stands have a greater impact on the hydrologic balance of groundwater than small trees and younger forest stands. However, mixed stands (small and large trees) may have a greater overall impact on the regional hydrologic balance than old stands, because trees in mixed stands draw on both soil water and groundwater reservoirs and thus can substantially increase total water discharge on scales from tens to hundreds of hectares.

There has been limited application of sapflow technology to small-stemmed species and across co-existing functional types, restricting its use in diverse floras such as the Mediterranean-type shrubland in South Africa. Our main objective was to test whether sapflow may provide an alternative to traditional gas-exchange measurements, which would permit comparative evaluation of transpiration at a previously unattained temporal resolution. We tested miniature external heat ratio method (HRM) sapflow gauges on three co-occurring functional types with contrasting stem or culm anatomies and examined the relationship between sapflow and shoot- and leaf-level water loss in both a controlled and field environment. Our sapflow gauges captured dynamic patterns of transpiration in both settings for all three functional types. In a controlled environment the relationship between sapflow and transpiration was linear in all three species with r2 values ranging from 0.78 for Cannomois congesta Mast. (Restionaceae) to 0.96 for Protea repens (L.) L. (Proteaceae) and Erica monsoniana L.f. (Ericaceae). In the field, r2 values were lower, ranging from 0.59 for C. congesta to 0.74 for P. repens. We discuss the efficacy and potential of this methodology to cast light on patterns of community ecology in functionally diverse shrublands by capturing continuous variation in transpiration.