the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Implementation of an eco-evolutionary approach to photosynthesis and acclimation in the JULES land surface model
Abstract. Realistic simulation of photosynthesis is needed for accurate prediction of the global carbon cycle. In addition to the well-known instantaneous response to changes in temperature, photosynthetic processes acclimate to long‐term environmental changes by adjusting the maximum photosynthetic capacities (Vcmax and Jmax) and stomatal behaviour The theoretical basis for acclimation can be understood as an outcome of eco-evolutionary optimality (EEO). Here, we have implemented an EEO‐based scheme to represent the universal acclimation of photosynthesis to environmental conditions independently of plant functional types in the JULES land surface model. We compare this implementation with two standard configurations of JULES (GL7.0, UKESM1). We evaluate model performance using daily, monthly and annual site-based observations of gross primary production (GPP) observations at flux towers from the PLUMBER2 data set. The EEO-based scheme, PJULES, produces better predictions of GPP than either GL7 or UKESM1, both of which substantially underestimate GPP at the higher end of the observed range. Comparison at individual sites shows that PJULES produces more realistic simulations of seasonal and diurnal cycles of GPP. The improvement in the estimation of GPP does not degrade simulated land-surface energy fluxes: predictions of the latent heat flux are similar to those from the UKESM1 and slightly better than those from GL7.0. The good performance of PJULES shows that a parsimonious model can offer a competitive alternative to more complex parameterisations for simulating land–atmosphere carbon and water exchanges.
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Status: open (until 06 Oct 2026)
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RC1: 'Comment on egusphere-2026-2804', Anonymous Referee #1, 11 Sep 2026
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The comment was uploaded in the form of a supplement: https://egusphere.copernicus.org/preprints/2026/egusphere-2026-2804/egusphere-2026-2804-RC1-supplement.pdfReplyCitation: https://doi.org/
10.5194/egusphere-2026-2804-RC1 -
RC2: 'Comment on egusphere-2026-2804', Anonymous Referee #2, 15 Sep 2026
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Summary of work
This study implements in the JULES land surface model a sub-daily, eco-evolutionary optimality-based model of C3 plant photosynthesis (the P-model) based on the well-used Farquhar et al description of C3 photosynthesis. The big-leaf P-model replaces the multi-layer Collatz photosynthesis and Jacobs photosynthesis-conductance closure schemes from JULES to yield a model called PJULES. The authors evaluate PJULES against two standard JULES configurations (GL7 and UKESM1) using model forcing and GPP and LE observations from 134 flux sites in the PLUMBER2 database. Results show that PJULES outperforms standard JULES without compromising modelled LE flux across a range of sites and conditions. This paper is a useful contribution to land-surface modelling in general and to the rapidly developing sub-field of EEO methods. The manuscript is well-written and well-presented throughout, uses appropriate methods and reaches sensible conclusions.
Main comments- It's disappointing to see only one week of sub-daily analysis. It seems likely that the results from this will be weather dependent, especially as the authors chose to use the same week at all sites, which wouldn't necessarily be a representative period for any particular site. Comparison of Figs 4 and 5 by eye seems to confirm this. What was the reasoning behind these choices?
- To what extent are the differences between PJULES and JULES down to the switch from Collatz to Farquhar, rather than the P-model hypotheses themselves? I wouldn't expect results from this in the main paper, but it would be useful if they could inform the discussion.
- What is the impact on NPP? As I understand it, JULES leaf- and canopy-level respiration is derived from Vcmax, so do the PJULES improvements in GPP transfer through to NPP or are they cancelled by similar changes in respiration?
- If the big-leaf P-model improves GPP without degrading LE, then what are the remaining benefits of the multi-layer configurations? Are the authors suggesting that multi-layer features like sunfleck penetration aren't needed? Some discussion on these implications would be appreciated.
Minor comments- Eqns 14, 15, 16: I assume that the authors have switched to using "=" as an assignment symbol rather than an equality symbol, as used in the rest of the paper, which will be confusing for readers. The authors should check these equations, as they appear not quite right for an exponential smoother.
- Eqns S26, S27: The supplementary info refers to "acclimated temperature (Taccl)" but I'm having trouble understanding what this is or relating it to the equations in the main text. In fact, I find it unhelpful that the SI frequently uses different symbols than the main text for the same variables, e.g., activation energies "\Delta H_a" versus "E_{a,c}". The authors should make their chosen notation internally consistent.
- L271: "acclimated values are updated at 12:30 LT": Is it a good idea to update these values at a time of day when GPP is high? Does this cause abrupt, unphysical changes in the model simulations around noon?
- L417: "green-up too early": I normally associate "greening-up" as referring to increasing LAI, but that is prescribed in these simulations. Is there a better phrasing that could be used here?
- Table 3: I'd like too see some uncertainties (standard errors) and N-values on these means, particularly for the observations. This would provide useful context for interpretation of the various model means.
- Fig 8: The regression statistics are small and hard to read - could the authors increase the size, as per other figures?
- L654: "promising way of improving the soil moisture stress function". There are already several options in JULES for modifying this function that are not used by GL7 or UKESM1. I think it would be worth acknowledging that these should be evaluated alongside anything new from Stocker et al.
Citation: https://doi.org/10.5194/egusphere-2026-2804-RC2 -
RC3: 'Comment on egusphere-2026-2804', Anonymous Referee #3, 21 Sep 2026
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This manuscript presents an implementation of the sub-daily P-model for JULES (PJULES), replacing the conventional PFT-specific Collatz photosynthesis scheme and the Jacobs stomatal scheme with an eco-evolutionary optimality-based formulation. PJLUES is evaluated against PLUMBER2 flux observations and compared with the standard GL7 and UKESM1 configurations for JULES. The manuscript offers a promising solution to the challenges of using prescribed PFT-specific parameters for studying long-term changes in vegetation dynamics and the carbon cycle under climate change, an important problem in the land-surface modelling community. The results look promising, with substantial improvement in PJULES for estimating GPP across different sites around the globe. The use of a large flux-tower dataset and evaluation over multiple temporal scales are valuable.
However, I have several concerns regarding the attribution of model improvement, the description of the acclimation scheme, and the interpretation of the GPP and LE results. In its current form, the analyses demonstrating model improvement are still not clear, with limited information on the structural aspect of the model for these improvements. In addition, some equations describing the central acclimation algorithm appear inconsistent or erroneous. I therefore recommend substantial revision before the manuscript can be considered for publication in the journal.
Major comments
- Attribution of the PJULES improvement in more detail. It would be important to understand why PJULES performs better. Several major changes have been implemented in PJULES compared with the GL7/UKESM1, including introducing eco-evolutionary optimality and an acclimation scheme, and changing the biochemical photosynthesis model, stomatal formulation, canopy representation and PFTs structure. The current comparison only demonstrates the overall improved performance of the complete PJULES configuration, but it is not clear which structural change mainly contributes to such improvement. If technically feasible, I strongly encourage additional ablation experiments to distinguish the effects of, for example, the fast and slow processes (e.g. response to instantaneous climate versus acclimation). Otherwise, it may limit the application of the model under varied environmental conditions when different temporal scales of climate signals come into play simultaneously.
- Analysis of GPP variability for understanding model improvement at varied temporal scales. This is in line with my previous comments. What aspect of GPP variability is improved? The current evaluation does not clearly distinguish improvement in the seasonal/diurnal cycle from improvements in temporal variability triggered by instant and/or acute climate signals around those cycles. Note that monthly and sub-daily R2 values are strongly influenced by the seasonal and diurnal signals themselves and may confound the evaluation. I suggest decomposing the time series signals into at least two components: one is the seasonal/diurnal cycles, and the other is daily/hourly residuals for evaluation. This could help understand how PJULES capture weather-driven and periodic signals in simulating GPP.
- Interpretation of the LE results and carbon-water coupling. The substantially greater improvement in GPP than in total LE deserves a more process-based analysis. Since the stomatal scheme that governs water-carbon coupling has changed, has the simulated transpiration also been improved? The current analysis of total LE is not really revealing; as noticed by the authors, it could be compensated by other hydrological components. If the selected site has transpiration measurements, I strongly suggest showing evaluations for transpiration as well. Otherwise, the gridded datasets with transpiration estimates could also be a good option. This could directly reflect the performance of the new stomatal scheme and could provide much stronger support for the overall improvement of carbon-water coupling in a land surface model when the P-model is implemented.
- Data selection for evaluation. The criteria for selecting the seven sites used for detailed seasonal and diurnal analysis are unclear; this is important as the statements of model improvement are mainly based on these analyses; however, the representativeness of employing such a small sample among more than 130 sites is questionable. For example, do those sites cover a large enough climate gradient and sufficient vegetation types?
Minor comments:
- Equations. 14-16: These acclimation equations appear inconsistent with the stated 15-day memory. For example, for Eqn. 14, this algebraically reduces to the previous day’s optimal value and therefore removes the intended temporal memory. Please double-check if there are any errors in the variable labelling.
- Please describe the actual representation of T_accl in Eqns. S26-S27.
- Please double-check the unit in Eqn. S17 and confirm whether the intrinsic quantum efficiency can be negative. Based on the current unit setting, the intrinsic quantum efficiency is likely negative.
- What does the “minn” function in Eq. 21 refer to?
- Lines 297-298: The statement that hourly forcing was “resampled to half-hourly resolution” requires clarification. Please describe the temporal disaggregation/interpolation procedure for each relevant forcing variable.
- Model initialisation and spin-up procedures should also be documented.
- Table 3 is numbered twice.
- Line 508: Fig. 8 -> Fig. 6?
Citation: https://doi.org/10.5194/egusphere-2026-2804-RC3
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