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At least 19 records

Dry and hot: the hydraulic consequences of a climate change-type drought for Amazonian trees

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016 an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SM), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapor pressure deficit but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation, and stomatal closure. Our results suggest extremely warm droughts greatly amplify the degree of trees’ physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.

Fontes, Clarissa G.↗

The xylem of anisohydric Quercus alba L. is more vulnerable to embolism than isohydric codominants

Abstract The coordination of plant leaf water potential (Ψ L ) regulation and xylem vulnerability to embolism is fundamental for understanding the tradeoffs between carbon uptake and risk of hydraulic damage. There is a general consensus that trees with vulnerable xylem more conservatively regulate Ψ L than plants with resistant xylem. We evaluated if this paradigm applied to three important eastern US temperate tree species, Quercus alba L., Acer saccharum Marsh. and Liriodendron tulipifera L., by synthesizing 1600 Ψ L observations, 122 xylem embolism curves and xylem anatomical measurements across 10 forests spanning pronounced hydroclimatological gradients and ages. We found that, unexpectedly, the species with the most vulnerable xylem ( Q. alba ) regulated Ψ L less strictly than the other species. This relationship was found across all sites, such that coordination among traits was largely unaffected by climate and stand age. Quercus species are perceived to be among the most drought tolerant temperate US forest species; however, our results suggest their relatively loose Ψ L regulation in response to hydrologic stress occurs with a substantial hydraulic cost that may expose them to novel risks in a more drought‐prone future.

Benson, Michael C.↗

Lianas and Trees From a Seasonally Dry and a Wet Tropical Forest Did Not Differ in Embolism Resistance but Did Differ in Xylem Anatomical Traits in the Dry Forest

One of the most prominent changes in neotropical forests has been the increase in abundance and size of lianas. Studies suggest that lianas have more acquisitive strategies than trees, which could allow them to take advantage of water more effectively when it is available in water-limited forests, but few studies compared across growth form (i.e., lianas vs. trees) and forest type (i.e., wet vs. seasonally dry). We measured hydraulic and anatomical traits of co-occurring lianas and trees that convey drought resistance (xylem embolism resistance and intervessel pit membranes) and water transport capacity (xylem vessel diameter and density) in a seasonally dry and a wet evergreen tropical forest to address: (1) Are there differences between vulnerability to embolisms (P 50 —water potential at 50% loss of hydraulic conductivity) and hydraulic safety margins (HSM) across growth form and forest type? (2) How do vessel diameter and density vary across growth form and forest type? (3) Are there differences in xylem intervessel pit membrane thickness across growth form and forest type and does it predict xylem embolism vulnerability in trees and lianas? We examined hydraulic and xylem anatomical traits of 32 species—eight lianas and eight trees in each forest type. We found no difference in P 50 and HSMs between lianas and trees and between the wetter and drier forest. Dry forest lianas had 81% greater maximum vessel diameter and 125% greater range in vessel diameter sizes than dry forest trees but, there was no significant difference between life forms in the wet forest. Dry forest species had 50% greater vessel density and 30% greater maximum pit membrane thickness than wet forest ones. Maximum pit membrane thickness was correlated to P 50 and HSMs. The main difference between lianas and trees occurred in the dry forest, where lianas had larger maximum xylem vessel size than trees, implying that they have proportionally greater hydraulic conductive capacity than the trees in seasonal forests.

Smith-Martin, Chris M.↗

Basin-wide variation in tree hydraulic safety margins predicts the carbon balance of Amazon forests

Abstract Tropical forests face increasing climate risk 1,2 , yet our ability to predict their response to climate change is limited by poor understanding of their resistance to water stress. Although xylem embolism resistance thresholds (for example, $$\varPsi $$ Ψ 50 ) and hydraulic safety margins (for example, HSM 50 ) are important predictors of drought-induced mortality risk 3–5 , little is known about how these vary across Earth’s largest tropical forest. Here, we present a pan-Amazon, fully standardized hydraulic traits dataset and use it to assess regional variation in drought sensitivity and hydraulic trait ability to predict species distributions and long-term forest biomass accumulation. Parameters $$\varPsi $$ Ψ 50 and HSM 50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both $$\varPsi $$ Ψ 50 and HSM 50 influence the biogeographical distribution of Amazon tree species. However, HSM 50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM 50 are gaining more biomass than are low HSM 50 forests. We propose that this may be associated with a growth–mortality trade-off whereby trees in forests consisting of fast-growing species take greater hydraulic risks and face greater mortality risk. Moreover, in regions of more pronounced climatic change, we find evidence that forests are losing biomass, suggesting that species in these regions may be operating beyond their hydraulic limits. Continued climate change is likely to further reduce HSM 50 in the Amazon 6,7 , with strong implications for the Amazon carbon sink.

Science & Technology - Other Topics↗

Living on the edge: A continental-scale assessment of forest vulnerability to drought

Globally, forests are facing an increasing risk of mass tree mortality events associated with extreme droughts and higher temperatures. Hydraulic dysfunction is considered a key mechanism of drought-triggered dieback. By leveraging the climate breadth of the Australian landscape and a national network of research sites (Terrestrial Ecosystem Research Network), we conducted a continental-scale study of physiological and hydraulic traits of 33 native tree species from contrasting environments to disentangle the complexities of plant response to drought across communities. Here, we found strong relationships between key plant hydraulic traits and site aridity. Leaf turgor loss point and xylem embolism resistance were correlated with minimum water potential experienced by each species. Across the data set, there was a strong coordination between hydraulic traits, including those linked to hydraulic safety, stomatal regulation and the cost of carbon investment into woody tissue. These results illustrate that aridity has acted as a strong selective pressure, shaping hydraulic traits of tree species across the Australian landscape. Hydraulic safety margins were constrained across sites, with species from wetter sites tending to have smaller safety margin compared with species at drier sites, suggesting trees are operating close to their hydraulic thresholds and forest biomes across the spectrum may be susceptible to shifts in climate that result in the intensification of drought.

59 BASIC BIOLOGICAL SCIENCES↗

A heuristic classification of woody plants based on contrasting shade and drought strategies

Woody plants vary in their adaptations to drought and shade. For a better prediction of vegetation responses to drought and shade within dynamic global vegetation models, it is critical to group species into functional types with similar adaptations. One of the key challenges is that the adaptations are generally determined by a large number of plant traits that may not be available for a large number of species. In this study, we present two heuristic woody plant groups that were separated using cluster analysis in a three-dimensional trait–environment space based on three key metrics for each species: mean xylem embolism resistance, shade tolerance and habitat aridity. The two heuristic groups separate these species into tolerators and avoiders. Furthermore, the tolerators either rely on their high embolism resistance to tolerate drought in arid habitats (e.g., Juniperus and Prunus ) or rely on high shade tolerance to withstand shaded conditions in wet habitats (e.g., Picea , Abies and Acer ). In contrast, all avoiders have low embolism resistance and low shade tolerance. In arid habitats, avoiders tend to minimize catastrophic embolism (e.g., most Pinus species) while in wet habitats, they may survive despite low shade tolerance (e.g., Betula , Populus , Alnus and Salix ). Because our approach links traits to the environmental conditions, we expect it could be a promising framework for predicting changes in species composition, and therefore ecosystem function, under changing environmental conditions.

59 BASIC BIOLOGICAL SCIENCES↗

Leaf Pressure Volume Data in Caxiuana and Tapajos National Forest, Para, Brazil (2011)

Pressure volume curve measurements on leaves of canopy trees from the from the Caxiuana and Tapajos National Forests, Para, Brazil. Tapajos samples were harvested from the km 67 forested area, which is adjacent to the decommissioned throughfall exclusion drought experimental plot. Caxiuana samples were harvested from trees growing in the throughfall exclusion plots. Data were collected in 2011. Dataset includes: date of measurement, site ID, plot ID, tree ID (species, tree tag #), leaf area, fresh weight, relative weight, leaf water potential, and leaf water loss. P-V curve parameters (turgor loss point, osmotic potential, and bulk modulus of elasticity) can be found in Powell et al. (2017) Differences in xylem cavitation resistance and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees. Global Change Biology.

54 ENVIRONMENTAL SCIENCES↗

Xylem vulnerability curves of canopy branches of mature trees from Caxiuana and Tapajos National Forests, Para, Brazil

Raw data for xylem vulnerability curves measured on upper canopy branches of mature trees from the Caxiuana and Tapajos National Forests, Para, Brazil. Tapajos samples were harvested from km67 transects, which is nearby the decommissioned throughfall-exclusion, drought-experiment plots. Caxiuana samples were harvested from trees growing in the throughfall-exclusion, drought-experiment plots. Data were collected in 2011 and 2012. Dataset includes: date of measurement, site ID, plot ID, tree ID (species, tree tag #), xylem pressure, percent loss of conductivity. Air injection method was used. Data reference: Powell et al. (2017) Differences in xylem cavitation resistance and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees. Global Change Biology.

54 ENVIRONMENTAL SCIENCES↗

Vascular defense responses in rice: peroxidase accumulation in xylem parenchyma cells and xylem wall thickening

The rice bacterial blight pathogen Xanthomonas oryzae pv. oryzae is a vascular pathogen that elicits a defensive response through interaction with metabolically active rice cells. In leaves of 12-day-old rice seedlings, the exposed pit membrane separating the xylem lumen from the associated parenchyma cells allows contact with bacterial cells. During resistant responses, the xylem secondary walls thicken within 48 h and the pit diameter decreases, effectively reducing the area of pit membrane exposed for access by bacteria. In susceptible interactions and mock-inoculated controls, the xylem walls do not thicken within 48 h. Xylem secondary wall thickening is developmental and, in untreated 65-day-old rice plants, the size of the pit also is reduced. Activity and accumulation of a secreted cationic peroxidase, PO-C1, were previously shown to increase in xylem vessel walls and lumen. Peptide-specific antibodies and immunogold-labeling were used to demonstrate that PO-C1 is produced in the xylem parenchyma and secreted to the xylem lumen and walls. The timing of the accumulation is consistent with vessel secondary wall thickening. The PO-C1 gene is distinct but shares a high level of similarity with previously cloned pathogen-induced peroxidases in rice. PO-C1 gene expression was induced as early as 12 h during resistant interactions and peaked between 18 and 24 h after inoculation. Expression during susceptible interactions was lower than that observed in resistant interactions and was undetectable after infiltration with water, after mechanical wounding, or in mature leaves. These data are consistent with a role for vessel secondary wall thickening and peroxidase PO-C1 accumulation in the defense response in rice to X. oryzae pv. oryzae.

NASA Discipline Plant Biology↗

Optimization of leaf morphology in relation to leaf water status: A theory

The leaf economic traits such as leaf area, maximum carbon assimilation rate, and venation are all correlated and related to water availability. Furthermore, leaves are often broad and large in humid areas and narrower in arid/semiarid and hot and cold areas. We use optimization theory to explain these patterns. We have created a constrained optimization leaf model linking leaf shape to vein structure that is integrated into coupled transpiration and carbon assimilation processes. The model maximizes net leaf carbon gain (NPPleaf) over the loss of xylem water potential. Modeled relations between leaf traits are consistent with empirically observed patterns. As the results of the leaf shape-venation relation, our model further predicts that a broadleaf has overall higher NPPleaf compared to a narrowleaf. In addition, a broadleaf has a lower stomatal resistance compared to a narrowleaf under the same level of constraint. With the same leaf area, a broadleaf will have, on average, larger conduits and lower total leaf xylem resistance and thus be more efficient in water transportation but less resistant to cavitation. By linking venation structure to leaf shape and using water potential as the constraint, our model provides a physical explanation for the general pattern of the covariance of leaf traits through the safety-efficiency trade-off of leaf hydraulic design.

59 BASIC BIOLOGICAL SCIENCES↗

Quantification of hydraulic trait control on plant hydrodynamics and risk of hydraulic failure within a demographic structured vegetation model in a tropical forest (FATES–HYDRO V1.0)

Abstract. Vegetation plays a key role in the global carbon cycle and thus is an important component within Earth system models (ESMs) that project future climate. Many ESMs are adopting methods to resolve plant size and ecosystem disturbance history, using vegetation demographic models. These models make it feasible to conduct more realistic simulation of processes that control vegetation dynamics. Meanwhile, increasing understanding of the processes governing plant water use, and ecosystem responses to drought in particular, has led to the adoption of dynamic plant water transport (i.e., hydrodynamic) schemes within ESMs. However, the extent to which variations in plant hydraulic traits affect both plant water stress and the risk of mortality in trait-diverse tropical forests is understudied. In this study, we report on a sensitivity analysis of an existing hydrodynamic scheme (HYDRO) model that is updated and incorporated into the Functionally Assembled Terrestrial Ecosystem Simulator (FATES) (FATES–HYDRO V1.0). The size- and canopy-structured representation within FATES is able to simulate how plant size and hydraulic traits affect vegetation dynamics and carbon–water fluxes. To better understand this new model system, and its functionality in tropical forest systems in particular, we conducted a global parameter sensitivity analysis at Barro Colorado Island, Panama. We assembled 942 observations of plant hydraulic traits on 306 tropical plant species for stomata, leaves, stems, and roots and determined the best-fit statistical distribution for each trait, which was used in model parameter sampling to assess the parametric sensitivity. We showed that, for simulated leaf water potential and loss of hydraulic conductivity across different plant organs, the four most important traits were associated with xylem conduit taper (buffers increasing hydraulic resistance with tree height), stomatal sensitivity to leaf water potential, maximum stem hydraulic conductivity, and the partitioning of total hydraulic resistance above vs. belowground. Our analysis of individual ensemble members revealed that trees at a high risk of hydraulic failure and potential tree mortality generally have a lower conduit taper, lower maximum xylem conductivity, lower stomatal sensitivity to leaf water potential, and lower resistance to xylem embolism for stem and transporting roots. We expect that our results will provide guidance on future modeling studies using plant hydrodynamic models to predict the forest responses to droughts and future field campaigns that aim to better parameterize plant hydrodynamic models.

54 ENVIRONMENTAL SCIENCES↗

Xylem-dwelling pathogen unaffected by local xylem vessel network properties in grapevines ( Vitis spp.)

Abstract Background and aims Xylella fastidiosa (Xf) is the xylem-dwelling bacterium associated with Pierce’s disease (PD), which causes mortality in agriculturally important species, such as grapevine (Vitis vinifera). The development of PD symptoms in grapevines depends on the ability of Xf to produce cell-wall-degrading enzymes to break up intervessel pit membranes and systematically spread through the xylem vessel network. Our objective here was to investigate whether PD resistance could be mechanistically linked to xylem vessel network local connectivity. Methods We used high-resolution X-ray micro-computed tomography (microCT) imaging to identify and describe the type, area and spatial distribution of intervessel connections for six different grapevine genotypes from three genetic backgrounds, with varying resistance to PD (four PD resistant and two PD susceptible). Key results Our results suggest that PD resistance is unlikely to derive from local xylem network connectivity. The intervessel pit area (Ai) varied from 0.07 ± 0.01 mm2 mm−3 in Lenoir to 0.17 ± 0.03 mm2 mm−3 in Blanc do Bois, both PD resistant. Intervessel contact fraction (Cp) was not statically significant, but the two PD-susceptible genotypes, Syrah (0.056 ± 0.015) and Chardonnay (0.041 ± 0.013), were among the most highly connected vessel networks. Neither Ai nor Cp explained differences in PD resistance among the six genotypes. Bayesian re-analysis of our data shows moderate evidence against the effects of the traits analysed: Ai (BF01 = 4.88), mean vessel density (4.86), relay diameter (4.30), relay density (3.31) and solitary vessel proportion (3.19). Conclusions Our results show that radial and tangential xylem network connectivity is highly conserved within the six different Vitis genotypes we sampled. The way that Xf traverses the vessel network may limit the importance of local network properties to its spread and may confer greater importance on host biochemical responses.

Plant Sciences↗

A test of the hydraulic vulnerability segmentation hypothesis in angiosperm and conifer tree species

Water transport from soils to the atmosphere is critical for plant growth and survival. However, we have a limited understanding about many portions of the whole-tree hydraulic pathway, because the vast majority of published information is on terminal branches. Our understanding of mature tree trunk hydraulic physiology, in particular, is limited. The hydraulic vulnerability segmentation hypothesis (HVSH) stipulates that distal portions of the plant (leaves, branches and roots) should be more vulnerable to embolism than trunks, which are non-redundant organs that require a massive carbon investment. In the current study, we compared vulnerability to loss of hydraulic function, leaf and xylem water potentials and the resulting hydraulic safety margins (in relation to the water potential causing 50% loss of hydraulic conductivity) in leaves, branches, trunks and roots of four angiosperms and four conifer tree species. Across all species, our results supported strongly the HVSH as leaves and roots were less resistant to embolism than branches or trunks. However, branches were consistently more resistant to embolism than any other portion of the plant, including trunks. Also, calculated whole-tree vulnerability to hydraulic dysfunction was much greater than vulnerability in branches. This was due to hydraulic dysfunction in roots and leaves at less negative water potentials than those causing branch or trunk dysfunction. Leaves and roots had narrow or negative hydraulic safety margins, but trunks and branches maintained positive safety margins. By using branch-based hydraulic information as a proxy for entire plants, much research has potentially overestimated embolism resistance, and possibly drought tolerance, for many species. This study highlights the necessity to reconsider past conclusions made about plant resistance to drought based on branch xylem only. As a result, this study also highlights the necessity for more research of whole-plant hydraulic physiology to better understand strategies of plant drought tolerance and the critical control points within the hydraulic pathway.

59 BASIC BIOLOGICAL SCIENCES↗

Gravitropism of cucumber hypocotyls: biophysical mechanism of altered growth

The biophysical basis for the changes in cell elongation rate during gravitropism was examined in aetiolated cucumber (Cucumis sativus L.) hypocotyls. Bulk osmotic pressures on the two sides of the stem and in the epidermal cells were not altered during the early time course of gravitropism. By the pressure-probe technique, a small increase in turgor (0.3 bar, 30 kPa) was detected on the upper (inhibited) side, whereas there was a negligible decrease in turgor on the lower (stimulated) side. These small changes in turgor and water potential appeared to be indirect, passive consequences of the altered growth and the small resistance for water movement from the xylem, and indicated that the change in growth was principally due to changes in wall properties. The results indicate that the hydraulic conductance of the water-transport pathway was large (.25 h-1 bar-1) and the water potential difference supporting cell expansion was no greater than 0.3 bar (30 kPa). From pressure-block experiments, it appeared that upon gravitropic stimulation (1) the yield threshold of the lower half of the stem did not decrease and (2) the wall on the upper side of the stem was not made more rigid by a cross-linking process. Mechanical measurements of the stress/strain properties of the walls showed that the initial development of gravitropism did not involve an alteration of the mechanical behaviour of the isolated walls. Thus, gravitropism in cucumber hypocotyls occurs principally by an alteration of the wall relaxation process, without a necessary change in wall mechanical properties.

Non-NASA Center↗

Xylem network connectivity and embolism spread in grapevine( Vitis vinifera L.)

Abstract Xylem networks are vulnerable to the formation and spread of gas embolisms that reduce water transport. Embolisms spread through interconduit pits, but the three-dimensional (3D) complexity and scale of xylem networks means that the functional implications of intervessel connections are not well understood. Here, xylem networks of grapevine (Vitis vinifera L.) were reconstructed from 3D high-resolution X-ray micro-computed tomography (microCT) images. Xylem network performance was then modeled to simulate loss of hydraulic conductivity under increasingly negative xylem sap pressure simulating drought stress conditions. We also considered the sensitivity of xylem network performance to changes in key network parameters. We found that the mean pit area per intervessel connection was constant across 10 networks from three, 1.5-m stem segments, but short (0.5 cm) segments fail to capture complete network connectivity. Simulations showed that network organization imparted additional resistance to embolism spread beyond the air-seeding threshold of pit membranes. Xylem network vulnerability to embolism spread was most sensitive to variation in the number and location of vessels that were initially embolized and pit membrane vulnerability. Our results show that xylem network organization can increase stem resistance to embolism spread by 40% (0.66 MPa) and challenge the notion that a single embolism can spread rapidly throughout an entire xylem network.

Wason, Jay↗

Hydraulic conductance, resistance, and resilience: how leaves of a tropical epiphyte respond to drought

Premise Because of its broad range in the neotropical rainforest and within tree canopies, the tank bromeliad Guzmania monostachia was investigated as a model of how varying leaf hydraulic conductance ( K leaf ) could help plants resist and recover from episodic drought. The two pathways of K leaf , inside and outside the xylem, were also examined to determine the sites and causes of major hydraulic resistances within the leaf. Methods We measured leaf hydraulic conductance for plants in the field and laboratory under wet, dry, and rewetted conditions and applied physiological, anatomical, and gene expression analysis with modeling to investigate changes in K leaf . Results After 7 d with no rain in the field or 14 days with no water in the glasshouse, K leaf decreased by 50% yet increased to hydrated values within 4 d of tank refilling. Staining to detect embolism combined with modeling indicated that changes outside the xylem were of greater importance to K leaf than were changes inside the xylem and were associated with changes in intercellular air spaces (aerenchyma), aquaporin expression and inhibition, and cuticular conductance. Conclusions Low values for all conductances during drying, particularly in pathways outside the xylem, lead to hydraulic resilience for this species and may also contribute to its broad environmental tolerances.

North, Gretchen B.↗

Embolism spread in the primary xylem of Polystichum munitum : implications for water transport during seasonal drought

Abstract Xylem network structure and function have been characterized for many woody plants, but less is known about fern xylem, particularly in species endemic to climates where water is a limiting resource for months at a time. We characterized seasonal variability in soil moisture and frond water status in a common perennial fern in the redwood understory of a costal California, and then investigated the consequences of drought‐induced embolism on vascular function. Seasonal variability in air temperature and soil water content was minimal, and frond water potential declined slowly over the observational period. Our data show that Polystichum munitum was protected from significant drought‐induced hydraulic dysfunction during this growing season because of a combination of cavitation resistant conduits (Air‐seeding threshold (ASP) = −1.53 MPa; xylem pressure inducing 50% loss of hydraulic conductivity ( P 50 ) = −3.02 MPa) and a soil with low moisture variability. High resolution micro‐computed tomography (MicroCT) imaging revealed patterns of embolism formation in vivo for the first time in ferns providing insight into the functional status of the xylem network under drought conditions. Together with stomatal conductance measurements, these data suggest that P . munitum is adapted to tolerate drier conditions than what was observed during the growing season.

Brodersen, Craig R.↗