On the importance of stoichiometric constraints for understanding global change effects on food web dynamics
Stoichiometric constraints modulate the effects of temperature and nutrients on biomass distribution and community stability
Recommendation: posted 22 January 2020, validated 29 January 2020
The constraints associated with the mass balance of chemical elements (i.e. stoichiometric constraints) are critical to our understanding of ecological interactions, as outlined by the ecological stoichiometry theory . Species in ecosystems differ in their elemental composition as well as in their level of elemental homeostasis , which can determine the outcome of interactions such as herbivory or decomposition on species coexistence and ecosystem functioning [3, 4].
Despite their importance, stoichiometric constraints are still often ignored in theoretical studies exploring the consequences of environmental perturbations on food web stability. Meanwhile, drivers of global change strongly alter biochemical cycles and the balance of chemical elements in ecosystems . An important challenge is thus to understand how stoichiometric constraints affect food web responses to global changes.
The study of Sentis et al.  makes a step in that direction. This article investigates how stoichiometric constraints affect the response of consumer-resource dynamics to increasing temperature and nutrient inputs. It shows that the stoichiometric flexibility of the resource, coupled with lower consumer assimilation efficiency when stoichiometric unbalance between the resource and the consumer is higher, dampens the destabilizing effects of nutrient enrichment on species dynamics but reduces consumer persistence at extreme temperatures. Interestingly, these effects of stoichiometric constraints arise not only from changes in species assimilation efficiencies and carrying capacities but also from stoichiometric negative feedback loops on resource and consumer populations.
The results of this study are a call to further include stoichiometric constraints into food web models to better understand and predict the consequences of global changes on ecological communities. Many perspectives exist on that issue. For instance, it would be interesting to assess the effects of other stoichiometric mechanisms (e.g. changes in the element limiting growth ) on food web stability and its response to nutrient enrichment, as well as the effects of other global change drivers associated with altered biochemical cycles (e.g. carbon dioxide increase).
 Sterner, R. W. and Elser, J. J. (2017). Ecological Stoichiometry, The Biology of Elements from Molecules to the Biosphere. doi: 10.1515/9781400885695
 Elser, J. J., Sterner, R. W., Gorokhova, E., Fagan, W. F., Markow, T. A., Cotner, J. B., Harrison, J.F., Hobbie, S.E., Odell, G.M., Weider, L. W. (2000). Biological stoichiometry from genes to ecosystems. Ecology Letters, 3(6), 540–550. doi: 10.1111/j.1461-0248.2000.00185.x
 Daufresne, T., and Loreau, M. (2001). Plant–herbivore interactions and ecological stoichiometry: when do herbivores determine plant nutrient limitation? Ecology Letters, 4(3), 196–206. doi: 10.1046/j.1461-0248.2001.00210.x
 Zou, K., Thébault, E., Lacroix, G., and Barot, S. (2016). Interactions between the green and brown food web determine ecosystem functioning. Functional Ecology, 30(8), 1454–1465. doi: 10.1111/1365-2435.12626
 Peñuelas, J., Poulter, B., Sardans, J., Ciais, P., van der Velde, M., Bopp, L., Boucher, O., Godderis, Y., Hinsinger, P., Llusia, J., Nardin, E., Vicca, S., Obersteiner, M., Janssens, I. A. (2013). Human-induced nitrogen–phosphorus imbalances alter natural and managed ecosystems across the globe. Nature Communications, 4(1), 1–10. doi: 10.1038/ncomms3934
 Sentis, A., Haegeman, B. & Montoya, J.M. (2020). Stoichiometric constraints modulate the effects of temperature and nutrients on biomass distribution and community stability. bioRxiv, 589895, ver. 7 peer-reviewed and recommended by PCI Ecology. doi: 10.1101/589895
Elisa Thebault (2020) On the importance of stoichiometric constraints for understanding global change effects on food web dynamics. Peer Community in Ecology, 100039. 10.24072/pci.ecology.100039
The recommender in charge of the evaluation of the article and the reviewers declared that they have no conflict of interest (as defined in the code of conduct of PCI) with the authors or with the content of the article.
Evaluation round #2
DOI or URL of the preprint: https://doi.org/10.1101/589895
Version of the preprint: 2
Author's Reply, 10 Jan 2020
Decision by Elisa Thebault, posted 02 Jan 2020
Many thanks for your thorough revision of the manuscript. There are only a few remaining minor comments to address from one of the reviewers before recommendation.
best wishes, Elisa
Reviewed by anonymous reviewer, 18 Dec 2019
Evaluation round #1
DOI or URL of the preprint: https://doi.org/10.1101/589895
Author's Reply, 25 Nov 2019
Decision by Elisa Thebault, posted 15 Oct 2019
Dear authors, I have now received two reviews of your manuscript. Both reviewers and I are in agreement that this is an interesting study considering how flexible stoichiometry of primary producers affect the dynamics of primary producers and consumers in response to both nutrient enrichment and climate warming. However, several issues have been identified which, in my views, require revision before recommendation. Such revised contribution would need to address all of the reviewer comments. In particular, as outlined by reviewer #2, several points would deserve to be further clarified and discussed (e.g. better justification of the use of different extinction thresholds, robustness of the results to different sets of parameters, consequences of abiotic pools such as detritus).
In addition to the detailed suggestions of the reviewers, I have a few additional comments: - Lines 352-358: The definition of the static and dynamic effects are at first not fully clear. Is the part on consumer energetic efficiency (lines 358-378) related to the static effect while the following part (lines 382-393) relates to the dynamic effect? I better understood the difference between the static and dynamic effects later on (line 395), when these two effects are directly compared. I would suggest modifying this section (from lines 347 to 414) to make the distinction and the comparison between the static and dynamic effects clearer to the reader.
Lines 387-389 “When consumer population increases, this decreases resource population growth leading to a negative feedback on consumer population growth rate”: isn’t it also the case in the RM model? Please explain more precisely why this feedback is different in the SRM model.
Discussion on the effects of temperature on assimilation efficiency (lines 481-493): It could be interesting to discuss a little further which mechanisms lead to reduced assimilation efficiency of the consumer (in relation to reduced resource nutrient quota) at high temperature in the model. This result could potentially also be related to existing experimental and empirical literature on the effects of temperature on the stoichiometry of primary producers (although the mechanisms involved might be indeed different in the model).
I am looking forward to seeing your revised manuscript addressing the reviewers’ comments, along with a point-by-point response.
Best wishes, Elisa Thébault