Proposal Status | MAINpeople.oregonstate.edu/~novakm/wp-content/uploads/2014/12/Nova… · The payoff is twofold, first the implementation and improvement of a new way of measuring

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Panel Summary #1

Proposal Number: 1353827

Panel Summary: Panel Summary

Population and Community Ecology Fall 2013 Advisory Panel

Results of Prior NSF Support (If Applicable)

Intellectual Merit: Prior support was DDIG to the PI. It supported the research that acts as the starting point of thecurrent proposal and was a very strong contributions that led to 5 publications Broader Impacts: This supported 4 undergraduates and is a strong contribution for a DDIG.

If this is a resubmission, how have previous criticisms been addressed?

NA

Criterion I: Intellectual Merit

Intellectual Strengths: Combining the methods for inferring functional responses previously developed by Novak andWootton with observational and experimental methods to address the stability of generalist consumers (incomparison with specialists) is an exciting area both in community ecology and even more so in basic populationbiology of consumers. The question is well developed by the PI and asks why generalist consumers show such lowlevels of population cycles when there are various ways in which such cycles should be more often seen. Theproposed solution, that multispecies functional responses are generally more stabilizing than previously thought andcan be quantified using extensions of the method of Novak and Wootton, is also well justified. If successful, thisresearch would shed novel insights into population cycles in food webs and have a broad array of implications forpopulation and community ecology.

Intellectual Weaknesses: Although some reviewers were confused about the relative importance of the goals of theproposal (refining the method vs applying it to ask about population cycles), few weaknesses were identified. Theseconsisted of asking if there were ways to better incorporate other factors into the theory and/or approach and thesuggestion that more effort might be invested into the proposed experiments to better isolate the process andpossibly examine the role of such other factors. The panel struggled with identifying the main goals of the proposal,particularly in the project summary which didn't adequately reflect important elements of the proposal.

Criterion II: Broader Impacts Broader Impact Strengths: Integrating research with teaching at OSU, with outreach with an existing program (SMILE) and with the AndrewsLTER are carefully developed and very strong.

The post-doc mentoring plan is carefully thought through and addresses the likely needs of the post doc very well.

Broader Impact Weaknesses: no important weaknesses were identified.

Data Management Plan: The data management plan is carefully considered and comprehensive as well as striking agood compromise between data retention and dissemination

Synthesis and Recommendation: This proposal outlines a promising approach and a novel but very plausiblehypothesis to address an important question in population and community ecology. It integrates theoretical,

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statistical, observational, and experimental approaches in an integrated approach. It supports a young andpromising investigator.

The panel recommendation is: High priority

This summary was read by the assigned panelists and they concurred that the summary accurately reflects thepanel discussion.

Panel Recommendation: High Priority

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Review #1


NSF Program: Population and Community Ecology Program

Principal Investigator: Novak, Mark

Proposal Title: What Stabilizes Population Dynamics in Nature? A New Framework for Quantifying thein situ Strength and Nonlinearity of Species Interactions

Rating: Multiple Rating: (Excellent/Very Good)

REVIEW:

In the context of the five review elements, please evaluate the strengths and weaknesses of the proposal with respect to intellectual merit.

Summary: this proposal will apply a novel method for measuring consumer resource attack rates (from simpleestimates of density, and relative fractions of feeding individuals) to evaluate the degree of functional responsesaturation in consumers. It will use this information to test the hypothesis that generalized consumers do nottypically show saturation of prey consumption (and thus differ from specialized consumers). This would explain aparadoxical pattern that shows that generalized consumers do not generally show population cycles otherwisepredicted from consumer resource theory. To do this, initial approaches to the methods will be improved to betteraddress parameter estimation and to validate the method using modeling. The method will be applied to data onthree coexisting sculpins and a metaanalysis of existing data in diverse systems will be implemented.

Strengths: The PI makes a fairly convincing case that the approach is valid and powerful. He also makes a strongcase that the issue is a potentially important contribution to resolving the paradox. Finally the project has anexceptional integration of statistical, observational, experimental and theoretical approaches to a complex problem. Iespecially like the proposed experiment to directly test the idea that it is the range of prey densities that regulatethe degree of saturation in functional responses in generalist predators and that this range is exceptional. Theimplications also likely go beyond explaining the paradox of generalist cycles because they may also play animportant role in explain food web stability in general.

Weaknesses: Of course this all depends on validating this method (admittedly this is part of the idea of thisproposal). Although the claim is that purely observational data is sufficient to use this method, I'm a bit skepticalthat this will always succeed given possible confounding among multiple possible factors. I therefore see theexperimental approach as being at least as important and was somewhat surprised to see that it was not more fullydeveloped. Finally, while the research may be important in helping resolve this paradox, it says less about thepossible influence of other processes such as spatial effects, refuges etc.. (i.e. H3).

In the context of the five review elements, please evaluate the strengths and weaknesses of the proposal with respect to broader impacts.

these will consist of integrating research into teaching at OSU, participating with SMILE programs that connect toschools and public via Andrews LTER. There will also be support for graduate students and underrepresented highschools students.

Please evaluate the strengths and weaknesses of the proposal with respect to any additional solicitation-specific review criteria, if applicable


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I do think there are some risks to this proposal but I believe the PI has thought as carefully about these issues asone might hope. The payoff is twofold, first the implementation and improvement of a new way of measuringinteraction strength in complex food webs, and second the application of these methods to address an intriguing andimportant puzzle in complex food webs.

Summary Statement

I do think there are some risks to this proposal but I believe the PI has thought as carefully about these issues asone might hope. The payoff is twofold, first the implementation and improvement of a new way of measuringinteraction strength in complex food webs, and second the application of these methods to address an intriguing andimportant puzzle in complex food webs.





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Review #2





Rating: Very Good

REVIEW:


1a) The work could significantly advance understanding of a core issue in population ecology ûthe regulation ofpopulations of generalist predators embedded in food webs. Previous work, cited in the proposal, has shown that theclassic pre-predator cycles only occur with specialist predators and that there is very limited understanding of thefactors regulating populations of generalist predators, especially those (the majority) that do not exhibit cyclicdynamics. Generating predictions using traditionally paramterized models makes impossible demands on data: inprinciple, they require knowledge of the "strength" (appropriately defined) of nonlinearities in growth rates of allprey, and in the functional responses of all predators. Novak and Wootton previously developed and tested a methodfor quantifying the nonlinearities in functional responses using more readily available data (mainly "snap-shots). Theoriginal method required assumption of a particular functional form, but the proposal describes a generalization(equation 3). The plan is to use experiments on a stream food web to characterize feeding rates and interferenceeffects among 3 sculpin species, and then explore the stability of the resulting dynamics using an elegant techniquerecently developed by Thilo Gross, Ulrike Feudel and others. No mathematical trick can completely circumvent theneed to have information on prey production. However, the intention is to compare the range of prey growthparameters that would led to stability with those that would occur with hypothetical specialist predators. Ifsuccessful, this would represent a very significant advance in understanding.

2) The creativity in the proposal lies in utilizing two strands of recent theory û the Novak-Wootton methods forcharacterizing nonlinearities in feeding rates and recasting of standard population models by Gross and collaboratorsin a form that gives insights that are do not depend on particular functional forms û together with carefulexperimental design and consideration of statistical challenges, to make progress on an important question.

3) With one exception, the ground work on all steps necessary to achieve the project goals has been covered. Theexcpetion is the model framwork to be used for the generalized analysis. Sculpins reproduce annually, but thefeeding processes in the stream are continuous. Thus neither standard ODE-based models nor discrete time modelsoffer a good modeling framwork for the food web. A hybrid model, sometimes called "semi-discrete" would be farsuperior (see for example Pachepsky et al., Ecology 2008 or Mailleret, Phil. Trans. Roy. Soc., 2009). The Pachepskypaper in particular shows some of the effects on stability of synchronized reproduction. It also contains recipes forstability analyisis to which the techniques of Gross could be applied. This limitation of the proposal is NOT a fatalflaw, as the PI and collaborators could easily add this task to their panned work, especially as it will come near theend of the project.

The proposal contains no explicit statement of a plan to assess success, but many check points along the way areidentified, and alternative hypotheses are stated.

4) The team is well qualified, and the PI has a track record of successful collaboration.


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5) I am not qualified to critically evaluate the adequacy of the resources.


1b) The sceintific work has intrinsic societal value. River systems experience mutiple stressors and understandingthe atbility of their food webs is obviously an important concern.

2) I do not see how a proposal by an assistant professor could reasonably be expected to have broader impacts thatinclude " creative, original, or potentially transformative concepts". Indeed a key strength of the planned outreach toelementary schools is intgration with a larger university-school parnership (SMILE) at OSU.

3) The first question is hard to answer without knowledge of the local organization. There is no stated assessmentmechanism, but I would guess that SMILE has an assessment component.

4) The team has the ability to carry out the planned work. As a reviewer, I annot evaluate the other members ofSMILE.

5)The connection to an exisiting program suggests that adequate resources will be available.


Summary Statement

This is a strong proposal that addresses a "big" issue in population ecology. The planned work is creative andthroughtly researched. The only substative problem I identified is technical and can be addressed as the projectprogresses. The broader impacts include an outreach component that is integrated with a larger effort at the PI'sUniversity





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Review #3





Rating: Very Good

REVIEW:


The overall role of the predator's functional response to prey in understanding ecological dynamics is well-described;in particular, the relationship between the scarcity of predator-prey cycles observed in nature and the paradoxes ofenrichment and prey dynamics is honed in on, providing a solid rationale for the inherent interest of the propsedwork.

The PI proposes to advance a new class of observational methods for estimating the strength and nonlinear natureof generalist predator-prey interactions in the field so as to provide insight into disparities between modelpredictions and observed patterns. He will use the simulataneous multi-species aspect of his method to test threehypoheses: (1) feeding rates of generalist predators are not sufficiently saturated to elicit cyclic dynamics viz-a-vizthe paradox of enrichment, due to nonlinear additive effects of feeding on mulitple prey; (2) predator and preydensities experienced by generalists tend to lie in the narrow range wherein the interactions are approximatelylinear and thus do not exhibit very strong density dependence; and consequently (3) traditional theoretcialexplanations for the scarcity of predator-prey cycles are not needed to explain observed phenomena.

The PI has previously developed a way of calculating predator per capita attack rates based on empirical informationreadily derived from observational data, which allows simultaneous estimation of rates across an in principlearbitrary number of prey and predators. This calculation heretofore was based upon careful consideration of timebudgets for type II functional repsonses. The rate simplifies to:

(# of predator j feeding of prey i) / (# of predator j observed not feeding)

times (1/( h(i,j)* Ni))

where h(i,j) is generlized handling time and Ni is the abundance of prey i. In the proposed research the PI willextend the method to account for spatio-temporal variation in attack rates and for predator interference as inBeddington_DeAngelis functional respsonses. The simultaneous nature of these observational calculations is the keystrength of the proposal in my estimation and gives hope of addressing the hypotheses as described. Here the firsttwo hypotheses amount to asserting that generalist predator-prey interactions occur along a restricted part offunctional repsonse curves where density dependence is weak. If this assertion is borne out it would be a significantstep toward better concurrence between modeling and field work. In my view this prospect makes the study aworthy effort.

The proposal would have been strengthened had the type II computational formula in the project description beensimplified as above so as to make its meaning clearer. To be fair, the relevant paper did appear in Ecology and thetime budget analysis approach is quite familiar. However, until I simplified the quantity, I could not see that it had a


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very straightforward ecological interpretation. So maybe I am quibbling but I think the proposal could have beensomewhat more self-contined.


The broader impacts here are at the individual investigator lab scale, which is entirely appropriate for a juniorinvestigator.


Summary Statement

The proposal tackles an important issue at the interface of theory and empirics in ecology, and makes a strong casefor advancing conceptual unification. It appears to be scientifically sound and reasonably well-described. Thepotential ramifications are important discipline wide. The development of highly qualified personnel described isappropriate for an early career investigator.





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Review #4





Rating: Very Good

REVIEW:


This is one of the best proposals I have read. The PI clearly outlines the knowledge gap and why improving uponthis through the innovative research described in the proposal is important, necessary, and timely. It had notoccurred to me before reading this proposal just how interesting the simple observation is that nature is pervadedby generalists that do not exhibit population cycles. This begs for an approach such as that articulated by the PI thatconjoins experimentation with empirically-driven population modeling to explain the in general terms thecharacteristic dynamics of specialists (which cycle) and generalists (which do not or should not). The PI supposesthat prey-switching and predator interference are not necessary in explaining the absence of cyclic populationdynamics in many of not most natural systems. If the proposed work finds support for the PI's alternativeexplanation that this is due to lack of saturation of predator feeding rates, I can imagine this finding becomingintegrated into introductory ecology coursework.

The research objectives are clearly outlined and throughly described, and the PI has the expertise to conduct theresearch. I believe the outcomes of the proposed work will constitute and important conceptual contribution to thefields of predator-prey dynamics and population dynamics in general.


I especially like the training in quantitative methods the PI will provide to high school students. The PI's involvementin OSU's SMILE program will provide a meaningful integration of the proposed work with local primary education andresearch experience for school age children.


Summary Statement



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Review #5





Rating: Good

REVIEW:


The main goal of this proposal as stated in the proposal (see below) is to develop a new class of observationalmethods for estimating the strength of predator-prey interactions in species-rich communities. Several specifichypotheses will be tested regarding the nonlinearity of the functional responses of generalist consumers and thepropensity of population dynamics to cycle. The PI is a new investigator who has published a number of papers onthis topic, and the proposed work builds on his DDIG results. The proposed work promises to advance ourunderstanding of the mechanistic processes that regulate population dynamics in species-rich systems.

I found this proposal rather confusing. In particular, the proposal states that the main goal is to develop a newapproach for quantifying the strength of predator-prey interactions that circumvents the limitations of previousmethods. However, it appears that the new approach has actually been developed though an NSF DDIG award (DEB-0608092, Novak & Wootton 2008). What the PI would like to do is to fine-tune the approach (e.g., by using theBayesian hierarchical framework to estimate variation and uncertainty), test several hypotheses using data from theAndrews LTER site, and evaluate the generality of their new results by conducting a meta analysis of relevant data inthe literature. I acknowledge that I have not read the PI's previous publications (and do not have time to read themnow), and I am not well suited for evaluating the technical details in this proposal. I assume that the 'new' approachwas validated with empirical data in the previous project. So, what has been done already, and what is new thatconstitutes significant contributions from the proposed work? It's also very odd hat the Project Summary of thisproposal states that the main goal is to develop a new class of methods but never explains what these methods are.Similarly the summary says nothing about the three hypotheses to be tested through the proposed research.

Certain passages of the proposal are difficult to follow (e.g. Conceptual Framework, p.5, last full paragraph on p. 8)though they seem critical to understanding the proposed research. There is substantial technical detail throughout,but rather than clarify, it often adds to the confusion (e.g. Table 1 is inadequately explained, confusingly labeled,and contains many variables that are only defined sporadically in the text). The proposal would be stronger iftechnical text was replaced by a clearer explanation of what is to be done and how it is an important advancementfrom previous work.

Aspects of the proposed Bayesian analysis are unclear. For examples: -- on p. 9, there is considerable detail aboutthe prey distribution, but no explanation about how this distribution will be linked to a larger model. Surely it is notassumed that the same distribution of prey applies to all transects and sites? Is the idea to model parameters vand/or p as functions of covariates? In that case, they don't need priors, but process model specifications.



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The proposed research has the potential to advance the theoretical understanding of prey-predator interactions andpopulation dynamics. Results from the project may offer insights into the mechanisms of large-amplitude populationcycles, which increase the risk of stochastic extinctions. Thus the work has some implications for biologicalconservation. The PI is a new assistant professor, and the project would contribute to the establishment of hislaboratory at OSU. The PI will also sponsor a postdoc (with a good mentoring program), a doctoral student, as wellas undergraduate and high school students. The project will also provide science and math enrichment forelementary school students through an established outreach program called SMILE at OSU. The PI is a productivescientist, and I have no doubt that research results will be widely disseminated through journal articles andconference presentations. However, the style of presentation in this proposal is complex and confusing enough thatparts of the proposal are not easy to follow. Thus I am unsure whether the PI would be able to communicate theexcitement of math and ecology to the general public, especially elementary and high school students.


Summary Statement

This project will test several hypotheses by quantifying how prey-dependent, predator-dependent, and adaptiveforaging processes contribute to the strength and nonlinearity of generalist interactions and their propensity tocause cycles. The project has the potential to advance the understanding of mechanisms controlling populationdynamics in species-rich system. A focus on specific unresolved research dilemmas is commendable. However, thedistinction between novel research and the previous findings (Novak pubs) is not clear in the proposal. For example,the proposal repeatedly refers to development of "a new class of observational methods," but the observationalmethods (i.e. "snap-shot" gut sampling) seem to be the same as those previously developed.

The PI is a new investigator, and the project builds on what he has accomplished though a DDIG award in 2008. Thebroad impacts component is quite strong, and includes a variety of educational and outreach activities. The PI needsto make the proposed research less opaque to the reader. For example, the project summary states that the maingoal is to develop 'a new class of new observational methods for estimating the strength of predator-preyinteractions in species-rich systems' but does not explain what the new methods are; nor does it mention thehypotheses to be tested. This is the sort of thing that could potentially turn biologists away from quantitative andtheoretical ecology, and I wonder if the PI would make a major effort to explain what this project is about toelementary and high school students and would be able to attract them to participate (as proposed under BroaderImpacts). It appears that the new approach to be developed through the proposed research has mostly beendeveloped through PI's PhD project, and that the new project would fine-tune the new approach, test severalspecific hypotheses in a new system (in stream communities of the Andrews Experimental Forest LTER), and test thegenerality of the new results via a meta analysis of data from the literature.





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Review #6





Rating: Very Good

REVIEW:


It is true that the focus on generalists is often ignored.

There is a nice arc of building upon the expertise of the PI which takes the work in expanded directions, and comesacross as feasible given the intellectual background of the PI.

The proposal is motivated in part by an argument that type II functional responses destabilize abundancesùleadingto cycles. We see lots of saturating, type II functional responses in the world, yet we see lots of non-cyclingdynamics. The proposal discusses some of the proposed stabilizing mechanisms such as refugees, prey switching, ortype III functional responses, arguing that these are not satisfying explanations because they will not be veryeffective at large population abundances which is where the saturating features of the interaction are realized. Ifound this argument a little dissatisfying because prey density-dependence was never discussed. Prey density-dependence is now the textbook explanation, arising out of Rosenzweig-MacArthur, as to how predator-preydyanamics can be stable over at least some parameter ranges. And density-dependence does act at high densities.This argument is actually congruent with one of the main ideas in the proposal, that perhaps the effective non-linearregions in the saturating functional response are not that important given the range of densities experiences. Givenhow central this model is in the development of the theory and how nicely it relates to a thesis of the proposal, itwould be nice to see its inclusion in the body of the argument.

The author makes a clear and convincing case that we lack a larger understanding of generalist-predator speciesinteractions and functional forms.

The proposed work utilizes a technique that, at least in part, quantifies species interactions utilizing data of wildanimals in nature. The value and importance of this perspective cannot be understated. This proposal is a niceexample of merging observational data with a theoretical framework to make insights that might otherwise beimpossible with an experimental, ANOVA hammer.

This proposal builds upon a very elegant and powerful technique, developed previously by the author, whichquantifies consumption rates. It is dependent on natural variation existing across samples, and dependent on theability to observe key features of prey handling. These key features will not exist in all systems, but in enoughsystems The Bayesian formulation is an important and powerful extension of previous work based on randomization.

The author describes using non-informative priors in the Bayesian analysis, which is appropriate. This is a minorpoint, but informative priors may be valuable in some situations.

My guess is that the simulation-based power analysis will result in general lessons similar to the previous


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randomization based power analysis. That said, there is a good plan for execution of this power analysis and thework does need to be done.

The system specific implantation is a fascinating part of this proposal and illustrates a component in which theempirical and theoretical are tightly linked. On the one hand, the author is arguing the nonlinearity in field systemsmay not matter, which runs counter to some of the motivation of the proposal which is to quantify nonlinearinteractions in the wild. This is not really a weakness of the proposal, just an observation of where the proposal ispositioning itself.

The proposed use of Generalized Modeling in this work illustrates the use of some of the newest theoretical tools,and represents a timeliness to the proposal. The proposed work again represents a nice feedback of empirical workinto theoretical work. Personally, I think the reach of Generalized Modeling is a currently a little overstated, but it isexactly the kind of work proposed here that needs to be conducted so that the discipline can sort out the utility andinference available from Generalized Modeling. I would like to see a little more of an appreciation, or speculation, onthe limits of Generalized Modeling in the proposal, but to be honest that is really manuscript Discussion material andnot proposal material.


The proposal includes a collection of outreach and education projects under the SMILE program, with some focus onparticipation of underrepresented groups. They are all appropriate and important. The current proposal is a vague asto how these educational modules will be anything beyond the mundaneùnot that I doubt they will all be good, itjust isn't stated in the proposal. In other words, how do they tie into the intellectual work of the proposal beyondbeing good science outreach. That said, the leverage of an existing program and the inclusion of the programdirector as a PI is a great strategy for successful outreach.


Summary Statement

The proposal has a very tight link between empirical and theoretical work. Furthermore, it connects speciesinteractions with food webs in a mechanistic manner that is not typically possible given the large number of speciesinvolved. It utilizes new theoretical techniques in Generalized Models. The components of the proposal fit togethernicely in creating a large argument across all papers that will come out of this proposal. For all of these reasons, itpromises to be a fruitful and successful project. The broader impact activities are appropriate.

If I had to identify a major weakness, it is that this method will be data intensive. That is mentioned briefly in theproposal, but it would be nice to see this issue more directly addressed. To be fair, the details on this issue will comeout in the proposed simulation/robustness part of the project. Furthermore, the combination of laboratory analysison clearance times, as described in the proposal, will reduce some of the burden of inference that will need to becarried by the field data.

The other minor weakness is that the major intellectual leap has already been done in previous work. However, theproposed work makes important steps in expanding that work such that it has impact on the discipline.

I am somewhat concerned about the message that may come out of this workùnonlinearities can be detected ifsnapshots from the field, therefore they don't matter. Small nonlinearities may still be really important for thesystem dynamics, even if they are not easily detected in field data. For example, Fussman and Blasius (2005.Biology Letters) show the very different population dynamics that can occur from the choice of three very similarshaped functions. The authors could begin to address this question in the power analysis study and perhaps in thegeneralized modeling approach. Perhaps my concern with this is the very reason it should be supported, becauseadvances in ecology occur when we address the unaddressed assumptions.



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