Ep 115: Does a porpoise have a purpose? Agency and goals in evolution (with Samir Okasha)

What is an agent, and does an organism have to be conscious to be one? How does organismal agency affect evolution?

In this episode, we talk with Samir Okasha, a Professor of Philosophy of Science at the University of Bristol. Samir studies fundamental philosophical questions in evolutionary biology, most notably how selection acts on various levels of biological organization. Our discussion focuses on his book “Agents and Goals in Evolution,” in which he unpacks various definitions of agency and outlines their evolutionary implications. We talk about whether genes and groups of individuals can be agents, whether agency is heritable, where variation in agency comes from, and the relationship between agency and adaptation.


Cover photo: Keating Shahmehri

  • Art Woods 0:00

    Hey Big Biology listeners a reminder that we're moving into a more intensive period of fundraising over the next few months. So you'll notice a couple of changes to our show.

    Cameron Ghalambor 0:08

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    Marty Martin 0:14

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    Art Woods 0:57

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    Marty Martin 1:02

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    Cameron Ghalambor 1:09

    Now, onto the show.

    Cameron Ghalambor 1:22

    Marty and Art it's good to see you two philosophers today.

    Art Woods 1:26

    Philosophers? Norwegian winters got you confused, Cam? We're biologists!

    Cameron Ghalambor 1:31

    Wait, but Ph. D stands for Doctor of Philosophy.

    Marty Martin 1:35

    Technically, Cam is right. We are philosophers in the sense that all sciences are rooted historically in philosophy. And that meant the pursuit of wisdom, scholarship and expertise.

    Art Woods 1:46

    Fail!

    Cameron Ghalambor 1:46

    But there is a discipline where the two meet. The philosophy of biology explores fundamental topics like how the meaning of terms and critical assessments of assumptions have changed through time.

    Marty Martin 1:57

    And if you're a regular Big Biology listener, you've probably heard past episodes with philosophers of biology, like Dan Nicholson, Dennis Walsh, and many others.

    Cameron Ghalambor 2:06

    I think most practicing biologists, myself included, rarely read the philosophy of biology literature, but those are some of my favorite episodes because they've made me stop and reflect on ideas and concepts that I often use, but never really thought deeply about their meanings.

    Art Woods 2:21

    In a 2019 article in the Proceedings of the National Academy of Sciences Lucie Laplane and colleagues argued that philosophy plays a critical role in the sciences

    Marty Martin 2:30

    An example they highlight is a critique of the self versus non self idea in immunology. Philosophers pointed out that the idea of self breaks down when we recognize that all organisms are made up of communities of non-self symbionts. And the shift has real life implications for developing medical treatments. Thank you, philosophers!

    Cameron Ghalambor 2:47

    One active area in the philosophy of biology is evolution by natural selection. Concepts like fitness, selection, adaptation, contingency and determinism pop up all the time. But talk to philosophers of biology and you find out that their meetings are far less clear than you would think.

    Art Woods 3:05

    For example, think about the phrase "survival of the fittest." When we define fitness as the ability to survive and reproduce, the phrase becomes tautological because we're saying essentially "survival of those who survive." This is why the philosopher Elliot Sober has said we need a better definition of fitness.

    Marty Martin 3:22

    Another topic we've covered a lot recently on the show, and that also shows up in a lot of philosophical analyses is agency. Agency has been defined very differently inside and outside of biology.

    Cameron Ghalambor 3:35

    Agency also carries a lot of baggage which puts a lot of biologists on edge. When you say that organisms have purposes and goals, it opens the door to conscious intention, and even supernatural effects and forces.

    Art Woods 3:48

    Yet, even if we recognize that most of what organisms do to survive and reproduce is done unconsciously. We often use casual and sometimes clumsy language.

    Marty Martin 3:58

    Think about the cuckoo, a bird that lays its eggs in the nest of other species, the cuckoo egg typically hatches first, and the cuckoo chick then pushes the host's eggs out of the nest.

    Art Woods 4:06

    We might casually say that the cuckoo chick does this because it consciously wants all the food for itself.

    Cameron Ghalambor 4:12

    We often use this kind of language as shorthand to help us explain why the chick behaves the way it does, even though it may not be making a conscious choice.

    Art Woods 4:21

    A more traditional explanation would say that this behavior evolved by natural selection acting on heritable variation in the context of an evolutionary arms race with its host.

    Marty Martin 4:31

    So while there is a growing interest in the notion of agency in evolutionary biology, there's still a lot of confusion and debate, even among us about what it means, as well as its relationship to fitness selection and adaptation.

    Cameron Ghalambor 4:44

    Our guest today is Samir Okasha, who's a Professor of Philosophy of Science at the University of Bristol in the UK, his 2006 book, Evolution and the Levels of Selection, won the Lakatos award. Today we talk with Samir about his 2018 book entitled Agents and Goals in Evolution.

    Art Woods 5:01

    In contrast to our past discussions of agency, Samir takes a different approach. He asks, in the book, quote, "does thinking about organisms as agents have a genuine scientific role to play in our understanding of adaptive evolution? Or is it simply another application of anthropomorphic language?"

    Marty Martin 5:17

    In our chat, we look critically at the different definitions of agency in biology. How is it different from phenotypic plasticity? And do we separate agency and fitness?

    Art Woods 5:27

    And what about agency below the organismal level? Could a gene be considered an agent if it acts selfishly to increase copies of itself like in cases of intergenomic conflict? And what about groups of humans or even groups of different species that exhibit cooperative, goal-directed behavior to groups have agency distinct from individual members?

    Cameron Ghalambor 5:46

    Samir argues that although we gain insight by anthropomorphizing some adaptive traits, there's still value in understanding how agency influences evolution, even if we don't know about consciousness in that organism.

    Marty Martin 5:58

    We also discuss whether the process of evolution by natural selection represents a form of agency. This might sound odd, but recall that Darwin himself depicted natural selection with the language of intent, as a background process that's always scrutinizing populations, keeping the good and eliminating the bad.

    Art Woods 6:15

    We debate some really difficult problems like whether agency can evolve? Is there heritable variation for it? And how would we measure that, something we've been thinking about a lot, but I'm not sure we've totally resolved.

    Marty Martin 6:26

    So stay tuned for a philosophical assessment of agents' goals, and evolution.

    Cameron Ghalambor 6:31

    I'm Cameron Ghalambor

    Art Woods 6:32

    I'm Art woods,

    Marty Martin 6:33

    And I'm Marty Martin, and you're listening to Big Biology.

    Cameron Ghalambor 6:51

    Samira Okasha, thanks so much for joining us on Big Biology today.

    Samir Okasha 6:54

    It's my pleasure.

    Cameron Ghalambor 6:55

    We're super excited to talk to you today about your recent book Agents and Goals in Evolution. Maybe to begin, could you start off by defining what is a biological agent?

    Samir Okasha 7:07

    Wow, that's a complicated and difficult matter. Let me give you my take on it. As I see it, the place we need to start is by saying, asking pretty much the question you did I mean, what do we mean by the term agent? And why do people think that this is a good concept or term to use when thinking about biology? And I think that's an important question to ask, because, I mean, there's this flurry of interest in the notion of agency and biology's many of your past contributors have discussed. And as you'll be aware, a lot of philosophers have joined this discussion too. But on the face of it, I mean, that's a bit odd, because if you open any biology textbook, in any branch of biology, I very much doubt that the term agent is going to appear in the index. So that immediately raises the question, well, why should this notion be thought important for sort of philosophical reflections about biology, if the science itself doesn't seem to have any use for it? So that's, I think, the first sort of skeptical question I would ask.

    Samir Okasha 8:07

    The next move I would make is then to say, well, if people do want to talk about this, let's do the sort of standard thing we do in philosophy of saying, but what exactly do you mean? It's always a good question to ask. And as I see it there, I mean, there are a number of different motions of agent at work, both in different sciences and in Vernacular English. So in my book, I distinguish between four different notions of agent as a sort of prelude to the discussion. The first is what in traditional philosophy, we would call an intentional agent, someone or thing that engages in reasoning, and in thinking, and in deliberating about what to do, and chooses courses of actions, because of particular goals that they want to achieve, where the goal is something that's mentally represented by the agent. So that means that at the very least, you've got to have a mind of some sort. Now, what exactly that I mean, that's clearly intended to include humans, whether it includes what else it includes, you know, whether it includes all vertebrates, or even some invertebrates, or even possibly microbes, it could be debated. But that, as I see, it is the sort of traditional philosophical notion of agent, an agent engages not just in behavior, but in action, where action means that they're deliberately trying to achieve a goal. And that's what I think of as the most restrictive notion of agents, the literal applicability of which is the least if you see what I mean.

    Samir Okasha 8:24

    But then in different sciences, we find broader notions of agents. So for example, in AI, one finds frequent reference to talk about intelligent agents, where that basically means anything, it could be a control system or piece of software, or a thermostat that doesn't always do the same thing, that does different things depending on the circumstance that it's in. So typically will sense the environment somehow, and change its behavior depending on what it senses I mean, so implements stimulus response conditionals. And it's an extremely inclusive notion of agent that, of course.

    Samir Okasha 10:13

    Similarly, but differently, one finds in the economics literature, talk about rational agents and a lot of talk about the rational agent model, where essentially, that means an agent who engages in consistent choice, so behaves as if they're trying to maximize, you know, some utility function or something like that, as whose choice or behavior is consistent engages satisfies conditions such as transitivity, things like that. Again, that's a really broad notion of agency of agent in that there are no psychological requirements. And then any organism that engages in choice, or behavior of any sort, could say, satisfy or fail to satisfy those conditions on being a rational agent, as understood in Rational Choice Theory in economics.

    Samir Okasha 11:05

    And then the final notion of agent that I operate with, or that some people operate with, that I distinguish, is what I have called a minimal agent. And that just means something that does something. So an agent in this sense, is just something that does something rather than something that to which things happen, if you like. And how exactly you cash that out, I think is a difficult philosophical question. But I mean, examples can make it clear. So to borrow one from the philosopher Fred Dretske that I like to use. And he said, "Look, contrast these two things, a rat moves its paw, in order to press a lever to get a pellet of food. And an experimental scientist puts their hand into the cage and forcibly moves the rat's paw." In the first case, the rat moved, it's poor, in the second case, it's poor was moved. And that he said that he says, illustrates the difference between doing something and having something happen to one.

    Art Woods 12:04

    Yeah it's sort of active versus passive.

    Samir Okasha 12:06

    Yeah, exactly. And then obviously, the next thing to say about that is, well, what how do we cash that out? Precisely, what does that really mean? And some philosophers have thought that the answer is simply lies in the question of whether the cause was internal to the agent or not, you know. So if a bird flies, then that's one thing, if a hurricane force wind blows the bird, that's another thing. In one case, the cause of the is the, you know, the motor output is internal. In the second case, there's an external cause. So that's what I call the most minimal notion of agents entirely. So those are the four notions I operate within some sort of rough hierarchy, the most demanding the intentional agent, the least demanding the minimal agent, and then the intelligent agent and the rational agent, somewhere in between those two extremes.

    Samir Okasha 12:57

    And then the next question to ask is, well, which event if any of those notions applies to biological systems? To which biological systems do they apply, and how do we know, and why? But here, a complication comes in because I maintain and this is my, I guess my real interest in my book, that much talk of agency in the biological realm is metaphorical rather than literal, particularly in evolutionary thinking. I claim that there's a lot of appeal to the intentional notion of agent, the most demanding of all, that I think of is in many ways the core notion where the agent must have a mental representation of the goal. But the concept is only applied metaphorically, when we say things like the bacterium swims up an oxygen gradient, because it wants to get to an area of higher oxygen concentration or something like that.

    Art Woods 13:48

    Yeah. So let's unpack that person a little bit, because they had a question lined up about that, that very thing. So you know, in the book, you make it clear that for this more restrictive idea of agency that it requires a mind and mental representation. And at least, like among those of us, the hosts, that are talking about agency, Marty, and I sort of kicked this back and forth, I think our conception of agency is a little bit broader. And it encompasses basically any complex system that's able to receive information, process it in some way, and make a decision. And that doesn't evoke anything about nervous systems or minds or mental representations. You certainly could have biochemical networks that are making decisions you could make. You have sort of genetic regulatory networks that are making decisions. And so I think that that immediately expands this idea of agency down to very simple organisms like an E. coli deciding whether or not to make the machinery to metabolize lactose or not. So in your conception, why do you have to have a brain and a mental map?

    Samir Okasha 14:55

    Well, I don't say that you do. I only say that you do in order to be a literal agent in one sense of the term agent, but I entirely hear your line of argument. And I think it's one I'm sympathetic to that says, look, there are, you know, other, what you might call hallmarks of agency have found in pretty much all complex systems, biological and maybe even non biological, certainly including cells and possibly sub cellular constituents and molecular pathways and things too. However, I would say that that involves a somewhat different notion of agent. So I would actually say that's closer to the Intelligent Agent notion of AI. So basically, what we mean is that we've got something that receives information from the environment and modifies its behavior, as a consequence. So in short, it doesn't always do the same thing. What it does depends on circumstance, if you like. But then I mean, then, if that's right, then it raises the question, do we just mean behavioral plasticity? When we talk about agency in that sense.

    Marty Martin 16:06

    Exactly what I wanted to ask. Is this a continuous grade from intelligent to intentional? I mean, is there some way to find a clear demarcation between these kinds of things?

    Samir Okasha 16:22

    Well, presumably, I mean, the intentional one is a relatively late arriver on the evolutionary scene, in that, you know, however, widely we think your true intentional agency is found in the animal world and, you know, cognitive ethologists have have, you know, often debated, for example, whether corvids are capable of genuine intentional behavior or not, and opinions divide on that issue. But what's very, very clear is that, well, I would say, at least the vast majority of organisms, including all invertebrates, are not literally intentional agents. I mean, if we, we might describe them as if they were, when we say, you know, a cell wants to achieve such and such an end. But that's, that's to deliberately anthropomorphize, and maybe for good reason, but it's not literally true. It's a bit like saying this, the suitcase doesn't want to close, you know, it's might be useful thing to say, it's not literally true, right.

    Marty Martin 17:17

    What do you think Lars Chittka might say about that? I'm not sure if you're familiar with his work, but we spoke to him quite a while ago. So he's very interested in insect cognition. And he's done really amazing work on all sorts of bees in terms of their capacity for solving puzzles. One of the really neat stories that he tells is, you know, how, how do bees decide, "decide" to build the nest that they do, and the configurations that they can take? You know, it's not like it's a sort of hardwired, always the same kind of nest, it's very responsive to the conditions of the environment. And he's done incredibly sophisticated experiments to show, you know, the amazingly elaborate kinds of outcomes from, you know, presumably a simple cognitive system. Yeah,

    Samir Okasha 18:02

    Yeah no, I mean, I think it's a genuine challenge that and I mean, at some level, it's got to be an empirical question, I think. One way to look at it would be to say, well, you know, what, what's to be gained? This is a formulation inspired by Daniel Dennett, where he says, something like "heuristically, what would we gain, if anything, by treating, you know, some organism or collection of organisms in a, in the B case, perhaps, as an intentional agent, as if it really was trying to achieve some goal and have beliefs about the world? And was processing information? And so on?" I mean, is that is that a useful perspective or not? And I mean, the answer might very well be yes, in that sort of example, but I take it that there are examples where it would not, where we have a purely mechanistic understanding of, you know, the movement of some microbe or something like that, or the growth pathway of some plant, for example. I mean, it would seem superfluous to give a psychological explanation of plant growth, you know, a plant grows towards the light or something, then I mean, we have a, we have a fairly good mechanistic understanding of, that why the plant responds to light that way.And if someone wanted to say, "Well, no, it's because it wants to get more sunlight or something, or wants to grow taller than its neighbor," you know, that that would not be, I'd take it, scientifically useful. So one test is just the scientific utility.

    Samir Okasha 19:34

    Now, some, some philosophers and others would oppose that methodologically by saying, look, the question of whether something is useful is distinct from the question of whether it's true, and that may be, but nonetheless, if I think if it isn't useful at all, then that's some indication that it's not literally true. So as I see it, I mean, I wouldn't at all insist that we know that it could only be in vertebrates that we find the hallmarks of true intentional agency. I mean, another point, though, to bear in mind is that, I mean, I started when I defined intentional agency by saying, look, there really is a mental representation of the goal. The organism, really, it's not just as if it was trying to achieve a goal, it really is. But you see, if you talk instead about decision, then that's perhaps a less psychologically demanding notion than having a mental representation of the goal. In that I think it's perfectly sensible and plausible to say, you know, even of a developing organism that it needs to decide which of which pathway to go down,

    Art Woods 20:34

    So developmental regulatory networks can make decisions, biochemical networks can make decisions. Yeah.

    Samir Okasha 20:42

    Yeah, I mean, but, you know, some people might insist that the literal meaning of the word "decision" is a conscious decision made by humans.

    Art Woods 20:50

    Yeah it's sort of inherently mental, yeah, right.

    Samir Okasha 20:52

    Yeah. But I don't see that I don't see that that's either the standard way that the word is used in science these days, or has anything particular to recommend it sort of theoretically. So I'm quite happy for decisions to be a pretty broad thing. I'm very happy with the idea then that, you know, all organisms, including microbes, make decisions, and maybe decisions are made at the level of regulatory networks, and, you know, subcellular levels too. But if so, then I would say that decision making and having a mental representation of a goal come apart, in that the former is taxonomically widespread, the latter isn't.

    Cameron Ghalambor 21:35

    So if we can circle back a little bit to the concept of plasticity and flexibility. For, I'd say most practicing evolutionary biologists, the capacity for being flexible, and whether it's at the level of development in response to some environmental cues, or in the context of behavior changing, say, in the presence and absence of a predator, a lot of those types of responses are captured currently under the sort of framework of phenotypic plasticity. And so just based on your description, it seems that being plastic and flexible is inherent to being agential in your response.

    Samir Okasha 22:25

    Yeah, I would agree in all of the senses of agency, in all of the four senses that I've operated, I've distinguished

    Cameron Ghalambor 22:31

    So are they so intertwined, that they are essentially capturing the same phenomenon? Or, or can we sort of decouple them from one another?

    Samir Okasha 22:39

    Yeah, no, it's I mean, it's tricky. I mean, I do think that, you know, the notion of phenotypic plasticity and behavioral plasticity, that being a subset of phenotypic plasticity, you know, given the novel phenotypes of behaviors, you know, captures a lot of what people who talk about agency are trying to say, and probably in a more sort of acceptable way to many practicing scientists. And I think that's exactly the source of some of the opposition to the language of agency that one finds these days where people say, look, you're just using this mystical sounding term, which the cash value of which is something that we could say anyway, right. Now, you see, I think that's a valid criticism of some projects that involve the notion of agency but not of mine.

    Marty Martin 23:30

    Touche

    Samir Okasha 23:31

    Some theorists, including my some of my colleagues, in philosophy of biology, seem to have convinced themselves that there's a deep and interesting sense in which some organisms or maybe all organisms, and maybe other biological entities too are agents, and their task is to figure out what that sense is, subject to the constraint that it had better be interesting and better not just reduce to behavioral plasticity. But that seems to me an odd way to proceed. So that's why I'm one of the people who's somewhat skeptical about the notion despite having worked on it myself.

    Samir Okasha 24:43

    But my project is a somewhat different one, in that. I was in writing my book specifically interested in a mode of thinking and speaking that is peculiar to evolutionary biology, I think, which involves deliberately anthropomorphizing for the purposes of tree of achieving evolutionary insight. And one example of which is that the practice of, you know, taking the adaptive function or rationale of some phenotype, but particularly a behavior, but not necessarily, in the sense that the reason why it's evolved, and then treating that as if it was the organism's own goal, a literal goal mentally represented, even when it really isn't. And I give many examples in my book of this practice, actually at work in evolutionary biology manifests by the use of intentional language. So when we say things like that the female rat kills its injured young, because it knows that they don't want to, they won't survive, it doesn't want to waste resources on for example. There we explicitly employ, you know, vocabulary that has its natural home in relation to conscious, deliberate human behavior to by way of giving an evolutionary explanation, or an adaptive explanation of infanticidal behavior of the female rat. And so that was the phenomenon I was specifically interested in the use of intentional vocabulary in an evolutionary context. And so I took my project, I mean, I approached it very much in the spirit of a kind of anthropologist of science. You know, my starting point was, look, there is this pre-existing practice out there in the scientific literature, that's really unusual and philosophically puzzling. And what on earth could be the point of talking that way, given that it seems to willfully invite a confusion of proximate and ultimate? And given that it seems to just be a form of anthropomorphism? Could it really be doing any scientific work? That was the question I started from. And I think of that as a somewhat different project, indeed, an orthogonal project, to the one that many people, you know, both biology and philosophy, who talk about agency are ultimately are engaged in.

    Marty Martin 27:01

    So I want to, I want to push back a little bit because you made a great case, and this adaptationist program, it has been in is pervasive in biology. But there's this particular word that I hear you using that I'd like you to address, and it's mental, right. So an important qualifier of agency that you write about is integration. So it's not just the, you know, the potential to perceive the problem or to sort of think about the problem, but then the system does something about it. So can you expand the definition of mental? And more importantly, it seems to me to be leaning on mental as the qualification of a general and, you know, justification of how to think about that, that seems akin to the assuming that, you know, anthropomorphizing and adaptationist perspective, right. It's a move that allows you to do other things, but I don't know, I guess I'm just not on board, that mental is something special, because you know, you have to have integration. And then as Art was saying a minute ago, there's a lot of different systems and even elements of systems that integrate.

    Samir Okasha 28:13

    Yeah, no, I mean, I agree that integration and mentality are not not exactly the same thing, certainly. So I mean, what I'm getting at when I use the term mental is that just the assumption that, you know, psychological descriptors, like believes, wants, tries, apply, literally in the human case, and quite possibly in in other non human cases, too. But don't don't apply literally, in all cases where there are sort of hallmarks of agency in the sense of, flexible behavior, plasticity, decision making, information integration, all of that. Now, I mean, that that move could be contested. I agree. That's not, that's not self-evidently true. But I think that's the natural default assumption that this is the home of this vocabulary. You know, it's really it's really weird to my mind to think that we those, the psychological language of that sort of visually characterizes what an earthworm does. I just don't think it's true. Right?

    Marty Martin 29:22

    Yeah. And that's fair. But I think that's where I'm coming from, I mean, to just sort of distinguish these forms of agency, using human cognition, as sort of a frame for how we think about all of these other things, I sort of approach it from the other direction. Life as a complex system, one of its manifestations ends up being the human brain, but even the vertebrate immune system can learn in an exquisite way. That has nothing to do with how nerves learn. I mean, if we had a better grasp on what consciousness was and the interplay between consciousness and mental, maybe I'd feel a little bit more comfortable but because those are such abstractions t0o, I don't know, it feels like a strange place to start.

    Samir Okasha 30:04

    Right., I see. Yeah. I mean, why do I start in that place? You see, I mean, I entirely agree with you. That, for example, the adaptive immune system learns. Yeah, that seems that seems absolutely clear. But why do I start with the mental? I mean, I think it's because my, my specific interest is in this anthropomorphizing tendency, or at least in that book, that's the thing I'm specifically interested in is this anthropomorphizing tendency within evolutionary biology. And my overarching question, in that book, was like, Is this tendency to use the language of intentional psychology in an evolutionary context in order to, to pause adaptive explanations? Is there any real good reason to do that? Or is it just a sort of casual anthropomorphic bias? I mean, is it? Does it pick out some genuine class of phenomena? Or is it does it really just more reflect the fact that the investigators are humans themselves and haven't entirely always managed to shed themselves of their anthropomorphic bias? But in terms of thinking about the origins of agency itself, and the origins of intentionality and consciousness? I mean, I think I agree with you that precursor forms of all of these things are found in a much simpler biological systems, and that somehow rather, you know, the evolutionary process gave rise to creatures with whose cognition was sophisticated enough, for whatever reasons exactly, that they could actually mentally represent their goals.

    Art Woods 31:43

    Yeah. So Samir, I gathered from your book that you sort of slot these different ideas about sort of levels of agency into what you call type one and type two. And type one is about organismal agency itself, which I think, you know, now many biologists are starting to hear about and having strong reactions to. Type two is this idea that the process of evolution itself and natural selection can be agential and, or that it may appear to be agential because of the sort of psychological, unintentional language that's used. And I would say, like, it feels like your book is largely in the end dismisses this idea that type two agency is important and actually captures something real that's happening in biology and that, we're sort of, in a sense, taken for a ride by the the language that we use to describe type to agency.

    Samir Okasha 32:42

    Yeah, that's exactly right. So it was, yeah, my sort of simple one line argument is a Type One good, in certain cases, scientifically justifiable, despite being anthropomorphic. Type Two, generally not good. Anthropomorphic, but bad. So type two, in the least the paradigm, sort I have in mind involves, you know, thinking of the evolutionary process, as you say, or the process of natural selection, as a form of agency, if you like. And Darwin himself was the first to do this in, you know, in a number of passages in the Origin of Species, where he says natural things like natural selection is "daily and hourly scrutinizing the world" see "looking for any advantage and preserving what's good, eliminating what's bad." So there he;s quite deliberately using this intentional language to characterize the process of natural selection itself. Now, Darwin went on to say, "Look, these are just metaphors, but don't worry about it. You know, it's not, it's not a big deal." And I think, in fact, he was cavalier on that point, because, I mean, one of the things that people didn't realize, in in the didn't understand, in the earliest days of post-Darwin was that the process is, of course, mechanistic, and doesn't have any foresight, there is no goal towards which evolution is striving or anything like that. And, you know, any attempt to, to think otherwise is really as to misunderstand Darwin's achievement.

    Samir Okasha 34:17

    But nonetheless, I do, I do think that sort of vestiges of that Type Two form of agential reasoning and thinking are still alive today. In fact, because you see, I think that we very often treat the simplest sort of natural selection as if it was paradigmatic. So by that I mean, the simplest sort of natural selection is just one we've got a number of variants in a population in a fixed environment, with no frequency dependence. And the evolutionary process when driven by natural selection simply involves fixation of the best variant and elimination of the worst ones and, you know, monotonic increase of mean population fitness until it reaches a maximum, pushes the population up the adaptive landscape. All ideas that are, you know, have their place and are useful but rests on. You know, that's only the very simplest case. And, you know, it's all evolutionists know, I mean those things hold true under restrictive model assumptions that almost never apply in the real world. So I think the tendency to adopt that language leads us to think that the simplest form of natural selection, as a sort of simple optimization process, is the paradigm example. And obviously people, you know, who really know the science know that that isn't true. But nonetheless, I think a vast number of people who sort of know a bit about evolution and natural selection, don't understand that that isn't true. And the use of this intentional language, thinking of the evolutionary processes, you know, either driven to achieve some goal or is in any case, as directed in some sort of way, or as pushing populations up landscapes by the steepest path, that sort of thing. Those mistaken notions go hand in hand, I think, with the use of these anthropomorphic intentional descriptors to characterize the process of evolution by natural selection. So that's why I say Type Two bad. But I want to say Type One, which involves thinking of an actual evolved entity, typically an organism, but possibly a group, or maybe even a gene, as akin to an agent, and as trying to achieve a goal, can be, in certain cases, a useful mode of description. I try and make this argument that although anthropomorphic, it does genuine work, in that it picks real phenomenon that, you know, certainly could be described otherwise, but that there is a real reason to describe this way. And that's when I start talking about integration and absence of of internal conflict, as being a prerequisite for thinking of an organism as agent-like

    Cameron Ghalambor 37:08

    Yeah, so this is a good place to really just ask the question, are genes agents, especially in the context of thinking about transposable elements, and gene for altruism or other kinds of intergenomic conflict? It seems like our language that we would use and our theory to describe why there would be selection for or against the spread of a gene within a genome fits within this concept of an agent.

    Samir Okasha 37:54

    Yeah, no, I agree with that. And I think I mean, the broad story I'm trying to tell about agency and the circumstances in which it's a valid concept or useful concept, at least to employ in an evolutionary context includes the sort of famous Selfish Gene notion of Dawkins, although I do think of the many different sort of lines of rationales and lines of reasoning that Dawkins uses to support The Selfish Gene notion, the one that I'm interested in is the one you allude to, that is where there's intergenomic conflict, that it makes most sense to think of a gene as trying to achieve an outcome by means of doing something that in this case, harms are the genes within the same genome. By now, a familiar point that and one that I think Dawkins himself made in his later work that the sense in which the gene is the unit of selection, where there's intergenomic conflict is really rather different from the sense in which the gene is always the unit of selection. I mean, the latter is true, in a sense, but it's in a different sense that we talk about genes as the entities being selected, where there's conflict, within a single genome. So I sometimes captured by contrasting the process of selection of gene-level selection, with a genic perspective on selection processes that take place at a different level.

    Marty Martin 39:24

    Yeah. Samir, can I ask you to tie back genes as agents to your four types of agents that we started with into which category would genes fit for you?

    Samir Okasha 39:35

    Well, I would say really, literally speaking, none of them because I mean, you know, DNA is a pretty inert molecule. I mean, obviously, the more we know about, you know, how genes work and how you know how gene expression works. You know, the more the situation is a little bit complicated, but I would still say that, if we're asking which of those ages concepts literally characterize genes, I would be inclined to say none. But nonetheless, it can be useful to use one or more of those concepts metaphorically to describe genes for certain projects. And I think that, you know, what Dawkins and Dennett and fellow travelers are doing is basically taking one of those notions of agency, namely the intentional agency one, and using that notion, applying that notion to genes, but in an overtly self consciously metaphorical sense. And I think that I've always been puzzled why people are so horrified by that. I think it makes extremely good sense to me. And so long as it's used carefully.

    Art Woods 40:45

    It's funny, I feel relatively comfortable with, you know, the idea of genes as a level of selection. It's hard for me to conceive, you sort of said this, but it's hard for me to conceive of genes as agents, because, you know, to me, agency itself comes from networks, doing computations and making decisions. And of course, genes don't contain those networks, right? They can be a component of another network, but they're just a part that is participating in a greater whole that maybe has agency, but they don't have agency themselves. What do you think of that?

    Samir Okasha 41:27

    Yeah, I mean, I think I would agree. But again, that's because you're using the terms in this literal way. I mean, I think that what you say that genes don't have themselves, any of these characteristic hallmarks of agency, I think is true. But I also think that that's compatible with heuristically treating them as if they did, in order to further our understanding of the evolutionary processes of certain evolutionary processes. And so I mean, if you think of, if you think of how people struggled to understand the phenomenon of intergenomic conflict, before they had The Selfish Gene paradigm, which was really invented for other reasons, really. You know, Dawkins was originally thinking about things like altruistic behavior, and so on. But I mean, I think, you know, once you have that idea, then you can immediately see how to make sense of this otherwise extremely puzzling phenomenon.

    Cameron Ghalambor 42:26

    I think it'd be useful to give an example of this type of inner genomic conflict, is there a particular one that you you really like,

    Samir Okasha 42:34

    No particular one, I mean, there are, you know, thousands of these examples. But take, for example, the phenomenon of cytoplasmic male sterility in flowering plants, where you have a gene, within a mitochondrion, that causes this plant to cease making pollen, which is a really odd thing to do. You know, the plant, in some cases, actually grows stamen. So, you know, to all intents and purposes, in order to make pollen, but then those mitochondrial genes frustrate the stamen actually doing the thing that they're meant to do. I mean, that's from the phenotypic perspective of the whole organism, you look at them, what on earth is that about? What's the rationale for that? Does this you know, is this an exception to the theory of evolution or something like that? How, how could it be? But it makes perfect sense when you realize that the mitochondrial genes are only transmitted maternally down the female line, so investing in pollen is a waste. That's the sort of case I have in mind, where I say that there's a threat to the idea that the whole organism is agent-like, because it doesn't have if you like, the sort of phenotypic integration and behavioral integration that one would need in order to apply the psychological language metaphorically to it. So what is it that the plant is trying to achieve? You know, is it trying to self=fertilize or not? While on the one hand, yes, because, you know, a lot of its apparatus is geared to that. But on the other hand, no, because they don't make any pollen.

    Marty Martin 44:10

    Yeah. And this might be a bizarre thing to say, Samir, to the author of a book on levels of selection. But, you know, those kinds of arguments in what you make a completely fantastic case, and there's no question that these conflicts are true. But because biology is organized across levels, to sort of use these as ways to understand organisms, you know, is it possible that these are great examples of genes as parasites, or something different, and the rest, or a large fraction of the genome and all its regulatory components, they aren't so much parasites is commensals or mutualists. I mean, you know, to use the parasitic argument to be foundational for everything organismal, I've always viewed that as an overreach.

    Samir Okasha 44:55

    Yeah, I see. I mean, I think I think I'm on board with the idea that we can have genomic parasites and that that actually accurately captures quite a lot of cases of intergenomic conflict. So what's the thing that you think is overreach?

    Marty Martin 45:11

    You know, your arguments about why not make pollen? That sort of framed in an adaptive context at the organismal level, where another explanation is some type of frozen accident happened in the past that produces this lineage that, you know, the system is still alive, natural selection is still going to operate, as natural selection does, as life does, it makes the best of what it has. But to use that, to sort of expand that out to a full way to understand the evolutionary process. That seems to be ambitious.

    Samir Okasha 45:45

    Well, you see, what I'm trying to do is to make an argument that says, look, it's because phenomena like that are relatively rare, that the language of agency has any purchase at all, in an evolutionary context. In that we have this sort of general philosophical line of argument, which says something like, look, talk of intention, goal, belief, and desire is really only appropriate when there is a kind of rational integration, indicative of the fact that in cases say, of multiple personality, then we really struggled to say what on earth it is that a person, you know, whether there really is one person there at all, or what it is that they believe or what it is they want, we can't say it, you know, the psychological language just ceases to apply in that sort of case. And what I'm arguing is that we have a kind of biological analogue of that, in the sense that if there were too much intergenomic conflict, the phenotypic consequences of that would make it impossible for us to think of the organism was trying to achieve anything. And it would be impossible to metaphorically characterize it as you're aiming at some goal or other, if you like. So I guess I'm suggesting that the psychological integration and the rational integration is a necessary condition of applying psychological language literally. And that the biological or phenotypic integration that one gets when intergenomic conflict is effectively suppressed, as in the vast majority of extant organisms is a necessary condition of applying that psychological language, metaphorically, to organisms.

    Cameron Ghalambor 47:33

    And we see those types of conflicts not only intergenomically, but we also see those conflicts, for example, between the goals of males versus females and how those play out, you know, in terms of toxic sperm that may help an individual male against other competitors, but actually does harm to the female. And so it's the same conceptual framework that we're applying in those cases as well.

    Samir Okasha 47:59

    But in that case, the conflict is between genes in different organisms,or between two separate organisms.

    Cameron Ghalambor 48:05

    Yeah. Or yeah, two sexes, I guess.

    Samir Okasha 48:09

    So for that very reason, I think it would be, it would be a mistake to think that a mating pair was agent-like, if you see what I mean, or to take it to talk about what the mating pair is trying to achieve. Obviously, their interaction is partially mutualistic,

    Cameron Ghalambor 48:24

    You use the term integration, but also, I think, along with that you also use the term unity of purpose. And so I think that's the outcome of the integration, that there, these actions are towards the goal, towards this purpose, and it requires this integration or coordination to achieve that goal.

    Samir Okasha 48:46

    Absolutely. But, I mean, one thing I do stress is that it's a matter of degree. So, you know, there are local violations of this unity of purpose do certainly exist and are, you know, relatively. I mean, intergenomic conflict, as you guys will know, has been, you know, found in almost every taxa where it's been looked for. But nonetheless, I do think that those violations are necessarily local, if you'd like, in that if you had too much a breakdown of unity of purpose, you probably wouldn't actually have an organism at all. And you certainly wouldn't be able to think of it as akin to a rational agent, trying to achieve an end to which its various phenotypes all contribute.

    Art Woods 49:31

    Yeah. So I have a related question here that builds on what you just said about the idea of whether a mating pair has agency and that's just to expand. It's also kind of a levels question, but going in the other direction. And thinking about groups of organisms and whether or not they show agency and how that relates to their evolutionary processes. And, like at some level, it's easy to think of groups that almost certainly have agency like eusocial insect colonies in which many of the individuals are closely related to one another, and they act essentially like a super organism. But what about, say interspecific groups, which occur all the time in the wild, and I'm thinking of things like so there's a retired professor here at Montana, Eric Greene, who's worked some on communication among different bird species and between mammals and birds in forests. And you may be aware of some of this work, but there are waves of communication about predators that travel between the members of these interspecies groups. And that really define a lot about where these individuals go. And you know, when they're feeding and when they're being fearful and paying attention to predators that are coming by. So would you say that a large multi-species group like that also shows agency in some way?

    Samir Okasha 50:57

    I guess I would hesitate to say that, although it might be true. I mean, I don't think you could rule it out. Definitely. But I think the reason I would hesitate to say it is just for the same reasons that people have always been reluctant to think of such a multi species group as an adapted unit itself. I mean, clearly, the organisms in it are adapted. And in part, they're adapting themselves to the social environment that they find themselves in, which includes living in these multi-species communities. But that's still different from saying that, you know, the multi species group itself is adaptive. I mean, this was the goes back to the point that George Williams made way back in 1966, in his book Adaptation and Natural Selection, where he wanted to say, look, distinguish between a group adaptation and a fortuitous group benefit. In that we can't really easily conceive of an evolutionary process by which multi-species groups could have come to be the adapted units themselves, although, I mean, it's possible.

    Art Woods 52:03

    Because their evolutionary interests are not closely enough aligned, consistently, right?

    Samir Okasha 52:07

    Right. Exactly. Yeah. Because we would think of the evolutionary process as taking place on the separate lineages within those multi-species groups. And to think of, you know, the multi-species group is really part of the environments relevant to the evolution of the population, the conspecific populations that make them up. So I mean, I think that's the reason that I would hesitate to apply the language of agency here. But I would want to say that it is ultimately an empirical question that mean, there are people who, who takes seriously the idea of community-level natural selection, where the community can be a multi species aggregate of some sorts,

    Cameron Ghalambor 52:52

    But I think if we sort of relax the definition of integration, and I think another way to capture integration is to call it some degree of cooperation, whether it's at the whole organism level, you have to have these coordinated responses. I could also see cooperation, you know, obviously, within a group, but if we kind of relax a little bit, what we what we mean by cooperation, I could see also applying to the examples that Art was giving of, you know, mixed species, because ultimately, then, through that cooperation, what emerges is this unity of purpose, they all have the same interest in avoiding the predators.

    Samir Okasha 53:38

    Yeah, I mean, I think that that's possible, presumably, that that overlap of evolutionary interests will be kind of contingent, if you like, and could break down.

    Cameron Ghalambor 53:51

    Especially if some of those bird species are also competitors, and don't have, under other contexts, aligned differences. So yeah, I think that kind of context dependency probably plays an important role.

    Samir Okasha 54:06

    Right. But and, of course, even if the groups were not conspecific, you know, comprised of a single bird species, then, I mean, it would still be problematic to think of the group, of such a group, as an adapted unit automatically. I mean, because I mean, I again, I would be happy with that, if a demonstrable process of group level selection had shaped the evolution of some phenotype. So then we get back into this question of, you know, what sort of selection process is necessary in order to produce an adapted group, as opposed to a group of adapted individuals, which I've always thought is that is a helpful framing that distinction that Williams made.

    Marty Martin 54:50

    Yeah. Do you think that groups of humans, given what we were just talking about with individuals within a bird population, would groups of humans count as agential, or do you have the same qualifiers for us?

    Samir Okasha 55:04

    I mean, I think potentially. So, I mean, if you think for example of, you know, not not a group of humans, like a tribe or something much smaller, like, say, a committee, or something like that. Many people in philosophy of social science take very seriously the idea that groupings like committees can quite literally be agent-like. We can talk literally about what the committee believes, and decides and did. And that certainly is manifest in the way we talk ordinarily, so yeah, you know, the firm raise prices, the committee decided what to do, we do use this language of choice and decision and belief, as a standard psychological language to to groupings of humans in certain contexts. And so I'm happy with the idea that that could indeed be literally true, yeah. And certainly that it could be heuristically useful to think that way. But again, only, I mean, only, of course, under certain circumstances. I mean, if the committee couldn't come to a decision, or if, you know, one person on the committee said "X" and the other said, "Not X" then of course, we couldn't sensibly talk about what the committee thinks about the topic.

    Marty Martin 56:31

    So we want to transition more directly into the interplay between agency, your ideas about agency and classic evolutionary theory. So to put this in an explicitly evolutionary perspective, and we could stick with humans, or we can go in any direction you'd like. Is there evidence or do you expect that in nature, there is heritable variation for agency? I think as you were talking about agency there it wasn't obviously in the adaptive the classic evolutionary adaptive sense, but in natural populations, what do you think about heritable variation for agency?

    Samir Okasha 57:07

    Well, I suppose I would say, you know, agency depends on, in any of the senses that we've been discussing, it depends on the possession of certain phenotypes, such as, you know, information processing, sensing the environment, cognitive abilities of various sorts. And there must, of course, have been heritable variation in all of those for them to evolve, whether there still is now is perhaps a slightly different question. But I still think of that, I think of the heritable variation, as being with respect to phenotypes, that are prerequisites of or building blocks of agency, rather than thinking of agency itself as being something that that is, is itself a phenotype.

    Art Woods 57:55

    Huh, that's interesting. So you wouldn't call agency itself a trait, for which there would be heritable variation?

    Samir Okasha 58:02

    No, I think it would, I think it's almost at the wrong level itself to be a trait.

    Cameron Ghalambor 58:09

    But that raises a sort of an interesting problem for me, because, you know, if we go back to Fisher, he talked a lot about in his fundamental theorem, he talks about essentially, heritability for fitness. And yet, fitness is also made up of all of these other components. And so I see a lot of parallels there. But yet, in our models, we don't have a problem thinking about, you know, as long as there's additive genetic variance for fitness, and maybe that's because we're thinking about it at a population level as opposed to individual level.

    Samir Okasha 58:44

    Yeah, no, I mean, that's, I think that's an absolutely fair point. Yeah. So if we take it that, you know, fitness ultimately depends on the vital rates, birth and the death rates, and that those are contributed by many, many different phenotypes, of course, will contribute to the, you know, the vital rates of an organism of any sort. So, how is it then that we're happy to talk about heritability of fitness? Yeah, I mean, I see the point. You're saying, well, why by parity of argument, why can't we talk about agency as a heritable trait or something exhibiting heritable variation? I mean, it in part, maybe the difference is, in part, just that it's not so easy how you would measure it.

    Cameron Ghalambor 59:29

    Yeah, I agree.

    Samir Okasha 59:30

    Right. Yeah. I mean, so do we find an organism that's got more of it, then one that's got less of it? I mean, how would you?

    Art Woods 59:38

    Yeah, right. What does that mean, to see a population in which there's a distribution of agencies, right?

    Samir Okasha 59:44

    Yeah, no, that's right. But nonetheless, I mean, if we take it that in contemporary biological populations, that we do see agency, and presumably if we go far enough back in their evolutionary ancestors, then we don't find agency or at least only precursor forms of it, then some evolutionary process must have taken place to produce it. So in that sense, it's got to be an evolved attribute. But I'm still a little bit uncomfortable with thinking of it itself as a trait, in part, just because it seems hard to measure, and to depend on too many other things that have sort of clearer status as traits, let's say.

    Cameron Ghalambor 1:00:23

    I agree. And I think that actually is a point of argument between me and Marty and Art that, you know, I recognize that organisms, behave and act agentially, but I see it more as sort of an epiphenomenon, it's an emergent property that probably has evolved over time, but as an emergent property, it's something that, in of itself, is really difficult to, as Art was saying, like, you know, plot a distribution of agency in a population.

    Samir Okasha 1:00:57

    Absolutely. I mean, I suppose someone, you know, might make the argument, well, you know, maybe it's fixed within every species or something. So it doesn't show any distribution, you know, within a human population, for example.

    Cameron Ghalambor 1:01:14

    But then it wouldn't be able to evolve

    Samir Okasha 1:01:15

    Not but maybe it's evolved already. Maybe it's, you know, it's like bipedalism or something interesting.

    Marty Martin 1:01:21

    Yeah interesting. Well, it's a good thing in evolutionary biology, we don't have any other great big emergent words like gene and fitness and things like that.

    Cameron Ghalambor 1:01:29

    So Samir, you spent considerable time or space in the book focusing on Alan Grafen's Formal Darwinism project. And I think the goal of Grafen is to move away from population measures of fitness to thinking about individual optimization, in particular, as it relates to this idea of agency. It seems like Gragen's goals are to separate selection from individual optimization. But then I think you made this point in the book that natural selection may not always then lead to adaptation. And this seems like a very counterintuitive conclusion. Could you help me understand how those gets separated from one another?

    Samir Okasha 1:02:24

    How natural selection and adaptation gets separated from one another?

    Cameron Ghalambor 1:02:28

    Yeah. And this idea that that selection alone can, in some cases, not lead to adaptation?

    Samir Okasha 1:02:35

    Right, yeah. No, I mean, what I was getting at with that was just the drawing attention to some points that, you know, are fairly well known among evolutionary theorists. Phenomena, for example, of cyclical dynamics, right, where even in simple one locus, population genetics, then we can point to certain circumstances under which equilibrium will not be attained. But rather gene frequencies will cycle indefinitely, never settling down and attaining equilibrium, even though natural selection is the only force at work, is the only evolutionary force at work, and it's not a stochastic effect. It's nothing like that. So I think simple examples like that, that seemed to illustrate, you know, fairly clearly, I think that although we usually think of natural selection as an engine for adaptation, as producing organisms that behave as if they're trying to achieve some end, some fitness related end, there's no sort of logical guarantee that that's got to be the case.

    Samir Okasha 1:03:39

    And I think one sees the same in models of frequency dependent selection. So if you look in the popular Adaptive Dynamics framework for thinking about that, although the same is true in other formal frameworks, as well, for thinking about frequency dependent selection, it's not always the case that natural selection will take the population to an equilibrium at which individuals are playing the optimal strategy, conditional on the rest of the population. I mean, often it will. And that's a nice case, in which we get an evolutionary stable strategy as the outcome of frequency dependent selection. But unpleasant, nasty, nonlinear dynamics or cyclical dynamics, for example, and other things are possible too, as people have known for a long, long time, even though natural selection is the only force at work.

    Samir Okasha 1:04:29

    So I think of I mean, what Grafen is doing, as I see it as, in part, a response to that situation, in that he's trying to say something like, "Well, I mean, can we say something about when natural selection will lead to adaptation," and he thinks that there's got to be an intimate and tight link between them. And I think, intuitively, that makes sense. I mean, the whole point of the theory of natural selection was precisely to explain two phenomena, adaptation and diversity. But then we faced this fact that when exposed to evolutionary models, you know obviously went beyond what Darwin himself was thinking of, we find that natural selection sometimes seems to result in something that we can't really call adaptation in any clear sense. So then we want to say, well, can we point out conditions under which it will, and that I think of as, as what Grafen is trying to do in his project, and I think it is very useful, although I don't think all the moves he makes are ultimately successful. I think, conceptually, it's extremely useful to say, look, on the one hand, we have the process of natural selection, and on the other hand, we have the phenomenon of adaptation to the environment. And the one is meant to explain the other. And the fitness concept, rather oddly, it sort of features on both sides of that equation.

    Cameron Ghalambor 1:05:52

    My follow up to that is, it seems like there's another concept here that is somehow lost in the arguments, which is variation. And so I completely agree in the simplistic models, you know, we think about selection leading to adaptation and the fixation of a single sort of variant. But whether it's frequency dependence, or density dependence, or just if there's fluctuating environments, it seems like one outcome of that is that no single variant will always be optimal. And a byproduct of that is that variation is maintained in the system, in the population, in this case. And so I'm just kind of curious, where variation among different agents or different individuals in a population falls within this? We touched on this a little bit that, you know, maybe a bit like bipedalism. It evolves and then becomes fixed. But I feel like, at some level, we have to incorporate variation into these ideas.

    Samir Okasha 1:07:04

    Yeah, no, I mean, absolutely. I mean, I think everybody would agree that, you know, you need variation in order for this to be natural selection at all. If all organisms are the same in some respect, and there's no way that some variants can out proliferate others, everyone has the same variant, if you like. So that I think is presumably common ground, that variation is necessary for natural selection. But I mean, there's also the different question of whether the outcome of natural selection will be to preserve variation, or to eliminate it.

    Cameron Ghalambor 1:07:37

    Yeah, we think about it, eliminating it usually.

    Samir Okasha 1:07:39

    Right, we do. We do. But of course, as you rightly point out, I mean, if there's, you know, environmental heterogeneity temporal, or spatial, or a host of other complications, then or non additive genetics, for example, then we can have scenarios where variation, genetic and phenotypic is, in fact, preserved by natural selection or natural selection can lead to adaptive diversification, for example, as well. So our sort of starting, you know, assumption that natural selection will reduce variation, although, you know, true, usually for most, for the simplest sort of natural selection need not always be right.

    Art Woods 1:08:24

    So, one of the issues you brought up is that there's lots of variation in the environment, and so often there's not going to be a single genotype that's optimal. Is it possible that things like plasticity, and then maybe on top of that, agential behaviors sort of writ large, are a response to that? An evolved outcome to deal with that very kind of uncertainty, right? So there's these systems that are much more flexible over much shorter time frames then, you know, alleles in genomes are that allow organisms to deal with that kind of uncertainty?

    Samir Okasha 1:09:03

    to deal with environmental uncertainty?

    Art Woods 1:09:06

    Yeah, yeah.

    Samir Okasha 1:09:07

    Yeah. So you're thinking of an environment that changes sort of within the lifetime of an organism?

    Art Woods 1:09:16

    Yeah, within or between. But I mean, either way, the systems that produce these organisms during development, inherently have this sort of flexibility and plasticity and agency built into them to sort of compensate for the fact that, of course, they're not going to have the ideal alleles at all the loci, there's no way they possibly could, right?

    Samir Okasha 1:09:37

    No, I mean, I think that that may well be true. I mean, of course, I guess it's not the only possible way that natural selection could adapt a species to environmental uncertainty. So, I mean, I think that's absolutely right. But, I mean, environmental fluctuations and heterogeneity and uncertainty are presumably pervasive in the living world, I mean, pretty much all lineages must face those. So then I guess the question is why some lineages evolved these agent-like behaviors or capacities for dealing with them, whereas others didn't? You know, so an alternative might be to engage in some sort of bet hedging or diversification, for example, where a given genotype produces lots of different phenotypes in order to hedge against the environmental uncertainty, you know, so one of those seed types will germinate, whatever the weather turns out to be, that well known idea. So for me, the question is, I mean, can we can anyone say something sort of concrete about the particular sorts of environments that led to the elements of agency being the adaptive response, as opposed to sort of sub-agential things that one finds more widely?

    Marty Martin 1:10:59

    Yeah, well Samir, we've really enjoyed the conversation. I really appreciate your effort. This has been wonderful, being great.

    Samir Okasha 1:11:05

    Oh, it's great. I mean, it's, you know, I could talk about this stuff all day, as I'm sure you guys could, too.

    Marty Martin 1:11:12

    I think we all could as well. Well excellent. So we always like to wrap with sort of giving you a blank page to say anything about the book or future plans or anything else you'd like to talk about. Is there anything else you wanted to hit?

    Samir Okasha 1:11:24

    Not particularly, I'm currently writing a book about the concept of fitness, and why it's caused so much controversy and difficulty and why it's not as simple a matter as it should be.

    Marty Martin 1:11:36

    Okay, the subject for our next conversation, please let us know when that one comes out. That would be wonderful.

    Samir Okasha 1:11:42

    Ok I will, hopefully this next year.

    Art Woods 1:11:46

    Yeah. Send us the book when it's out.

    Marty Martin 1:11:49

    Okay, well, thank you so much.

    Art Woods 1:11:50

    Yeah, thanks, Samir.

    Cameron Ghalambor 1:11:51

    Thank you.

    Samir Okasha 1:11:52

    My pleasure. Thank you all.

    Marty Martin 1:12:11

    Thanks for listening. If you like what you hear, let us know via X, Facebook, Instagram or leave a review wherever you get your podcast and if you don't, we'd love to know that too. Write to us at info at big biology.org

    Art Woods 1:12:22

    Thanks to Steve Lane who manages the website and Molly Magid for producing the episode.

    Cameron Ghalambor 1:12:26

    Thanks also to Dayna De La Cruz for her amazing social media.

    Marty Martin 1:12:30

    Keating Shahmehri produces our awesome cover art.

    Art Woods 1:12:33

    Thanks also to the College of Public Health at the University of South Florida and the National Science Foundation for support.

    Cameron Ghalambor 1:12:38

    Music on the episode is from Podington Bear and Tieren Costello

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