Sean Carroll's Mindscape: Science, Society, Philosophy, Culture, Arts, and Ideas - 185 | Arvid Ågren on the Gene's-Eye View of Evolution

Episode Date: February 21, 2022

One of the brilliant achievements of Darwin's theory of natural selection was to help explain apparently "purposeful" or "designed" aspects of biology in a purely mechanistic theory of unguided evolution. Features are good if they help organisms survive. But should we put organisms at the center of our attention, or the genetic information that governs those features? Arvid Ågren helps us understand the attraction of the "selfish gene" view of evolution, as well as its shortcomings. This biological excursion has deep connections to philosophical issues of levels and emergence. Support Mindscape on Patreon. Arvid Ågren received his Ph.D. in Ecology and Evolutionary Biology from the University of Toronto. He is currently a Wenner-Gren Fellow at the Evolutionary Biology Centre at Uppsala University. Previously he worked at Cornell and Harvard. His recent book is The Gene's-Eye View of Evolution. Web site Google Scholar publications Amazon author page Twitter

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We contrasted this human-scale way of thinking about the world, the social world, where you make up categories, and then they're based on what really happens, but there's a lot of freedom there. We contrasted that with the physics level, right, with the electrons and the photons, where it's pretty clear how you want to think about the pieces out of which reality is made. But there are a lot of layers in between the physics level and the social level, the human level, right? There's all of biology, for example. Today we're going to dig into a puzzle, or at least an important research area within biology, that involves exactly this. What is the right way to conceptualize the world of species and populations and evolution?

Famously, Richard Dawkins in the 1970s popularized a view known as the selfish gene. Back in the old days, in the beginning of the theory of evolution, you might have focused on individual organisms and imagine these organisms wanted to reproduce their own genomes, right? They wanted to have kids and have their genetic heritage be passed on to future generations. Then came the math, then came a way of thinking about population genetics that pointed out that you can pass on the genes inside of you, even without you being involved, if all of your relatives are very good at passing on genes. So it became not only Dawkins, but other people point out that it's as if it's the genes that want to pass on their heritage, not the individuals carrying them.

you could say that the individuals are just kind of a bus full of genes. They're a vehicle that carries on the genes because individuals die, right? Organisms are born, they have a life, they die, less than a century for most species, whereas the genes can live on a very, very long time being passed from generation to generation. So surprisingly, not everyone agrees, not so surprisingly, of course, this is a complicated thing. What is the best way to conceptualize levels of selection. This is what we call this particular debate in biology. Is it at the level of genes, of species, of populations, of individuals? And it's exactly this kind of how do you divide up reality kind of question. The answer from our guest today, Arvid Ogren, who's worked as a

biologist on this problem, is he's actually pretty pro the selfish gene view, or as he calls it, the gene's eye view of evolution, but he completely admits that it's not the end all. Biology is more complicated, as we've often found out here on Minescape, than something easy like physics is. There's leakage between the levels of genes and molecules, up to organisms, up to populations. So there's insight to be gained by thinking about things from the selfish gene point of view. There's also insight to be gained by thinking about higher levels of abstraction. It's a wonderful real-world example of the philosophical questions that we have about emergence, complexity, how to chunk the universe up into pieces that we can analyze and theorize about.

So I think it's a great kind of discussion. It's very mindscapy as I make the joke in the podcast. Both me and the evil Sean Carroll, the biologist, who is a previous Minescape guest, are referenced in Arvid's book. So it's a very natural kind of conversation for us to have here. Let's go. Arvid Orgren, welcome to the Minescape Podcast. Thank you for having.

me. So I think we can assume that most of the audience is familiar with Darwinian evolution theory, natural selection, and maybe also a little bit of Mendel and genetics. But let's start with the groundwork at the synthesis of these two great ideas, right? The modern synthesis. It still boggles my mind a little bit that Darwin didn't know about genetics. He went pretty far without knowing about that. But tell us how that came together in the 20th century. Yeah. So this is really one of the kind of striking moments in the history of biology, what has become known as the modern synthesis of biology in the 1920s, 1930s. And as you say, this kind of came together by bringing two different insights into one theory. On the one hand, it was Darwin's theory of evolution by natural selection, and a theory

about individual organisms, organisms vary in how well they survive and reproduce, and if any of those traits are heritable, they will become more common over time. Now, the big problem for Darwin then was, as he said, he said, he said, he said, he said no functioning theory of inheritance. And while he kind of had some ideas of his own, none of them really came to work. And that remained a conundrum for him and kind of viewed as a weakness for a long time of his theory. This was in parallel then with the second insight that was brought to it. And that was a functioning theory of inheritance. And that was provided by Gregor Mendel's work, where the realization that inheritance functioned with its inheritance

of this discrete particular entities, which we now refer to as genes. And so, I mean, if we didn't know about genes and we didn't know about how discreet they were, clearly there's some degree of inheritance. Everyone can see that in hair color and things like that. But you might have thought that things would just blend together every time, right? And then you would never diverge, everything would just become some kind of homogeneous unified organism or something like that, in terms of sharing the same equilibrated characteristics. But with discrete units, all sorts of funny things can happen. Exactly. And indeed, that Darwin for a while believed in his sort of blending inheritance, and that the realization that you were quickly run out of variation

for natural selection to act upon was a kind of serious issue for him. The problem then with this new fear of inheritance in the form of discrete entities was that how could that be reconciled with the gradualism of, on the one hand, variants as we see it in the natural world, but also with a gradualism as envisioned by Darwinian evolution, the gradual change over time. And this is really what the emergence of what we now referred to as population genetics, the idea that you can describe evolution as changes in certain variants of genes or allele frequencies over time. That is what kind of what evolution almost has come to be defined as if you open an evolution textbook today, that the evolution is changed in allele frequencies

in a population over time. So I, I guess I don't quite see how that answers the question of the apparent smoothness of variation. I would have just said that variation is smooth because there's a lot of genes and you can just change one at a time. It looks almost like it's continuous. And exactly. That is what the solution that population genesis, strong upon, in particular the English statistician and evolution genesis, Ronald Fisher, showed that you could get the appearance of gradual variation from the, if a trait was caused by many alleles with small effects, you could get. You can reconcile the gradualism with the discrete inheritance. And that really represented this major achievement.

That was part of, and it is generally synthesis of bringing together multiple parts of biolid into one. I think you already explained it, but I'm going to ask you again to explain the word allele, because it's not exactly the same as the word gene, right? It is not. No, so allele is a version of a gene. So, for example, we may think of a gene for, say, eye color, but then you can have multiple alleles, one for blue eyes, one for brown eyes, for example. Okay.

So we have, what we can say we have one gene for that as a human. We have one allele from our father and one allele from our mother. that we have inherited. So I think probably the person on the street would talk about a gene for blue eyes or a gene for brown eyes, but really there's one gene for eye color and there's an allele for brown eyes or blue eyes. Yeah. Yeah.

And then in most of the cases, you have many genes with multiple alleles underlying most traits, most of the time. Okay. Very good. And what is a gene? Ah. I ask everyone. I ask every biologist this question.

They always get uncomfortable when I'm. I ask it. It's a practical little word in biology that has come to mean quite different things, depending on what corner of biology you are from. In kind of theoretical population genetics, it is simply something that kind of stably inherited, something that you can say affects a trait to a molecular biologist has a much more physical or materialistic definition of it, that is the sequence of DNA that often has a function or encodes a protein or RNA that does something.

And indeed, it's kind of multiple definitions of genes within biology has often led to misunderstandings and disagreements. So I guess even when we had Mendelian genetics, Mendel was still in the 19th century, but then it was sort of glued on or incorporated into evolutionary theory in the 20th century. But we still didn't have DNA at that point, right? So, I mean, happily, we have DNA now.

change the way we thought about this a lot or did it just get serving as an underpinning for the existing discussion about genetics? I think the first approximation it changed a lot in what we think of what genes are. We can study in a completely different way. We have a son so the number of genes that a species has and how they evolve. At the same time it is important to remember that population genetics as a an approach as a field, as you say emerged before we we knew what the material basis of Heresy was. Before we knew that it was DNA, before we knew that DNA was a double helix. And those mathematical models that we still use in evolutionary biology

were developed before that, before we had any idea of those. And there you rely on a definition of a gene that is completely agnostic about the material basis. And I do think it's rather remarkable that they work so well despite being completely ignorant about what the material basis actually is. Except that it let us invent a new meaning for the word gene, right? So we had the word gene, and then we found DNA, and then they invented a new meaning for the word gene, namely a sequence of DNA that encodes for a protein. Yes, yes.

I mean, the population genetics gene, as we use it, still in population, is much closer in a way to the original Mendelian gene, as kind of as it was originally. But it is certainly one of those words, one of many in biology that have similar, enough meaning that people can think that they are talking about the same thing, but they are different enough to cause frustration. Well, you've written a book called the Gene's Eye View, and what do you mean by the word Gene in the title of your book? Or does it not matter?

So the Gene's Eye View uses the word gene in this much more old-fashioned way of simply defining something that's stably inherited, a part of a chromosome that's stably inherited across generation. And this means that it, can be almost of arbitrarily long a gene. It can be anything that's kind of inherited together, which means that the parts of the Y chromosome, for example, that never recombines with the X

is like, can be thought of as one gene or kind of large swastal chromosome. Or indeed, very small part of a chromosome. It's also thought of as a gene here. But it is kind of this definition agnostic about any sort of material basis. And how do we know,

How much do we know about the genetic composition of, let's say, the human genome? Does everyone agree that it's divided into so-and-so-many genes, the human DNA? To first approximation, yes. I think everyone is in the same order of magnitude. I don't know exactly what the latest number is. When I was taught it, it was somewhere between 20 and 25,000 genes. I think someone who, you know, with the latest updates of the human genome, you're going to find a much more precise number. And I think I'm sure experts disagree, but to first approximation, I think everyone kind of agrees that this is the number. This is all off topic a little bit, but since I have you here, I'm just sort of satisfying my curiosity about a bunch of these questions.

Do we have differential data about the persistence of different genes over time? I mean, are some genes very malleable over evolutionary timescales and other? stick around? Yeah. No, certainly we know that we have something called pseudogenes, which are genes that are still around. They're still recognized as genes, but they've lost their function in humans.

So, for example, my favorite example is that we have the gene that allows you to synthesize vitamin C from scratch, which is present in most mammals. Yeah. But so we have that gene. It's clearly recognizable, but it doesn't work. We don't do it. It's in us, some of our primate relatives, and I think in fruit bats that have lost that ability and carry this around.

So there it's around. Selection hasn't gotten rid of it. But they're also certainly an example of where if you compare species that are distantly related, you can see that they've retained some genes in common, but sometimes they've lost genes or gained other genes. Hey, everyone, it's Cal Penn. And I'm the host of Earsay, the Audible and I Heart Audiobook Club. This week on the podcast, I am sitting down with Ray Porter, the narrator of Andy Weir's audiobook Project Hail Mary, massive sci-fi adventure about survival and science.

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That's great. Because it served the story. People will say like, oh, my God, I cried at the end. It's like, yeah, dude, me too. Listen to Earsay, the Audible and IHeart Audio Book Club on the IHeart Radio. app or wherever you get your podcasts. My best skin ever at 45? Give me a theme song and a best skin care award because it feels like this.

Right there. That's farmhouse fresh skin all right. I'm blowing and everyone asks how. The best skincare is farmhouse fresh and the award is you, your best you. Visit farmhouse fresh skin care.com and use code radio for a free starter routine with any purchase. So in my naive physicist's way of thinking about things, I would imagine that in any stretch of DNA, there's sort of a chance per unit generation that a mutation happens. And in those

stretches that really matter to our functioning, if a bad mutation happens, it gets weeded out, it gets selected out of the population. And in places where it doesn't matter that much, those can just continue to mutate away. So is it true that certain stretches of DNA maintain their integrity longer than we would think because they're really, really important to us? Yeah, and indeed, that is often how at the heart of the methods that are used

to estimate how much natural selection is acting across a genome. So if you think of all the DNA in an organism, we can ask questions like, how much of this is under constant natural selection and how much of it is evolving just by chance or neutrally. as we say, which is going to be a function of things like mutation rate and population size, essentially. And this is something like how much questions like how important is natural selection versus

evolution by randomness or genetic drift used to be a quite contentious debate. Okay. Population genetics, but one that has matured a lot in light of the the abundance of data that we have now and is much more kind of empirically informed, which is a much more insightful now because you simply go out and ask how much of this genome in this species is under selection and so on, which we couldn't before. And let me guess. The answer is that some parts selection is really important and other parts it's not. Yeah. And in some species, most of it is under selection and some large sports is not.

And am I right to think that maybe in the back of our minds when we think about selection pressures, we're thinking about, you know, some species going out there in hunting and becoming faster or stronger and therefore competing? but really a lot of the selection pressure is just that if you get the wrong mutation, you die or you're still born. It's not really that sort of detailed capacity to get food or have sex or anything like that. Yeah, I think that that represents obviously a crucial part of fitness for any organism, but there's so much more going on in the life cycle of any organism. A lot of it is that we can't see or it happens at the molecular level.

And often those machinaries are evolutionary old and often quite, sophisticated in keeping check that mutation rates are not too high or kind of the repair mechanisms are in place. Well, I guess so is it, again, my naive physicist thing is every base pair would have a random chance of mutating. And then the selection acts upon those mutations. But is it, are there even more sophisticated repair mechanisms than that? Are there ways biologically or molecular biologically that the really important parts of DNA are protected from mutation? Oh, that I don't dare answer that in a confident way. How that varies. Certainly, selection is stronger on important parts of the genome.

And I think, like, for example, that we carry around this gene for vitamin C, where you can kind of accumulate almost any sort of mutation without selection doing anything about it. how this kind of the efficacy of the kind of DNA repair mechanisms vary across the genome. I'm just I'm just wondering, you know, if there's in quantum computing, we care a lot about what we call error correction, right, or even classical communication theory. I can imagine that it would be evolutionarily advantageous to develop a cellular biology level mechanism that would just say, no, no, I'm not going to let any mutations happen in this part of the genome, but I have no idea whether or not that's actually true.

Yeah, now I'm curious too. Okay, good. We'll bring you back if you invent one. Okay, so I think that I completely interrupted you multiple times there while you were trying to explain the modern synthesis. What is the modern synthesis at the end of the day? At the end of the day, the modern synthesis was kind of, in a way, the founding of the feel of evolutionary biology as we know it today. It is an event that brought together rather quite disparate fields of biology, everything from kind of paleontology to ecology to genetics into one cohesive science, the science of evolution. So on the one hand, it was kind of this crucial event of showing that, I mean, Dilean inheritance and Darwinian evolution are compatible and part of the same process.

But in many ways, you can also think of it as a big institutional event. That's when the Society for the Study of Evolution was founded and the journal. and kind of starting the emergence of evolutionary biology as kind of as a department in university and so on. So kind of the modern feel of evolutionary biology was really born in that event as well. It's interesting. I bet you don't always see. I'm just saying, again, I'm completely speculating. I wonder how often, I should say, you see a parallel evolution of scientific knowledge and institutional support. or institutional organization for thinking about it.

So you're saying that in the case of evolutionary biology, they went hand in hand. I would think so. I've recently kind of come around to this view of that part of the history of our, of the field of evolution and the importance of the kind of institutional aspects of building a new field. Yeah. And especially that this was, so this kind of perhaps happened in the 1940,

it was just before the emergence of molecular biology. And it was in some ways you can think of it. This is almost this assault on old-fashioned natural history, organismal-based biology, but it's more modern science, which was considered to be the future in kind of, and recently this is Edward O. Wilson at Harvard describes it very well. And he's kind of, when he was hired at the department, was then the Department of Biology at Harvard, around the same time as Jim Watson, co-meanor of discovery of the double helix,

and kind of their battles of what, by all you ought to be like. And Wilson described this rather nicely in his autobiography, a naturalist. And that kind of combined with that, we have recently had the 75th anniversary of the Journal of Evolution, the Evolution Journal. So it kind of made me reflect upon that part and the importance of people like Wilson. Yeah, I mean, it is really fascinating because, of course, we hope that science gets at some true facts about nature. But we all know that it's done by human beings. And it's not just done by individual human beings, right?

Getting support from an institution, having jobs, getting funding, all this stuff matters in a really interesting way. And, of course, the other thing that matters that happened, I think, around that time is math, right? I mean, once you were beginning to be thinking about evolution in terms of population genetics, population is a number. And you can talk about the fraction of different genes and suddenly a bunch of equations first on the scene. It's very much so, and population genetics remain imminent with the theoretical backbone of evolutionary biology. Evolutionary biology, I would say, is part of biology that is perhaps the most mathematicized and kind of population genetics, often borrowed models from physics, statistical physics, and similar kinds of models, describing changes in allele frequencies from how physicists have described motion of objects.

Of course, there are other traditions. Evolutionabiali happily stole game theory from political science and economics and incorporated perhaps even more successfully as a way of doing things. And there are also other kind of quantitative methods, for example, in what they know is quantitative genetics. We simply describe changes in phenotypes over time, for example, used in animal breeding and so on. But I would say the kind of the emerging population genetics and during the modern synthesis was instrumental in some ways in making evolutionary biology be viewed as a more exact science and something to be taken seriously. I mean, it's certainly true that once you write down equations, even if your substantive content is exactly the same, you suddenly look more respectable because now you have equations in your papers.

People go, oh, okay, now you're doing serious work. Yeah, especially when you convince others you are not shared the same obsession as you are that you are to be taken seriously. It can be proven to be quite important. And when we say that there is math, I mean, what are the quantities that are being followed by these equations? In physics, you know, we have the position and velocity of particles. So what are we keeping track of and deriving differential equations for in population genetic? Population genetics have typically been considered kind of the frequencies of a specific allele in a population. And you kind of can say if you have an allele with these properties that has, say, this effect on fitness,

under what circumstances can it invade a population? So can it become more common as the generations go by? And also things you are interested in human track of are things like mutation rate, how often does this mutation happen, the specific mutation, population size. So in large population, natural selection will be more effective, whereas in small population chance will play a bigger role. If it's strongly advantageous,

So it was known as the selection coefficient. So how strongly is the selection acting on this specific allele? Those, I would say, are kind of the most basic parameters. And then you can introduce all sorts of complexities in terms of population structure and the relatedness between individuals interacting assumptions about the life cycle, whether you sexually or asexually and how mating is determined, and almost any aspect that you're interested in. But in kind of traditional population genetics,

these are the kind of the basic parameter of population size, mutation rates, selection coefficients and stuff. Well, that's interesting. I think we run immediately into some philosophy of science questions here because things like the frequency of alleles or the mutation rate, those sound very measurable. You know, if I were a positivist, I would say, that's good. I can see what's going on there.

But then you have some kind of model, right, some kind of theoretical framework that it fits into. So you already talked about selection coefficients or fitness. I mean, maybe explain a little bit more about what fitness is supposed to be and how empirical are those ideas versus concepts we need to introduce to make sense of our equations. One of the architects of population genetics, the Brit J-Bis Holden, once described fitness as a bugger.

It's one of those kind of central entities in evolutionary theory. In many ways, you can say we are kind of one trait discipline. The end of the day, that is what matters. At the same time, it is awfully hard to kind of agree on a proper definition and even harder to measure in context that really matters in natural population. Usually we think of it as something along the lines of the genetic contribution that a given individual makes to the next generation. So how much the genes in certain individuals,

how much they contribute to the next generation. But then we also talk about fitness of specific mutations. So then we're talking about things at the genic level. We have a long debate whether it's best to define fitness only at the individual level or sometimes have notions where we also should account for the interactions we have with individuals around us, this notion of inclusive fitness. And then this is even before, we try to measure anything.

So if you, those who study kind of evolution in the wild, for example, you want to compare, if you have, say, you have sample individual plants from two different populations

and you put them in a third one and see which one has a higher fitness. Well, then you often have to rely on like, well, make something like a number of seeds that they produce or kind of maybe even something like size and all these kind of indirect measures of fitness,

which means that it is this, central entity that has been a lot of both conceptual debate, how we should auto define it theoretically, but also kind of debate about how do you measure it and what are good kind of indirect measures? Because that's usually what we have to rely upon at the end of day, if you want to study it in natural populations. When I wrote my book, The Big Picture, I talked about fitness landscapes a little bit. I talked a little about evolution. And when I ran it by some biologists, some thought it was fine. Others are like, why are you talking about fitness?

This is an outmoded concept. We can't measure this. This is, it's one of those, they thought that it was one of those things that, you know, gives you the illusion of understanding. But when you get right down to it, it's hard to really quantify in a reliable way that everyone would agree on. Yeah, I think it's often like to do it well, you would have to keep track of the features, not only of the focal individual that you're interested in, but also of others in a population in a way that's really quite hard in the in the long run, especially if you study anything other than plants or kind of, kind of. things in the lab, like you can do it really well with fruit flies or microbes in a controlled lab setting where you can keep track of anything. But if, like I do believe that at the end of the day, what truly matters in biology is try to understand organisms in natural populations, it is much

harder. Well, for one thing, there are things like floods and earthquakes and asteroids hitting the planet and, you know, it's hard to predict ahead of time what your fitness is going to be in those circumstances. Yeah, yeah. Pandemics. Okay. So, but anyway, to get back to the main line of thought here. We've mathematicized a little bit of population genetics on the basis of the modern synthesis and what we realize

correct me if I'm wrong, my impression from looking at your book in other places, is that it's not just about me and my offspring. In some sense if you can speak of

these goings on in slightly overly anthropomorphic language, what wants to happen is that your genetic heritage wants to be shared. And one way to do that is to have kids, but the other way to do it is for your brothers and sisters to have kids, right? And so we get the idea of kin selection and inclusive fitness. So this is an insight that appears perhaps in the generation after the architects of the modern synthesis.

So in the 1960s, and at the heart of this is this English biologist William Bill Hamilton, who already as a struggling graduate students write these two tremendously important papers. One published in 1963 and the other. It's a two-part majestic piece in the following year in 1964. And there he introduces both concepts, both kin selection, which is not a term that he coined, but it was coined by someone else, but which is simply the idea that selection involves you and the relatives around you. But at the heart of it was his idea of inclusive fitness.

and to kind of get a sense of why he thought we needed an update of the basic notion of fitness. You have to think of situations where the classic definition doesn't really work. And those often happens in the context of social behaviors that we had known for a long time that in social insects, like in bees and ants, you have individuals that are completely sterile and instead devote their life to helping other members of the same population to reproduce. the queen. This is recognized already by Darwin, who described as his one special difficulty of his theory. And what Hamilton realized that one way to explain this is to think of that you shouldn't just think about the organisms, the offspring that a specific individual has, but it should also

account for other offspring that this focal individual is causal responsible for, and you should add those to our focal individual's fitness scaled by the relatedness that you have to those individuals. So in a way that if your brother has it, that counts more than if your cousin does it and so on. And then it also has to do this, to do the mass properly, you also then have to subtract from this sum be part of your or the focal individual's personal fitness that someone else is causally responsible for. And doing this kind of sums is where it can get mess pretty quickly, but that is basically the hard. You end up with its quantity, inclusive fitness, that you can show that individual can

appear to maximize that. Then under certain circumstances, inclusive fitness can be maximized even though this individual does not personally have any offspring, but they can help enough others to do so. And then still, you can have the process of work whereby natural selection maximizes a kind of fitness, but it's a new definition of it. And this is kind of what emerged in the 1960s. And does the witty remark from JBS Haldane that he would lay down his life for two brothers or eight cousins, right?

Exactly, exactly. That is kind of predates actually this, but it's very much sums up the basic insight very well. You're confused about your credit score. One site has one number and another site something completely. What? That can't be right. It's okay.

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and the Quaternians and things like that, two different William Hamilton's. Also, there are two different Sean Carrolls. And there's a biologist, Sean Carroll. And when I got your book, I think you will laugh at this. When I got your book, I realized that this is a rare book,

given that you wrote about levels of selection and sort of philosophical things that I care about, but also biology and evolution that the other Sean cares about. This is the rare book that might have both of us,

in the index. And indeed, there is one entry for Sean Carroll in the index, but it refers to two different appearances, and one of them is about me and the other one is about the other one. Oh, is that true. That's great. So it's a very Sean Carroll-centric kind of topic that we're talking about here. That's what I'd like to see. Yeah. Okay. So, yeah, we get the idea that the motivation for thinking this way. I mean, maybe the motivation is that it's true, but part of the motivation is we see in nature individual organisms acting in ways other than just trying to have the most offspring as possible. And maybe this kin selection or inclusive fitness can account for that. Exactly. That has been a tremendously influential way to think and make sense of these observations.

It is also an approach that has been criticized from the get-go, often from mathematically-minded parts of the field and the kind of a criticism that continues to this day, that the way that you, it's not as kind of mathematically robust of a construct as some people with, perhaps a more mathematical, strong background would like it to be given the centrality that it has in evolutionary biology. It has kind of survived because it provides a tremendous, this powerful rule of thumb and a very is something that kind of empirical biologists and fibres can go out and try to guide their experiments. And to kind of cut this long debate short, my kind of impression of it is, is partly what you want models to do in evolutionary biology, how kind of comfortable you are with these kind of mathematical limitations. Because at the end of the day, this kind of summing up what, at the addativity or summing up what you are responsible for versus other parts of the population, often is, becomes,

prohibitively difficult in other cases and the most simple ones. Similarly, you have to make assumptions about genetics and how strong selection are and so on to make it work. But it has pointed us to something really valuable. And that, perhaps one way to sum up is the third leg of the kind of Hamilton and contribution to evolution body is the kin selection, the inclusive fitness. And sometimes what is known that is Hamilton's rule that a trait will be favored if the benefit times, the relationship is greater than the cost, which is kind of summed up in J.B.'s Holdings.

I laid down my life with two brothers, right cousin, quip. The cost being that he would lay down his life. Exactly. And that as a guiding principle has been very helpful. Okay, but I mean, are you alluding to the criticisms by mathematicians? In that, are you alluding to Martin Novak and Tarnita and Wilson? They had this paper about group selection and use sociality a few years ago that really stirred things up. It really quite did. And I thought in some ways, quite exciting way, I had just started graduate school when the paper came out.

And so it's kind of been with me. Your whole life. Ever since, it was kind of one of the first paper we read in Journal Club when I was a new grader student. And I kind of was really taken by these kind of conceptual debates and really kind of reinvigorated in my interest in that part of the field. So they in some ways represent the latest iteration of the debate. They are far from the first to make these kinds of criticisms of. inclusive fitness. They were kind of made from the get-go in a way,

that given the centrality that is taken, it's not as mathematical robust as you would want it to be. But it's a bait that perhaps has calmed down a little bit the last few years, and I think we are in the field once the dust has settled is better off. I think we're kind of more on the same page of what the limitations are. And you can kind of have a more, I think, a more interesting discussion about when can you live with those limitations and when can you not and kind of what the the benefits of the approach are? And then I think I'm quite comfortable with biolids at the end of using different approaches depending on the kind of temperaments and kind of what they want the model to achieve in a specific situation.

But I guess I'm kind of confused a little bit because the inclusive fitness idea, I mean part of what it wanted to do, at least in principle, was to explain not only why certain ants don't reproduce and nevertheless contribute to the life of the colony, but also maybe even altruism in human beings or other mammals and so forth, it sounds like it is quintessentially mathematical. Like, you know, you, Haldane's joke is very mathematical, right? You know, two brothers or eight cousins. But you're saying that the criticisms have come from the more mathematically inclined? Why is that? Is that explicitly So I think that there are parts of the concept is kind of that relatedness is going to be really important. It's in a way mathematical, but it's also quite simple intuition.

So it led a lot of kind of feel like you can go out and measure what are the relatedness between interacting individuals in this specific population and the kind of studying that in specific population in the wild. You can do it phallogenetically by comparing, if you look at this group of species that have developed this quite complex social behavior, do they have a population structure of higher relatedness? So you have interacting generally between relatives compared to this part or this group of speeches on another part of a phallegeneratic tree. That do they have less relatedness among interacting individuals? So it's kind of guided us a lot in those kind of approaches. And that kind of like kin selection ideas there, I think has been. mathematical, but they're not that, not too complex in a way. They just kind of points us in their right direction.

Okay. Now, to get inclusive fitness, so, inclusive fitness has, to its proponents, one of, a kind of uniquely good property of it is that it's a property of the individual organism. That is, and it can be modeled as a property that an individual organism should appear as if trying to maximize. And this is good, because then. you can kind of treat it as the kind of design the maximum of or design principle

of evolution or natural selection. And this kind of goes back to some part of evolution biology, the key problem that we should try to explain is the appearance of design. That like the fact that organs appear so perfectly suited for the environment that they live in, that's kind of adaptation or appearance of that is the problem we should try to explain.

And you then take a step back and say, in order to do that, we ought to have a kind of a principle that shows like what should organisms appear designed for? And to its performance, and inclusive fitness is that, that an organism should appear trying to maximize that. And you can derive a model by which you can get the expectation that that is what the organism should try to do, to try to do in inverted commas. And that is that when you try to get inclusive fitness to have that property, that it should be both what should try and be maximized and be the property of an individual.

In order to get there, the maths is a little shakier in a way. You have to rely on more restrictions either to get the work or the assumptions are considered to be almost the opposite problem. There are too general to really become useful, because they almost end up with this kind of truism where it's always true. But there is where I think more mathematically, even more mathematically minded people have had its criticisms. And I should say, all of these people are at the max the end of the spectrum

within biology. But if you come there from a biologist by training interested in theory versus someone who has the original training in, say, math or physics and then come into biology in graduate school or even later on kind of degrees of mathiness. And so, again, just to get the landscape, as it were, perfectly clear, is the alternative to inclusive fitness or is the alternative to kin selection, group selection, or is it more complicated than that? I'd say it's a alternative. Historically, that has often been the kind of viewed as the two contrasts. And what does it mean, group selection? The group selection, again, is one of those terms that mean different things to different people, and that is part of the problem.

And it has historically kind of meant two slightly different things. Again, coming back to how we measure fitness. So by now, so I say kind of historically, the first way we thought of group selection It wasn't kind of, whereas now often even by its proponents referred to as naive group selection, this kind of fully that individuals may sacrifice their life for the good of the group, meaning for the good of the species. And that's also kind of done in a rather unreflexive kind of way. But then you often define group, kind of group fitness in terms of groups producing new daughter groups

that like you measure group fits in terms of new groups produce. Most today, most models of group selection. So if you pick up one of the current journals and someone can send the group selection model, fitness is usually measured in terms of individuals, producing other individuals, but that you account for some sort of group properties, some part of the group that's not really reducible in a meaningful way into individual level, kind of individual level properties. That is what now these days meant by group selection.

I can sometimes think that it's a little unfortunate that that is Kim, known as group selection, but that is, you know, me and others lost that debate a long time ago, that this is what we've settled on as a field. But those models then can can often account for the same things and often rely on similar kind of properties that a kin selection model would do. So one kind of observation or insight that in some ways calm things down in this kind of ongoing debate between inclusive business and group selection is that you can show that often they are kind of formally equivalent, but they give the same prediction. And I think that that is a good thing to realize that often you have the same properties.

There's an interesting kind of, I think, more philosophical point there that often this equivalence claim relies on generating rather abstract statistical models that you get almost quite far removed from the kind of causal processes happening in order to achieve that kind of equivalence. and I'm kind of two minds of this. Sometimes I'm kind of thinking, oh, this is great, I wish I'm not their equivalent. Sometimes I'm a little concerned that like this equivalence relies on such an abstract notion and that surely we are interested in what causally happen in a specific event, whether it is better described in terms of inclusive fitness, which is an individual level property or in terms of group level properties. Good. Well, this is, that's a perfect segue because I do want to finally hit.

the payoff here, which is that, you know, once you start thinking about things in this way, evolution in this way, of population genetics and equating my life to that of an equivalent number of other people's sharing the same genome, that's when you get to the genes-eye view, right? That's when it begins to maybe make sense that what we should think of as competing in the world of evolution is different genes rather than different organisms. Yeah, I think the origin story of the gene side view, which is a perspective that kind of really comes to his own in the 1960s and 1970s, stand on the, it's exactly these three topics that we have covered.

It's a approach to evolutionary biology that takes adaptation as the central problem that we're trying to explain. It builds on the insight from population genetics that evolution can be described as changes in allele frequencies. And then the kind of the third debate from which emerges is the one over group versus individual level selection or the kind of was sometimes known as the broader levels of selection at debates and in a way kind of combines these three into one perspective thinking that adaptation is what we are trying to explain and kind of then goes about thinking well

when we say that adaptation is for for the good of something what is that something and the answer to that then is the gene because genes then are conceptualized as these entities that are passed on intact from one generation to the next, whereas the kind of the main alternative organisms or sometimes groups don't have that evolutionary longevity required to play that role. An organism, by this reasoning, is a unique combination of its genotype and environment and their interaction is here in one generation gone in the next, whereas a gene then is passed on. So by this reasoning, the gene is ultimate beneficiary of any. adaptation.

And lastly, it does this, kind of takes this insight from population genetics, that evolution is this lineages of genes over generation. But because it's very much going to emerge out of the study of social behavior, the kind of group selection debate, it combines that with a form of kind of anthropomorphizing or gential thinking. I think if I was a gene, what would I do in this situation? And this is where it comes that we expect genes to behave selfishly and selfish, simply meaning here, trying to maximize their own.

representation in the next generation. And just to give credit words to, the idea obviously became famous and was developed by Richard Dawkins with the selfish gene, but it was essentially the idea itself was already there. Yeah, so I think most of us learned about the genes that we through the writings of Richard Dawkins, especially the selfish gene. Ten years earlier, the American George Williams published a book called Adaptation and Natural Selection, which I hold as one of the most important books in Evolution and Biology in the second half of the 20th century.

But this is a book primarily directed towards other biologists, on whom it did have a kind of profound influence. It really sharpened the discussion about adaptation and the study of adaptation. But it's, you know, it had the kind of fate that successful academics books have. It's read by the peers and but not so much more. Whereas, you know, there are a few terms in modern science that have had as in the reach that selfish genes has had. But as you say, many of those arguments are there already 10 years earlier, but they're laid out in an even more explicit and a much more forceful way in the selfish. It's a much better title. Come on.

Yeah, it's an incredible title. It matters, right? Except that we're still arguing about what selfish and what gene means. That still works. That's okay. The purpose of the title is to make people buy the book. So it's an incredibly successful title in that sense. Yeah, that you have to admit.

And just to be, just to clarify what it means to say the sort of, I like that you call it the, the genes, sorry, what did you call it? I've forgotten it. The genes idea. Right. So it's a little bit, you're lowering the temperature a little bit from the selfish gene way of putting it. But, I mean, one distinction may be worth drawing is that when we say gene here, And correct me if I'm wrong, we mean the instantiation of every single copy of that particular gene in all the individuals. That's the thing that is competing. It's not like the particular

stretch of DNA just in one cell or one individual. Exactly. Most of the time, that is indeed what we mean. They're kind of the, in a way, the type, the gene type that is present in this, all its copies in a population. And you think of that kind of whatever that is, is trying to maximize. to mytesone representation. It turns out that this way of thinking, this kind of genes eye view, is very effective at also thinking of situations where indeed the material part of a specific genes

are in conflict with other genes within a body, what is known as this sort of genomic conflicts in biology where kind of the fitness interests of different genes inside an organism diverge. And this is kind of one of those empirical fields that really was stimulated by the kind of, shift in perspective from organisms down to the gene level. Hey, everyone, it's Cal Penn.

I'm the host of Earsay, the Audible and I Heart Audio Book Club. This week on the podcast, I'm sitting down with Ray Porter, the narrator of Andy Weir's audiobook Project Hail Mary, massive sci-fi adventure about survival and science, and what happens when you wake up alone very far from Earth? I really had to make a decision because I caught myself getting that frog in my throat and starting to get teary as I'm narrating some of these sections. And it's like, okay, yo, yeah, yo, is this indulgent? And I really thought about it.

I was like, no, at this point, it would kind of be betraying the trust the author and the listener have in telling this story if I don't go through it. But there's places in this book that deeply emotionally affected me and I left it on the mic. That's great. Because it served the story. People will say like, oh my God, I cried at the end. It's like, yeah, dude, me too. Listen to Earsay, the Audible and IHeart Audio Book Club on the IHeart Radio app or wherever you get your podcasts. My best skin ever at 45?

Give me a theme song and a best skincare award because it feels like this. Right there. That's farmhouse fresh skin, all right? I'm blowing. And everyone asks how. The best skin care is Farmhouse Fresh, and the award is you, your best you. Visit Farmhousefreshskincare.com and use code radio for a free starter routine with any purchase. And one of the reasons why people got their hairs up on end a little bit,

and also one of the reasons why other people loved the idea, is because it does seem to anthropomorphize a little abstract notion, right? I mean, Richard Dawkins and no one else, believes that genes have attitudes or have selfishness or anything like that. So the claim is purported to be that given this mathematical description of population genetics and kin selection and inclusive fitness, it is as if the genes are being selfish. Exactly. I mean, and the as if here is crucial, especially because, I mean, selfish is, of course,

a very loaded term. I think, you know, Dawkins Spreps deserve some criticism. that the use of the word selfish, even in the book, slides a little bit what I mean, because on the one hand, you have selfishness in terms that all genes are expected to be selfish. And you can say that if something applies to everyone,

how useful is the term to begin with, because you also then use the same word to describe individual organisms behaving selfish. And you contrast that with them saying behaving altruistically or mutualistically. And then like something that is used slightly slightly differently. You kind of get it, but it also kind of slides somewhat. And it's not even to begin with the kind of psychological motivations of genes or organisms.

Where again, we use self-gages, just kind of daily language in a slightly, though, related kind of way. And is the genes-eye view or the selfish gene idea supposed to be exactly equivalent to the existing formalism, but just highlighting things in a conceptually more clear way? Does it actually make slightly different predictions? So most of the time, the idea is that it will be the same. So in the book that was followed up, followed the self machine, the extended phenotype, which kind of is the only book that Richard Dawkins wrote that was aimed at professional biologists.

All other books, as you know, has been for the public. But the extended phenotype are for professional biologists. And it comes with, it's fully referenced as you can expect a book like that to be. That thing said, it's a book that can be understood by anyone kind of willing to make the effort, but you get less for free. Yeah. That you compare to his other books. But there he starts with kind of the, this Nekker cube illusion that if you draw a cube on a paper with straight lines, you can often see from two perspective, which lines come first, so to speak. And he kind of would say that this in a way is the gene-centered versus the kind of inclusive fitness view of the world.

that there are kind of two ways of looking at the same thing. And in many ways, that is the kind of prevailing attitude, I would say. Early on in its selfish teen is written a bit more that like this is a kind of empirical rival to individual group level selection. But I'd say most of the time is considered that it's equivalent to inclusive fitness. And I do, there is an interesting kind of history about the relationship between the Deanside View and Inclusive Fitness. As far as I can tell, the very first time that the termed selfish genes appears is in the notes that Dawkins developed when he was a graduate student.

And his PhD advisor was away on sabbatical and asked him to step in and lecture on animal behavior. And it's in the mid-1960s. So the papers on Inclusive Fitness had just been published and Dawkins wanted to cover it. But these are tricky papers to get right. So he had tied up his notes. And then there, he kind of. explains the insight of inclusive fitness using this kind of gene-centric approach that we should expect genes to behave selfishly or trying to maximize their own. That being said, while Dawkins

has kind of viewed Hamilton as arguably the greatest Darwinians in Darwin, he's also sometimes thought that Hamilton is one of those people who never really finished their own revolution and never committed fully to the insight that you should just get rid of the organism and go down to the genic level. So the first answer is that yes, they're most of the time equivalent. I think the nuanced answer is that there is some kind of, I think, underappreciated tension between the gene-side view and inclusive fitness that sometimes, I think, ought to be explored more. Well, so I was going to ask, I was going to ask about how you could get new insights,

even if they were exactly identical. But maybe I'm not clear on how they're, what the tension is. You know, maybe there's rhetorical tension, but is there empirical tension? So I'll say the tension arises that the way we can, if you want to construct an inclusive fitness model, that is a kind of an individual organism property. And that often you have to kind of ignore things that happen inside of the organism that like that all the whole part of the organism share the same purpose. That has a unity of purpose. Whereas so I've done most of my empirical research has been on this example of genomic. conflicts, so situations where genes differ in their inner fitness interest, where that assumption

breaks down, that you don't have kind of all genes working for the same goal. You have certain genes working for their own transmission, even if that comes at expense of the thickness of the individual organism that cares it. And these are kind of has somewhat confusing in terms that they're not in a selfish genetic elements, which is, you know, strikingly close to selfish genes. Here, selfishness is then used in a much more explicit way that they are selfish with respect to other genes of the same genome or indeed the thickness of the individual organisms. I think I see. So the question is, who are the genes competing with? Is it other alleles?

I mean, the alleles that are competing. So they are competing with other alleles and other organisms? Or if really they just want to pass themselves on, who cares of other ones are also passing themselves on? And they might even fight against other genetic components of their own organism. Exactly. And this is truly a weird and wonderful world. And you have examples like, so you have things like the meotic drivers that. So normally you expect that meiosis, so the process of production of sperms and eggs,

is this kind of fair lottery, that you have two copies of a specific gene, like in humans, and that each of them has a 50% chance of being transmitted every time you produce a sex cell. But then you have these kinds of genes that, can distort that process. So in general, rather than ending up in 50% of the sex cells, they end up in 99% of them. And they can then become very common in a population,

even if by doing so, they reduce the number of offspring, in other words, fitness of the individual in which this kind of distortion happens. And you have almost any part of the mechanism by which you copy genes and put them into sex, that has been hijacked by some sort of self-genetic element to improve their chances of being transmitted.

And I think these kind of things are making, in many ways, make perfect sense from a gene center or a gene side view. It's often been considered the perfect, kind of empirical vindication of the approach. And I think this kind of, the tradition, in evolutionary biology that emphasizes the individual organism too much, which I think I put inclusive fitness in that category,

I often have to downplay the importance of this kind of phenomena too much for my liking. It's not that you can't use those approaches, but in a way it's kind of an issue of temperament, I think, or what observation you think are important in trying to explain. Well, it's interesting because I forget whether we said this explicitly, but one of the motivations or sort of paradigmatic examples in discussions of kin selection and inclusive fitness are ant colonies, right, where many individuals have no offspring, right? Depending on what kind of ant they are, you know, they work for the betterment of the colony. And the idea being that their genetic heritage is passed down by them serving their queen and their, you know, et cetera. even though they themselves don't have offspring.

And so this is kind of a twist on that idea. We're saying that a certain gene can benefit itself. Maybe it's the evil twin of that idea. A certain gene can benefit itself by shutting down some of the reproduction of reproductive capacities of an organism, but guaranteeing that its cells or its heritage will be passed on. Yeah, and at the heart, in some ways, you are faced with the same kind of conceptual question and that you have an allele that is harmful

in a way to the individual that carries it and you're trying to make sense of how can that ever evolve that kind of conundrum and like that happens when you have the evolution of sterility in your social insects and in the situation of these kind of genetic conflicts or self genetic elements that you have the spread of an lease that harmful to the individual carries it and so how that how can we explain that evolutionarily uh So to me, there's always been that kind of conceptual kinship between the two kinds of questions. And even if, to go back, okay, that was a very, very interesting example there. But even if the two views like Dawkins originally said are more or less equivalent,

it still can bring conceptual insight if you have a different way of formalizing the same set of ideas. And so by thinking of genes as the agents in some sense, you think of all biology is slightly different, right? I mean, there's this whole vocabulary of genes as replicators and organisms as just the vehicles. All the organisms are are buses to carry on genes from generation to generation. Yeah, in many ways, that is the central claim of the genes I view,

that evolution by natural selection requires two entities. It requires something to play the role of replicators, which is how information is passed on from one generation, to the next and then something that kind of packages that and transport is around and that is called been known as vehicles and this is typically a role typically played by by organisms but in principle it can also be played by things like like a cell like in a situation like cancer so that there you have kind of to break down or you used to have kind of think of it as one vehicle now we have some vehicles that have broken out on on their own competition and and in in in practice you can in principle it can also

be something like the whole group could be could be the vehicle but you're absolutely right that in a way like the choice of words here it's important that vehicle is kind of inherently a passive term relative to the replicator you kind of clear kind of what is valued in this viewpoint is the become a replicator and the organism in a way is downplayed and it kind of Dawkins was part of formalizing this but it's also the the philosopher of biology david hall who kind of came up with the same kind of way of formalizing the insight of the genes I view, but he preferred the term interactor over vehicle because you think that by calling it vehicle, you're downplaying too much of what's interesting about organisms.

So, so, but Dawkins famously said that he didn't coin, he coined the term vehicle not to praise it but to bury it that he really wanted kind of the field to move his focus down to some of my best friends are organisms. I do want to give them credit for, you know, for doing something. playing some role here. But this is probably out of left field, but that distinction kind of reminds me of touring machines a little bit. You know, the idea of some physical or even just, you know,

touring machines is probably being too highbrow about it. Computers, right, where you have hardware and software. I've always wondered going back to touring in his machines, where he has an instruction tape and then some little head that goes back and forth and does the job. how natural is this division of labor into sort of physical doer in the world versus instructions or information about how to do it? I mean, at the end of the day, it's all physical stuff. So where and why and how universal is this distinction?

It certainly seems central to the genes eye view. Yeah, and it's an interesting layer to that, that Dawkins himself seemed to be very taking. by computers. He was one of the first to learn how to program a computer when he was still an active researcher. In several of his book, it does emphasize that

genes or DNA as this kind of information, just like a CD or any kind of other storage. And then the organism is this machine that on which it just runs. There is a debate in philosophy biology to what extent the machine metaphor

is helpful or not and in the kind of thinking of organisms as machines it goes back a long way and I think historically has been quite helpful I'm I'm a little bit of two minds because I think there is something interesting about organisms because on the one hand of course there are physical object just like everything else as you say there are this subject to the same laws of physics like anything else and there's nothing kind of magical about it so there are physical object just like anything else at the same time organisms are physical objects unlike anything else, that they have this kind of like,

almost sometimes the, a sense of purpose and a goal directness in them. And this is kind of what's given rise to kind of biol, uneasy relation with teleology. And how much should we allow ourselves to talk in terms of purpose? And that is something that really runs quite deep in biology. You can vile up even the most senior and sober people on questions like this, whether this is an embarrassment that we talk.

in terms of purposes in biology, whereas others think that is almost impossible not to do it, especially when we study animals and animals' behaviors, that kind of attempts to try to use some sort of language that's meant to be neutral. It's not just unhelpful, it's downright silly. And press my attitude is somewhat that there has been the classic debate that did Darwin banished teleology from biology, from biology, or did he naturalize it?

Did he make it okay by kind of natural selection is a way that makes it okay? Because then you can talk about goals and purpose in terms of fitness. And in a way, kind of like, I think of the inclusive fitness situation, in a way coming from that naturalizing way, that like here you have kind of something that organ is ought to kind of appear as striving towards. So there's no magic here. There's no land with hell. There's nothing kind of spooky going on.

But you have kind of allow yourself then to use a kind of land. which otherwise isn't really okay in science. I mean, you don't use it in physical chemistry. Part of science that biology often looks towards. But biology, I think, straddles this study on the one end is a physical science, but on the hand it's a behavioral science, and it's awkwardly trying to balance the two. But it's certainly, I think, a part of science

where purpose enters into our explanation in one way or another. And I am on the view that we should just accept that, but try to do it in a formalized or try to be so kind of as formal about it as possible in the way we approach it. Yeah, good. I'm completely on your side. I have hopes to write, I've already written in various places

toward the side that says that purpose and teleology can be naturalized. Basically, I would say not just because of evolution, but because of the arrow of time, because of entropy increasing over time. But to do it, I say, it can be naturalized. I didn't say that I did it,

or we have a formal theory of it, and I think that that's something that would be very interesting to understand better than we do. Yeah, so a colleague of mine, Manus Pat and I, just submitted a paper where we try to defend

what we call license anthropomorphicizing. Oh, good. Which is a concept that you're allowed to anthropomorphize and say, if I was a gene, I would do so and so. Yeah. But then you need to back it up

with a model, a mathematical model, and that is what provides the license. And it kind of just goes back to the inside this basic point that is almost impossible to do biology without using this. But we should try to do it then in an organized or licensed kind of way. And that we argue is kind of things like mathematical population genetics or other part of mathematical biology provides that license to do so. I would say personally the same exact things about free will. But let's not get into that debate right now. That's for another discussion.

Then there's that. Let me give you one more chance, just in case we've missed anything, to give good consequences of being a genes-eye view person or selfish gene person. Are there other predictions that this viewpoint leads us to or other perspectives that are helpful to biology? So I think that the genes-eye view is a very powerful thinking tool in biology. I think it's one of the best tools we have to make sense of this messy world. But like with all tools, to get the most out of it, you must understand where it came from and what it was what is trying to achieve. So it's an approach to then to try to understand adaptations and kind of the logic of natural

selections that can give rise to that. So it works especially well when those adaptations that we are kind of counterintuitive to the individual organism's point of view, which I have. that's been the traditional focus of biologists. So we talked a little bit about genomic conflicts, as I think it's a great example of where the benefits of the genes of you comes through. The others in the study of social behavior to understand the work of sterile team. One that was kind of like a new thing that people handle is the notion of extended phenotypes.

So these are defined as phenotypes that are located outside of the body in which the genes are kind of caught responsible for, that phenotype reside. So, kind of typical example has been the things that animals build, like the beaver's dam or the nests of birds. So these kind of structures that clearly has a genetic component to them. So in a way, should be thought of as a phenotype. But that the phenotype is not part of the organism. So do my, wait, do the clothes I wear count as my extended phenotype? So the synodemines are typically considered to be, to be part that has some influence on the fitness of the organism in a consistent way across time and population. I think it's pretty clear that how well people dress has an impact on their reproductive success.

It do, though, humans are famously an abherent species to study. Fair enough. I'm just going to believe this on purely theoretical grounds. Don't worry. Yeah, yeah. But those going to build structures of animals. I think that's kind of like thinking of those in turn from a genie point of view. I think helped stimulate that.

Also, kind of individuals where individuals of one species can manipulate the behavior of another, so kind of this gruesome example like zombie ants, where ants who are infected by a particular kind of fungi completely changed their behavior from kind of shying away from parts of the planet where they live to go, where it's beneficial for the fungi to live and eventually kind of end up behaving completely bonkers because they've been infected by this fungi. And here, so that again, there's a behavior of the ant that has a genetic component, but the genes of for that, if you will, if you allow us off that short hand,

for that behavior is not located in the ant, it's located in another organism. And that again is something that this kind of extended phenotypes are, I think, easier to think about from a genic point of view than trying to shoehorn it into that of the individual organism. So except let me bring up that not everyone agrees, right? Like this is why you wrote a book because there are skeptics and critics of the genes eye view point of view. And at least, I mean, hopefully you'll fill me in on what all of the critics are,

the most important critics are, but one of them is that even the modern synthesis is no longer how we should be thinking about evolution, right? I mean, there's horizontal gene transfer, there's epigenesis, factors, the whole idea that there's just a simple tree and things branch off and never cross-pollinate has become a lot messier in recent years. There's mitochondrial DNA as well as nuclear DNA. So is there any legitimacy to this idea that the genes-eye view is just a codification of a very old-fashioned way of thinking about evolution? The genes-eye view certainly shares a lot with the modern synthesis, especially kind of ideas of that adaptation and natural selection are

importance and kind of framing that in terms of genes. But there are a couple of things I think that are worth mentioning here. So one that the modern synthesis is perhaps more or some more diverse than what critics of it gave it credit for. There's often presented as this very kind of strict dogmatic viewpoint, which it wasn't included many different kinds of people and many different kinds of views. Second of all, criticism of it has existed pretty much as soon as it was kind of codified or kind of as it became expressed that often of the nature, I think, of people not really thinking that what they studied gutted is the respect that it deserved. And this has come different kind of versions of this over time.

I think it is interesting both Richard Dawkins and George Williams, so the kind of the two architects of the Dean's Review, did emphasize that what they were doing. it was to articulate the kind of orthodox view in a new way, that they kind of very much align themselves with that kind of traditional approach. So the larger criticism of the modern synthesis, and the genevieve, often is taken to be kind of the poster child of that, is that biology has changed so much that this all way of thinking about is no longer helpful. And yeah, this is a contentious topic. And I think many sensible people can disagree on it.

I don't tend to think, tend to view this new observation as being lethal to the modern synthesis anyway. I think it's modern synthesis is also quite a, it's a flexible framework. And evolution by all this is not as dogmatic as even its internal critics would like to present it. That's been quite, so that's like to incorporating it. We are probably putting too much emphasis on certain things and not enough emphasis on other things. perhaps the kind of the greatest challenge, I think, to the genes I view is this notions of that we ought to consider a more inclusive notion of inheritance that we are giving genes too big of a role in that that parents pass on more to their offspring than their genes. So this can be things like epigenic signals, methylation patterns, but also kind of cultural, forms of cultural inheritance, some other kinds of maternal effects. So that I think there may well be some truth too.

One thing that I would call it like to see, I don't know if anyone is done, is that the reason why the genes I view proponent likes to talk about replicators rather than genes is that replicators is completely divorced, divorced from any sort of material basis, and that what really matters is that it can make copies of itself. So I wonder how some of these kind of other forms of inheritance can that be incorporated into this kind of traditional framework. framework. I mean, certainly Richard Dawkins is in favor of thinking about memes as an important method of cultural transmission. Yeah, I mean, memes also, I think is interesting. Of course, it was coined in the last chapter of the first edition of the selfish gene. It was meant to kind of free replicator from the kind of association with genes in a way. And that was the idea that if organic evolution requires something to play a replicator and a vehicle, so should, should cultural evolution.

Ironically, though, I think that despite wanting to free itself from the material basis of genes, it is actually exactly those kinds of properties that makes the genes have you worked so well in organic evolution, but not in cultural evolution. That thinks that where does one gene start and the other one ends? How long is a gene? What does mutation actually look like or competition between alleles, what are really? a lot of those things that in organic evolution, we try to downplay some of those kind of properties,

but in a way, that is kind of exactly why it worked at the end of the day in organic evolution, but not in cultural evolution, where those boundaries become so poor is that I think to lose its value as a concept. Yeah, a real biological gene is something at the end of the day you can hope to point to there, the DNA. Yeah, it's anchored in some sort of material reality

in a way that I think is hard to do in memes. So despite being a very successful meme, of itself, my impression is that it's influencing current study of cultural, it's somewhat limited. That makes sense. I guess, and this is a good place to sort of wrap up, the

part of the discussion that is most fascinating to me, it's taken us a while to get here, but it's the level of selection, levels of description discussion, right? One thing that one can say, it's not a very useful thing to say, but one can say it is

if I could be Laplace's demon, If I knew the exact state of everything in the universe, and I knew all laws of physics, and I had infinite calculational capacity, I could just tell you what was going to happen. And I think an underappreciated aspect of that is that Laplace's demon doesn't need to know about cells and genes and organisms, right? It just needs to know about the microscopic state of the world. But I can talk about things in terms of cells and genes or organisms or societies, etc. is there how I mean this is a far too big a question to answer but say what inspires you to say what is the right way to think about the different levels of description that we have to capture biological reality and can they coexist or is there a best one that we should focus in on

I think I've very much come on the view these days that there isn't one best way to do it in biology. I think that biology is a kind of science that is so messy, and especially the part of biology, like evolution, where we're trying to start a historical process, that we are better of trying to retain multiple perspectives. And that it's quite comfortable that some people prefer to model things in a group section way and some in inclusive fitness kind of way

and sometimes in a genetic kind of way. And that it's not necessarily going to be a true answer. Well, there may be one. I don't know it's meaningful, necessary to spend all our time trying to find that. That being said, I think we ought to spend some time worrying about when we have two different kinds of approaches. So one example is perhaps in population genetics and inclusive fitness methodology. So inclusive fitness, you've often relied on these kind of optimization methods, quite popular in the study of animal behavior. But optimization is also something that population genesis is rejected in the 1960s, that we should never expect selecting to optimize anything.

that is the thing is going to be in this kind of may reach an equilibrium, but they're not going to be kind of optimized in any sort of meaningful way. So some part of our field has rejected it and some uses constantly. And in some ways, I'm fine with that.

At some time, I think we ought to spend some time worrying about it. But kind of on the larger philosophical point, I think that we have, you know, I think you have one reality, but you have many ways of talking about it. And I think becomes expressed very well on a day-to-day level in this study of biology.

Well, I guess the complication or the interestingness of the question comes in because, well, for me personally, I don't know about for the rest of the world, but as someone who is trained on pretty cut and dried examples of different levels of descriptions, such as atoms and fluids, right? We can literally derive fluid mechanics from large collections of atoms interacting with each and we know that you don't need to understand atoms to understand fluids or vice versa. So there's this almost too good to be true simplicity of autonomy of the different levels. And I get the feeling I'm sort of slowly dawning to the realization that in biology and in the

human sciences, that those levels are harder to disentangle from each other. I think that is exactly right. that you have these kind of layers of descriptions that both can work really quite well. So kind of the quantitative genetics that animal breeders use, where you're kind of selecting on phenotypes and you kind of know that there is heritability because you can compare parents and offspring, but they kind of quite precise quantitative methods that you use there don't really care about DNA sequences. They can be informed by those.

And I think, like, you know, with the advance of what, you know, sequencing and all that, part has also been improved, but it functions rather well without it as well. But it's not that, so you can kind of derive those kind of phenotypic approaches from studying genetic sequences or vice versa. It's not necessarily going to happen, I think. And I don't really feel that pessimistic saying that. I think that's just kind of the nature of the endeavor. So absolutely, I think biology is kind of perhaps where that kind of, that you can derive one from that is starting to really, break down.

Well, I think, but I think, I certainly agree with that 100%, but there's sort of like a deeper, trickier issue. I mean, I think that far too much emphasis has been put on derivability, because outside the very simplest examples, of course you can't derive. I cannot derive the solidity of the table in front of me from the standard model particle physics, but I don't think it's incompatible with that, right? But what is important, I think, or not just what is important, what I'm interested in at the moment is, I don't need to know about the standard model of particle physics to describe the

behavior of the table. There is an autonomous theory of it. And in biology, forgetting about derivability, it's not even clear to me that there's that, right? Yeah, no, as an interesting point, I don't thought about it in that way. But yes, I think there is less clear that you have that kind of asymmetry. Yeah. No, I think that it's absolutely true, that you can kind of go about it. I mean, everything has to be compatible. Of course, it was going on at the lower level. But like how you begin to kind of worry about that, I think is less clear. Well, I mean, I have an example in mind, which is very contemporary, which is the germ theory of disease, right?

So, you know, we can have a theory of how diseases are transmitted through breathing and contact and things like that without actually knowing that they're microbes, right? We don't need that lower level theory, but it is helpful. And so I honestly don't know the answer to this question, but I would like to know is there some sense in which there is a self-contained theory of, you know, let's say disease transmission that never refers to microbes? Or if there is such a thing, is it so baroque and backwards that you should always, you can always improve your description by just helping yourself to the microbial description. Yeah, I guess part of the answer may lie in that like epidemiological models are in some ways very generic in the sense that you rely on like it just transmitted from one individual to the next. But they can certainly be improved by knowing how it's transmitted. So like learning that the COVID-19 virus is primarily airborne rather than like taking around on surfaces improves models, I think, in how you want to, that it become more exact. But you can certainly, it's not necessarily, you can do sound models, regardless of how that,

how they kind of transmission, what is most important for transmission of this particular virus. But it's kind of where one part where the biology of it matters, or the kind of the lived reality of it. Yeah, I do think these are these are good questions. We don't know the answers to yet. So, but I'll give you one last chance, one final question. What is all this bode for the future? evolutionary biology. Like it's interesting to me as an outsider to see the flare-ups,

not just intellectual but emotional, when things like the Novak et al paper came out about group selection or just the discussion around the idea of the selfish gene, you know, gets pretty heated. You've mentioned a couple times that data has sort of stepped in and made things a little calmer. Is that the future, or are we still in for, you know, a whole bunch of heated debates in the pages of the New review books. I think data certainly has helped a lot. But there is also this issue that in evolutionary biology, that we tend to have these flare-ups over time.

There was a fun paper probably a couple of years and simply called what is wrong with evolutionary biology. And the argument there is that they think they're kind of two things that evolution by national selection on the one, and it's such an abstract principle that it can kind of be applied to almost anything. But also the evolutionary biology is such a diverse feel in terms of what we study. So if you can't convince the paleontologists, maybe you can get the molecular genesis to listen to you.

If they don't, maybe you can try the plant ecologists. And those two things in combination mean that you can always get a small enough audience to keep it going. But you also may have a hard time explaining, like kind of convincing everyone. Kind of the chief thing that I hope to achieve with this book is to bring some nuance to the debate over the gene side view. that I noticed that one, that people kept having these debates, but also when I was a graduate student, I would ask more senior people in my department about it. You walk to one person and they will tell you,

oh yeah, this is the correct way to think about evolution. And everyone knows that. And then you walk a few doors down and he said, oh, no, this was debunked in the 70s and everyone knows that. And there was something there that I kind of was frustrated both by the proponents and the critics that I often thought that they painted, kind of an unfair picture of what the debate actually is, the strength of the case before and against this way of thinking about the evolution.

And so that's kind of what I wanted to capture really with the book. Yeah, no, I think it's great. And I think this was extremely helpful conversation for me in clarifying things. So maybe whatever debates we have in the future, they'll be at least on a more informed basis going forward. So Arvid Ogren, thanks so much for being on the Mindscape podcast. Thank you very much for having me.