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Classifying the diverse ways that plants respond to hydrologic stress into generalizable ‘water‐use strategies’ has long been an eco‐physiological research goal. While many schemes for describing water‐use strategies have proven to be quite useful, they are also associated with uncertainties regarding their theoretical basis and their connection to plant carbon and water relations. In this review, we discuss the factors that shape plant water stress responses and assess the approaches used to classify a plant's water‐use strategy, paying particular attention to the popular but controversial concept of a continuum from isohydry to anisohydry.

A generalizable and predictive framework for assessing plant water‐use strategies has been historically elusive, yet recent advances in plant physiology and hydraulics provide the field with a way past these obstacles. Specifically, we promote the idea that many metrics that quantify water‐use strategies are highly dynamic and emergent from the interaction between plant traits and environmental conditions, and that this complexity has historically hindered the development of a generalizable water‐use strategy framework.

This idea is explored using a plant hydraulics model to identify: (a) distinct temporal phases in plant hydraulic regulation during drought that underpin dynamic water‐use responses, and (b) how variation in both traits and environmental forcings can significantly alter common metrics used to characterize plant water‐use strategies. This modelling exercise can bridge the divide between various conceptualizations of water‐use strategies and provide targeted hypotheses to advance the understanding and quantification of plant water status regulation across spatial and temporal scales.

Finally, we describe research frontiers that are necessary to improve the predictive capacity of the plant water‐use strategy concept, including further investigation into the below‐ground determinants of plant water relations, targeted data collection efforts and the potential to scale these concepts from individuals to whole regions.

A freePlain Language Summarycan be found within the Supporting Information of this article.

 

The Craig–Gordon type (C–G) leaf water isotope enrichment models assume a homogeneous distribution of enriched water across the leaf surface, despite observations that Δ¹⁸O can become increasingly enriched from leaf base to tip. Datasets of this ‘progressive isotope enrichment’ are limited, precluding a comprehensive understanding of (a) the magnitude and variability of progressive isotope enrichment, and (b) how progressive enrichment impacts the accuracy of C–G leaf water model predictions. Here, we present observations of progressive enrichment in two conifer species that capture seasonal and diurnal variability in environmental conditions. We further examine which leaf water isotope models best capture the influence of progressive enrichment on bulk needle water Δ¹⁸O. Observed progressive enrichment was large and equal in magnitude across both species. The magnitude of this effect fluctuated seasonally in concert with vapour pressure deficit, but was static in the face of diurnal cycles in meteorological conditions. Despite large progressive enrichment, three variants of the C–G model reasonably successfully predicted bulk needle Δ¹⁸O. Our results thus suggest that the presence of progressive enrichment does not impact the predictive success of C–G models, and instead yields new insight regarding the physiological and anatomical mechanisms that cause progressive isotope enrichment.

 

Multi‐year lags in tree drought recovery, termed ‘drought legacy effects’, are important for understanding the impacts of drought on forest ecosystems, including carbon (C) cycle feedbacks to climate change. Despite the ubiquity of lags in drought recovery, large uncertainties remain regarding the mechanistic basis of legacy effects and their importance for the C cycle. In this review, we identify the approaches used to study legacy effects, from tree rings to whole forests. We then discuss key knowledge gaps pertaining to the causes of legacy effects, and how the various mechanisms that may contribute these lags in drought recovery could have contrasting implications for the C cycle. Furthermore, we conduct a novel data synthesis and find that legacy effects differ drastically in both size and length across the US depending on if they are identified in tree rings versus gross primary productivity. Finally, we highlight promising approaches for future research to improve our capacity to model legacy effects and predict their impact on forest health. We emphasise that a holistic view of legacy effects – from tissues to whole forests – will advance our understanding of legacy effects and stimulate efforts to investigate drought recovery via experimental, observational and modelling approaches.

 

Severe droughts can impart long‐lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole‐forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late‐season droughts (August–September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.

 

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.

 

Globally, intrinsic water-use efficiency (iWUE) has risen dramatically over the past century in concert with increases in atmospheric CO 2 concentration. This increase could be further accelerated by long-term drought events, such as the ongoing multidecadal “megadrought” in the American Southwest. However, direct measurements of iWUE in this region are rare and largely constrained to trees, which may bias estimates of iWUE trends toward more mesic, high elevation areas and neglect the responses of other key plant functional types such as shrubs that are dominant across much of the region. Here, we found evidence that iWUE is increasing in the Southwest at one of the fastest rates documented due to the recent drying trend. These increases were particularly large across three common shrub species, which had a greater iWUE sensitivity to aridity than Pinus ponderosa , a common tree species in the western United States. The sensitivity of both shrub and tree iWUE to variability in atmospheric aridity exceeded their sensitivity to increasing atmospheric [CO 2 ]. The shift to more water-efficient vegetation would be, all else being equal, a net positive for plant health. However, ongoing trends toward lower plant density, diminished growth, and increasing vegetation mortality across the Southwest indicate that this increase in iWUE is unlikely to offset the negative impacts of aridification.