the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Non-steady-state Stomatal Conductance Modeling and Its Implications: From Leaf to Ecosystem
Yujie Wang
Troy Magney
Christian Frankenberg
Abstract. Accurate and efficient modeling of stomatal conductance (gs) has been a key challenge in vegetation models across scales. Current practice of most land surface models (LSMs) assumes steady-state gs and predicts stomatal responses to environmental cues as immediate jumps between stationary regimes. However, the response of stomata can be orders of magnitude slower than that of photosynthesis, and often cannot reach a steady state before the next model time-step, even on half-hourly time scales. Here, we implemented a simple dynamic gs model in the vegetation module of an LSM developed within the Climate Modeling Alliance, and investigated the potential biases caused by the steady-state assumption from leaf to canopy scales. In comparison with steady-state models, the dynamic model better predicted the coupled temporal response of photosynthesis and stomatal conductance to changes in light intensity using leaf measurements. In ecosystem flux simulations, while the impact of gs hysteresis response may not be substantial in terms of daily or monthly integrated canopy fluxes, our results suggested the importance of considering this effect when quantifying fluxes in the mornings and evenings, and interpreting diurnal hysteresis patterns observed in ecosystem fluxes. Furthermore, prognostic modeling can bypass the A-Ci iterations required for steady-state simulations and can be robustly run with comparable computational costs. Overall, our study highlights the implications of dynamic gs modeling in improving the accuracy and efficiency of LSMs, and for advancing our understanding of plant-environment interactions.
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Ke Liu et al.
Status: final response (author comments only)
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RC1: 'Comment on egusphere-2023-1706', Anonymous Referee #1, 11 Sep 2023
This manuscript describes a new stomatal model that does not assume steady state responses to climate. Instead, this approach models gs with prognostic updates at each time step, which allows the model simulate limitations to gas exchange caused by stomatal speed and simplify leaf energy balance calculations.
The model was coupled with an LSM and evaluated at leaf and ecosystem levels. At leaf level the model could predict better the stomatal responses to changes in light regime. At ecosystem level, the model makes only a small difference at the canopy fluxes at mornings and evenings.
The paper is well written, and the results are generally clearly presented. I find the results of the paper interesting, and I really like the idea of the dynamic stomatal model because it seems a better representation of plant stomatal behaviour in a (rapidly) changing environment. I believe this approach have the potential to improve how LSMs represent ecosystem carbon and water dynamics. My only criticisms are relatively minor methodological issues explained in detail below. Additionally, I think the paper would be improved if the authors had also validated their dynamic model against ecosystem level observations (i.e. eddy flux data) similarly to what they did for leaf-level data.
Specific comments:
L9: In the results you claim the daily effect of the dynamic model is notable (L244)
L24: The idea of using optimization to predict stomatal behaviour goes back to Cowan & Farquhar 1977, so I am not sure if using the term “more recently” is really appropriated.
L25: Most of the models discussed in these papers do not really optimize water use efficiency; they optimize the balance between carbon gain with some penalty, which can be related to plant hydraulics, non-stomatal limitations to photosynthesis, etc.
L34-35: I assume it depends when the “next change occurs”? It would be useful to provide some quantitative examples of the time scales of stomatal response to environmental changes to clarify that.
L100: What steady state model do you use to calculate the target gs?
L120: How many leaves/plants were used?
L140: This vcmax value should be micromol instead?
L170: Wouldn’t a linear interpolation homogenize the environmental conditions over time and “mask” the real importance of the non-steady state model? I assume the dynamic model would only make a bigger difference in environments with rapidly changing environmental conditions. Would that explain why you have a relatively small impact of the dynamic model on the leaf and canopy simulations?
L175: This section needs more details on the LSM configuration for these simulations to allow reproducibility. For example, how canopy light diffusion, carbon allocation/vegetation dynamics and soil moisture were handled. Maybe add this information as supplementary material.
L205: check vcmax units
L245: Do the daily differences “disappear” at monthly scales because you have opposite RD in different days that cancel each other out? If that is the case, it would still be interesting to show the total monthly differences between SS and NSS (using for example the mean absolute error between SS and NSS). I believe the differences between models throughout the month could still result in different trajectories for the vegetation over time, for example, if the higher carbon gain of a model in a given day resulted in more leaves being produced on that day, this model productivity advantage that would accumulate over the other one.
L276: I could not find the computational cost differences in the results.
L295: It could be interesting to see the cumulative effect of this unnecessary water loss on xylem hydraulic damage in future studies.
Fig. 2: I don’t consider this figure really essential for the manuscript. Considering you already have 9 figures maybe it would be best to leave it as supplementary material.
Fig. 6: Its hard to visualize the RD, please use darker shading.
Citation: https://doi.org/10.5194/egusphere-2023-1706-RC1 -
RC2: 'Comment on egusphere-2023-1706', Anonymous Referee #2, 15 Sep 2023
The manuscript by Liu et al. presents how including dynamic stomatal responses impacts gas exchange predictions from leaf to canopy. The topic is of high importance as research carried out at the leaf level has shown the importance of stomatal dynamics but there is still a debate about how these dynamics influence gas exchange at a larger scale. The manuscript is well written and I like that the authors are addressing this gap in the literature, however, I have some concerns about the methods used.
What should be clearer is what are the challenges to simulating gas exchange at different scales from leaf to canopy and from seconds to months. For example, during a diurnal period, the slow temporal response of stomatal conductance will lead to the transient limitation of photosynthesis and potential damage to the PSII reaction centres (due to excessive energy received compared to the sink strength). These effects accumulate during the day and impact plant growth, which will in turn affect the gas exchange of the following days. These effects are not clearly accounted for by the model. Averaging the fluxes over a long period is useful for quantifying the ecosystem exchange but does not reflect that every day the environmental conditions will impact the leaf functioning and acclimation. The effect of stomatal dynamics is also dependent on the fluctuation of the environment. The more it fluctuates, the more the stomatal behaviour will matter in the gas exchange.
The methods section is missing important information and requires more clarity. From lines 90 to 105, it is not clear how gbc was set or calculated. How the simplified model was solved and with which procedure is missing. Did the authors use an ODE solver? How are the environmental variables included in this model? Did the authors use them as forcing variables that change continuously? How is the steady state gs calculated? Which model was used: Ball or Medlyn? How were they calibrated? At different places in the manuscript, the time steps used are different and without a clear explanation of why, it is confusing. The authors did not include leaf energy balance to calculate leaf temperature, but this can have an important impact on their simulations. The vapour pressure gradient between the leaf and atmosphere depends on this and drives the water exchange. This should be acknowledged at least as a limitation in the discussion. Overall, it is difficult to judge the validity of the simulations, because important details are missing.
For the exponential model used here, there is a time constant for stomatal opening and another one for stomatal closure. Stomata have very different rapidity of response for opening and closing. Here the authors only used one time constant and only estimated it for the opening part. The asymmetry of the time constants is strongly influencing the hysteresis between the morning and afternoon part of the simulation. This should be corrected.
The authors concluded that their optimisation procedure improves the parameter estimation compared to traditional linear fitting methods. It is not clearly explained in the text how these methods differ and what are the improvements. The need to reach steady state gs to estimate the parameter values is known. Using a coupled dynamic gs model and steady-state target model to fit the data of a light response curve has been done previously. In general, it is an interesting result but not the focus of this study.
Line 25: the optimisation theory does not optimize water use efficiency.
Citation: https://doi.org/10.5194/egusphere-2023-1706-RC2
Ke Liu et al.
Ke Liu et al.
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