Sperry JS, Venturas MD, Anderegg WRL, Mencuccini M, Mackay DS, et al. (2017). Predicting stomatal responses to the environment from the optimization of photosynthetic gain and hydraulic cost. Plant Cell Environ 40(6): 816-830.
Stomatal regulation presumably evolved to optimize COfor HO exchange in response to changing conditions. If the optimization criterion can be readily measured or calculated, then stomatal responses can be efficiently modelled without recourse to empirical models or underlying mechanism. Previous efforts have been challenged by the lack of a transparent index for the cost of losing water. Yet it is accepted that stomata control water loss to avoid excessive loss of hydraulic conductance from cavitation and soil drying. Proximity to hydraulic failure and desiccation can represent the cost of water loss. If at any given instant, the stomatal aperture adjusts to maximize the instantaneous difference between photosynthetic gain and hydraulic cost, then a model can predict the trajectory of stomatal responses to changes in environment across time. Results of this optimization model are consistent with the widely used Ball-Berry-Leuning empirical model (r > 0.99) across a wide range of vapour pressure deficits and ambient COconcentrations for wet soil. The advantage of the optimization approach is the absence of empirical coefficients, applicability to dry as well as wet soil and prediction of plant hydraulic status along with gas exchange.
Tai X, Mackay DS, Anderegg WR, Sperry JS, Brooks PD (2017). Plant hydraulics improves and topography mediates prediction of aspen mortality in southwestern USA. New Phytol 213(1): 113-127.
Elevated forest mortality has been attributed to climate change-induced droughts, but prediction of spatial mortality patterns remains challenging. We evaluated whether introducing plant hydraulics and topographic convergence-induced soil moisture variation to land surface models (LSM) can help explain spatial patterns of mortality. A scheme predicting plant hydraulic safety loss from soil moisture was developed using field measurements and a plant physiology-hydraulics model, TREES. The scheme was upscaled to Populus tremuloides forests across Colorado, USA, using LSM-modeled and topography-mediated soil moisture, respectively. The spatial patterns of hydraulic safety loss were compared against aerial surveyed mortality. Incorporating hydraulic safety loss raised the explanatory power of mortality by 40% compared to LSM-modeled soil moisture. Topographic convergence was mostly influential in suppressing mortality in low and concave areas, explaining an additional 10% of the variations in mortality for those regions. Plant hydraulics integrated water stress along the soil-plant continuum and was more closely tied to plant physiological response to drought. In addition to the well-recognized topo-climate influence due to elevation and aspect, we found evidence that topographic convergence mediates tree mortality in certain parts of the landscape that are low and convergent, likely through influences on plant-available water.
Sperry JS, Wang Y, Wolfe BT, Mackay DS, Anderegg WR, et al. (2016). Pragmatic hydraulic theory predicts stomatal responses to climatic water deficits. New Phytol 212(3): 577-589.
Ecosystem models have difficulty predicting plant drought responses, partially from uncertainty in the stomatal response to water deficits in soil and atmosphere. We evaluate a 'supply-demand' theory for water-limited stomatal behavior that avoids the typical scaffold of empirical response functions. The premise is that canopy water demand is regulated in proportion to threat to supply posed by xylem cavitation and soil drying. The theory was implemented in a trait-based soil-plant-atmosphere model. The model predicted canopy transpiration (E), canopy diffusive conductance (G), and canopy xylem pressure (P) from soil water potential (P) and vapor pressure deficit (D). Modeled responses to D and Pwere consistent with empirical response functions, but controlling parameters were hydraulic traits rather than coefficients. Maximum hydraulic and diffusive conductances and vulnerability to loss in hydraulic conductance dictated stomatal sensitivity and hence the iso- to anisohydric spectrum of regulation. The model matched wide fluctuations in G and Pacross nine data sets from seasonally dry tropical forest and piñon-juniper woodland with < 26% mean error. Promising initial performance suggests the theory could be useful in improving ecosystem models. Better understanding of the variation in hydraulic properties along the root-stem-leaf continuum will simplify parameterization.
McDowell NG, Fisher RA, Xu C, Domec JC, Hölttä T, et al. (2013). Evaluating theories of drought-induced vegetation mortality using a multimodel-experiment framework. New Phytol 200(2): 304-21.
SUMMARY: Model-data comparisons of plant physiological processes provide an understanding of mechanisms underlying vegetation responses to climate. We simulated the physiology of a piñon pine-juniper woodland (Pinus edulis-Juniperus monosperma) that experienced mortality during a 5 yr precipitation-reduction experiment, allowing a framework with which to examine our knowledge of drought-induced tree mortality. We used six models designed for scales ranging from individual plants to a global level, all containing state-of-the-art representations of the internal hydraulic and carbohydrate dynamics of woody plants. Despite the large range of model structures, tuning, and parameterization employed, all simulations predicted hydraulic failure and carbon starvation processes co-occurring in dying trees of both species, with the time spent with severe hydraulic failure and carbon starvation, rather than absolute thresholds per se, being a better predictor of impending mortality. Model and empirical data suggest that limited carbon and water exchanges at stomatal, phloem, and below-ground interfaces were associated with mortality of both species. The model-data comparison suggests that the introduction of a mechanistic process into physiology-based models provides equal or improved predictive power over traditional process-model or empirical thresholds. Both biophysical and empirical modeling approaches are useful in understanding processes, particularly when the models fail, because they reveal mechanisms that are likely to underlie mortality. We suggest that for some ecosystems, integration of mechanistic pathogen models into current vegetation models, and evaluation against observations, could result in a breakthrough capability to simulate vegetation dynamics.
Adelman JD, Ewers BE, Mackay DS (2008). Use of temporal patterns in vapor pressure deficit to explain spatial autocorrelation dynamics in tree transpiration. Tree Physiol 28(4): 647-58.
To quantify the relationship between temporal and spatial variation in tree transpiration, we measured sap flow in 129 trees with constant-heat sap flow sensors in a subalpine forest in southern Wyoming, USA. The forest stand was located along a soil water gradient from a stream side to near the top of a ridge. The stand was dominated by Pinus contorta Dougl. ex Loud. with Picea engelmannii Parry ex Engelm and Abies lasiocarpa (Hook.) Nutt. present near the stream and scattered individuals of Populus tremuloides Michx. throughout the stand. We used a cyclic sampling design that maximized spatial information with a minimum number of samples for semivariogram analyses. All species exhibited previously established responses to environmental variables in which the dominant driver was a saturating response to vapor pressure deficit (D). This response to D is predictable from tree hydraulic theory in which stomatal conductance declines as D increases to prevent excessive cavitation. The degree to which stomatal conductance declines with D is dependent on both species and individual tree physiology and increases the variability in transpiration as D increases. We quantified this variability spatially by calculating the spatial autocorrelation within 0.2-kPa D bins. Across 11 bins of D, spatial autocorrelation in individual tree transpiration was inversely correlated to D and dropped from 45 to 20 m. Spatial autocorrelation was much less for transpiration per unit leaf area and not significant for transpiration per unit sapwood area suggesting that spatial autocorrelation within a particular D bin could be explained by tree size. Future research should focus on the mechanisms underlying tree size spatial variability, and the potentially broad applicability of the inverse relationship between D and spatial autocorrelation in tree transpiration.
Ewers BE, Mackay DS, Samanta S (2007). Interannual consistency in canopy stomatal conductance control of leaf water potential across seven tree species. Tree Physiol 27(1): 11-24.
We investigated interannual variability of canopy transpiration per unit ground area (E (C)) and per unit leaf area (E (L)) across seven tree species in northern Wisconsin over two years. These species have previously been shown to be sufficient to upscale stand-level transpiration to the landscape level during one growing season. Our objective was to test whether a simple plant hydraulic model could capture interannual variation in transpiration. Three species, wetland balsam fir (Abies balsamea (L.) Mill), basswood (Tilia Americana L.) and speckled alder (Alnus rugosa (DuRoi) Spreng), had no change in E (C) or E (L) between 2000 and 2001. Red pine (Pinus resinosa Ait) had a 57 and 19% increase in E (C) and E (L), respectively, and sugar maple (Acer saccharum Marsh) had an 83 and 41% increase in E (C) and E (L), respectively, from 2000 to 2001. Quaking aspen (Populus tremuloides Michx) had a 50 and 21% decrease in E (C) and E (L), respectively, from 2000 to 2001 in response to complete defoliation by forest tent caterpillar (Malascoma distria Hüber) and subsequent lower total leaf area index of the reflushed foliage. White cedar (Thuja occidentalis L.) had a 20% decrease in both E (C) and E (L) caused by lowered surface water in wetlands in 2001 because of lower precipitation and wetland flow management. Upland A. balsamea increased E (L) and E (C) by 55 and 53%, respectively, as a result of release from light competition of the defoliated, overstory P. tremuloides. We hypothesized that regardless of different drivers of interannual variability in E (C) and E (L), minimum leaf water potential would be regulated at the same value. Minimum midday water potentials were consistent over the two years within each of the seven species despite large changes in transpiration between years. This regulation was independently verified by the exponential saturation between daily E (C) and vapor pressure deficit (D) and the tradeoff between a reference canopy stomatal conductance (G (S)) and the sensitivity of G (S) to D, indicating that trees with high G (S) must decrease G (S) in response to atmospheric drought faster than trees with low G (S). Our results show that models of forest canopy transpiration can be simplified by incorporating G (S) regulation of minimum leaf water potential for isohydric species.
Predicting stomatal responses to the environment from the optimization of photosynthetic gain and hydraulic cost.
Plant hydraulics improves and topography mediates prediction of aspen mortality in southwestern USA.
Evaluating theories of drought-induced vegetation mortality using a multimodel-experiment framework.
Use of temporal patterns in vapor pressure deficit to explain spatial autocorrelation dynamics in tree transpiration.
Interannual consistency in canopy stomatal conductance control of leaf water potential across seven tree species.