I was never in theoretical ecology or really in evolutionary theory, but I've always had an interest. It's basically an open question how to deal with the interplay between the two. Ecology works on a much faster timescale than evolution, but the two both operate on a population-- so what are the true population dynamics?
I ran across https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10614967/ and it's almost like we had the same idea hit at the same time! I even wonder if maybe David Schwab went to the same APS March Meeting Session to get the idea. The basic idea is very simple: ecology is one set of equations, evolution is represented by another set of equations, and to understand the interplay between the two, you just couple the two equations. David Schwab and his co-author found the coupling-- mutations lead to new phenotypes, which then lead to a change in the ecology. Interestingly, the way that I would approach the problem is very different. It basically boils down to a different model and a totally different analysis that leads to solutions that can approach the fully correct solution more and more accurately. The model I would use for evolution would be the Fisher-Wright model, and the model I would use for ecology would be a master equation version of the generalized Lotka-Volterra model. Then, since ecology operates on fast timescales, you could solve for the probability distribution over species number in the generalized Lotka-Volterra model first, and plug it into the evolutionary equations to get a full solution. The generalized Lotka-Volterra model takes some explaining. Basically, in the generalized Lotka-Volterra model, there are three "chemical reactions": one birth, one death, and one death via eating by another species. If you write down the master equation for these reactions, you can use moment equations to find better and better approximations via a Maximum Entropy approach to the true probability distribution over the number of each species. The moment equations don't close, but you could artificially at some order just assume that the higher-order moments have an independence property of some sort. At the very least, you can get a Gaussian approximation to what the ecological equations might look like. Of course that's not correct, but maybe it's good enough. This avoids a thorny problem with pretending that the ecological equations solve quickly. In reality, the rate equation approximation to the master equation can yield chaos. This chaos goes away if you consider the fact that the number of species has to be a nonnegative integer. Whether or not these MaxEnt approximations are good is up to the field to decide, but you can get better and better approximations by including more and more moments. The Fisher-Wright equations are well-known to take one generation and produce a binomial distribution over the next generation's population number with properties that depend on selection coefficients and mutation rates. So basically, the next generation is sampled, it settles down to something having to do with a MaxEnt approximation, and you plug the resultant MaxEnt approximation into the next generation (convolve it) to see the probabilistic updates. You can automatically assume that this is the right thing to do because of the separation of timescales. Altogether, you essentially get a complete solution to the probability distribution over population number as a function of time. This isn't my field, and my schizophrenic voices basically don't want me to do this project, so I won't, but I'm hoping somebody will. I don't think it's a terrible idea. First of all, apologies that the hyperlink feature isn't working.
I've been keenly interested in nonequilibrium thermodynamics for a while, ever since graduate school. Originally, my idea for being interested was based on the idea that it provided insight into biology. This is not a bad idea-- the minimal cortical wiring hypothesis (https://www.sciencedirect.com/science/article/pii/S0896627302006797) says that neural systems minimize cortical wiring while retaining function. Why would they do that? Well, the usual reason is that the brain of the child has to fit through the mother's vagina, but you could also cite timing delays as being costly for computational reasons (such as in https://journals.aps.org/prresearch/abstract/10.1103/PhysRevResearch.5.033034) and also cite energy considerations via Attwell and Laughlin. And in fact, there is a paper by Hausenstaub (https://www.pnas.org/doi/abs/10.1073/pnas.0914886107) that suggests that energy minimization is crucial when deciding what kinds of channel densities and kinetic parameters still produced action potentials in neurons. And I have an unpublished manuscript (still in my advisor Mike DeWeese's hands) that suggests that eye movements are driven by energy considerations ala visuomotor optimization theory (https://www.nature.com/articles/35071081) reinterpreted-- maybe. However, I have been somewhat sad at the match between nonequilibrium thermodynamics and biophysics. Basically, the question is this: do Landauer-like bounds matter? A long time ago, Landauer proposed that there was a fundamental limit on the energy efficiency of engineered systems. This limit is far from being obtained. But you might hope that evolved systems have attained this limit (https://worrydream.com/refs/Landauer_1961_-_Irreversibility_and_Heat_Generation_in_the_Computing_Process.pdf), or something like it (https://journals.aps.org/prl/abstract/10.1103/PhysRevLett.109.120604). There is some evidence that biology is somewhat close to the limit, but the evidence is sparse and could be easily reinterpreted as not being that close. We are about half an order of magnitude off of the actual nonequilibrium thermodynamic bound in bacterial chemotaxis systems (https://journals.plos.org/ploscompbiol/article?id=10.1371/journal.pcbi.1003974), in simple Hill molecule receptors (https://link.springer.com/article/10.1007/s11538-020-00694-2), with a match to an improved nonequilibrium thermodynamic bound being given with networks that I am not sure are biologically realistic (https://journals.aps.org/pre/abstract/10.1103/PhysRevE.108.014403). There is a seeming match with Drosophila (https://www.pnas.org/doi/abs/10.1073/pnas.2109011118), which tends to have surprisingly optimal information-theoretic characteristics (https://www.pnas.org/doi/full/10.1073/pnas.0806077105), and also with chemosensory systems again (https://www.nature.com/articles/nphys2276). I am sure I'm missing many references, and please do leave a comment if you notice a missing reference so I can include it. This may look promising. To me, it's interesting what we need to do to get rid of the half an order of magnitude mismatch. Now, half an order of magnitude is not much. Our biological models are not sufficiently good that half an order of magnitude means much to me, so the half an order of magnitude basically has error bars the size of half an order of magnitude because we just don't understand biology. But beyond that, we should have more examples of biology meeting nonequilibrium thermodynamic bounds. I think we can get there by incorporating more and more constraints into the bounds. For example, incorporate the modularity of a network, and you get a tighter bound on energy dissipation (https://journals.aps.org/prx/abstract/10.1103/PhysRevX.8.031036). I want to use nonequilibrium thermodynamic bounds in my research, but haven't for a while. I just need better bounds with more constraints incorporated. Does the network you're studying have modularity or some degree distribution? Are there hidden states? Is it trying to do multiple things at once with a performance metric for each, e.g. not just Berg-Purcell extended but Berg-Purcell revisited so that we are trying to not just estimate concentrations but predict something about them as well at the same time? Is the environment fluctuating and memoryful, which was one way in which the Berg-Purcell limit was extended (https://journals.aps.org/prl/abstract/10.1103/PhysRevLett.123.198101)? I'm going to talk to a real expert on this stuff in a few months and will update this post then to reflect any additional links. Please do comment if there are links I've missed while being out of the field for years. I am quite curious. |
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May 2024
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