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The response of forests to climate change depends in part on whether the photosynthetic benefit from increased atmospheric CO 2 (∆C a = future minus historic CO 2 ) compensates for increased physiological stresses from higher temperature (∆T). We predicted the outcome of these competing responses by using optimization theory and a mechanistic model of tree water transport and photosynthesis. We simulated current and future productivity, stress, and mortality in mature monospecific stands with soil, species, and climate sampled from 20 continental US locations. We modeled stands with and without acclimation to ∆C a and ∆T, where acclimated forests adjusted leaf area, photosynthetic capacity, and stand density to maximize productivity while avoiding stress. Without acclimation, the ∆C a -driven boost in net primary productivity (NPP) was compromised by ∆T-driven stress and mortality associated with vascular failure. With acclimation, the ∆C a -driven boost in NPP and stand biomass (C storage) was accentuated for cooler futures but negated for warmer futures by a ∆T-driven reduction in NPP and biomass. Thus, hotter futures reduced forest biomass through either mortality or acclimation. Forest outcomes depended on whether projected climatic ∆C a /∆T ratios were above or below physiological thresholds that neutralized the negative impacts of warming. Critically, if forests do not acclimate, the ∆C a /∆T must be above ca . 89 ppm⋅°C −1 to avoid chronic stress, a threshold met by 55% of climate projections. If forests do acclimate, the ∆C a /∆T must rise above ca . 67 ppm⋅°C −1 for NPP and biomass to increase, a lower threshold met by 71% of projections. 

Optimal stomatal control models have shown great potential in predicting stomatal behavior and improving carbon cycle modeling. Basic stomatal optimality theory posits that stomatal regulation maximizes the carbon gain relative to a penalty of stomatal opening. All models take a similar approach to calculate instantaneous carbon gain from stomatal opening (the gain function). Where the models diverge is in how they calculate the corresponding penalty (the penalty function). In this review, we compare and evaluate 10 different optimization models in how they quantify the penalty and how well they predict stomatal responses to the environment. We evaluate models in two ways. First, we compare their penalty functions against seven criteria that ensure a unique and qualitatively realistic solution. Second, we quantitatively test model against multiple leaf gas‐exchange datasets. The optimization models with better predictive skills have penalty functions that meet our seven criteria and use fitting parameters that are both few in number and physiology based. The most skilled models are those with a penalty function based on stress‐induced hydraulic failure. We conclude by proposing a new model that has a hydraulics‐based penalty function that meets all seven criteria and demonstrates a highly predictive skill against our test datasets.

 

Herein we review the current state‐of‐the‐art of plant hydraulics in the context of plant physiology, ecology, and evolution, focusing on current and future research opportunities. We explain the physics of water transport in plants and the limits of this transport system, highlighting the relationships between xylem structure and function. We describe the great variety of techniques existing for evaluating xylem resistance to cavitation. We address several methodological issues and their connection with current debates on conduit refilling and exponentially shaped vulnerability curves. We analyze the trade‐offs existing between water transport safety and efficiency. We also stress how little information is available on molecular biology of cavitation and the potential role of aquaporins in conduit refilling. Finally, we draw attention to how plant hydraulic traits can be used for modeling stomatal responses to environmental variables and climate change, including drought mortality.

 

Plant functional traits provide a link in process‐based vegetation models between plant‐level physiology and ecosystem‐level responses. Recent advances in physiological understanding and computational efficiency have allowed for the incorporation of plant hydraulic processes in large‐scale vegetation models. However, a more mechanistic representation of water limitation that determines ecosystem responses to plant water stress necessitates a re‐evaluation of trait‐based constraints for plant carbon allocation, particularly allocation to leaf area. In this review, we examine model representations of plant allocation to leaves, which is often empirically set by plant functional type‐specific allometric relationships. We analyze the evolution of the representation of leaf allocation in models of different scales and complexities. We show the impacts of leaf allocation strategy on plant carbon uptake in the context of recent advancements in modeling hydraulic processes. Finally, we posit that deriving allometry from first principles using mechanistic hydraulic processes is possible and should become standard practice, rather than using prescribed allometries. The representation of allocation as an emergent property of scarce resource constraints is likely to be critical to representing how global change processes impact future ecosystem dynamics and carbon fluxes and may reduce the number of poorly constrained parameters in vegetation models.

 

Trees may survive prolonged droughts by shifting water uptake to reliable water sources, but it is unknown if the dominant mechanism involves activating existing roots or growing new roots during drought, or some combination of the two.

To gain mechanistic insights on this unknown, a dynamic root‐hydraulic modeling framework was developed that set up a feedback between hydraulic controls over carbon allocation and the role of root growth on soil–plant hydraulics. The new model was tested using a 5 yr drought/heat field experiment on an established piñon‐juniper stand with root access to bedrock groundwater.

Owing to the high carbon cost per unit root area, modeled trees initialized without adequate bedrock groundwater access experienced potentially lethal declines in water potential, while all of the experimental trees maintained nonlethal water potentials. Simulated trees were unable to grow roots rapidly enough to mediate the hydraulic stress, particularly during warm droughts. Alternatively, modeled trees initiated with root access to bedrock groundwater matched the hydraulics of the experimental trees by increasing their water uptake from bedrock groundwater when soil layers dried out.

Therefore, the modeling framework identified a critical mechanism for drought response that required trees to shift water uptake among existing roots rather than growing new roots.

 

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.