According to classic stomatal optimization theory, plant stomata are regulated to maximize carbon assimilation for a given water loss. A key component of stomatal optimization models is marginal water‐use efficiency (mWUE), the ratio of the change of transpiration to the change in carbon assimilation. Although the mWUE is often assumed to be constant, variability of mWUE under changing hydrologic conditions has been reported. However, there has yet to be a consensus on the patterns of mWUE variabilities and their relations with atmospheric aridity. We investigate the dynamics of mWUE in response to vapor pressure deficit (VPD) and aridity index using carbon and water fluxes from 115 eddy covariance towers available from the global database FLUXNET. We demonstrate a non‐linear mWUE‐VPD relationship at a sub‐daily scale in general; mWUE varies substantially at both low and high VPD levels. However, mWUE remains relatively constant within the mid‐range of VPD. Despite the highly non‐linear relationship between mWUE and VPD, the relationship can be informed by the strong linear relationship between ecosystem‐level inherent water‐use efficiency (IWUE) and mWUE using the slope,
Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon‐maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle.
more » « less- Award ID(s):
- 1714972
- PAR ID:
- 10058812
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Ecology Letters
- Volume:
- 21
- Issue:
- 7
- ISSN:
- 1461-023X
- Page Range / eLocation ID:
- p. 968-977
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract m *. We further identify site‐specificm * and its variability with changing site‐level aridity across six vegetation types. We suggest accurately representing the relationship between IWUE and VPD using Michaelis–Menten or quadratic functions to ensure precise estimation of mWUE variability for individual sites. -
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Summary The classical theory of stomatal optimization stipulates that stomata should act to maximize photosynthesis while minimizing transpiration. This theory, despite its remarkable success in reproducing empirical patterns, does not account for the fact that the available water to plants is dynamically regulated by plants themselves, and that plants compete for water in most locations.
Here, we develop an alternative theory in which plants maximize the expected carbon gain under stochastic rainfall in a competitive environment. We further incorporate xylem hydraulic limitation as an additional constraint to transpiration and evaluate its impacts on stomatal optimization by incorporating the direct carbon cost of xylem recovery and the opportunity cost of reduced future photosynthesis as a result of irrecoverable xylem damage.
We predict stomatal behaviour to be more conservative with a higher cost induced by xylem damage. By varying the unit carbon cost and extent of xylem recovery, characterizing the direct and opportunity cost of xylem damage, respectively, our model can reproduce several key patterns of stomatal‐hydraulic trait covariations.
By addressing the key elements of water limitation in plant gas exchange simultaneously, including plants’ self‐regulation of water availability, competition for water and hydraulic risk, our study provides a comprehensive theoretical basis for understanding stomatal behaviour.
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null (Ed.)Abstract. Evaporation (E) and transpiration (T) respond differentlyto ongoing changes in climate, atmospheric composition, and land use. It isdifficult to partition ecosystem-scale evapotranspiration (ET) measurementsinto E and T, which makes it difficult to validate satellite data and landsurface models. Here, we review current progress in partitioning E and T andprovide a prospectus for how to improve theory and observations goingforward. Recent advancements in analytical techniques create newopportunities for partitioning E and T at the ecosystem scale, but theirassumptions have yet to be fully tested. For example, many approaches topartition E and T rely on the notion that plant canopy conductance andecosystem water use efficiency exhibit optimal responses to atmosphericvapor pressure deficit (D). We use observations from 240 eddy covariance fluxtowers to demonstrate that optimal ecosystem response to D is a reasonableassumption, in agreement with recent studies, but more analysis is necessaryto determine the conditions for which this assumption holds. Anothercritical assumption for many partitioning approaches is that ET can beapproximated as T during ideal transpiring conditions, which has beenchallenged by observational studies. We demonstrate that T can exceed 95 %of ET from certain ecosystems, but other ecosystems do not appear to reachthis value, which suggests that this assumption is ecosystem-dependent withimplications for partitioning. It is important to further improve approachesfor partitioning E and T, yet few multi-method comparisons have beenundertaken to date. Advances in our understanding of carbon–water couplingat the stomatal, leaf, and canopy level open new perspectives on how toquantify T via its strong coupling with photosynthesis. Photosynthesis can beconstrained at the ecosystem and global scales with emerging data sourcesincluding solar-induced fluorescence, carbonyl sulfide flux measurements,thermography, and more. Such comparisons would improve our mechanisticunderstanding of ecosystem water fluxes and provide the observationsnecessary to validate remote sensing algorithms and land surface models tounderstand the changing global water cycle.more » « less
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