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Hey, Big Biology listeners! A reminder that we're

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for just a few dollars a month.

1:02

Patrons also get cool insider stuff, like

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access to behind-the-scenes audio from our guests

1:07

about their lives, their hobbies, and their

1:09

careers. Now, on to the show. Marty

1:22

and Art, it's good to see you

1:25

two philosophers today. Philosophers? Norwegian

1:27

Winter got you confused, Cam.

1:29

We're biologists. Wait, but PhD

1:32

stands for Doctor of Philosophy.

1:35

Technically, Cam's right. We are philosophers

1:37

in the sense that all sciences

1:39

are rooted historically in philosophy, and

1:42

that meant the pursuit of wisdom,

1:44

scholarship, and expertise. Fail. But

1:47

there is a discipline where the

1:49

two meet. The philosophy of biology

1:51

explores fundamental topics like how the

1:53

meaning of terms and critical assessments

1:55

of assumptions have changed through time.

1:57

And if you're a regular big biology listener, you

2:00

probably heard past episodes with philosophers

2:02

of biology like Dan Nicholson, Dennis

2:04

Walsh, and many others. I

2:06

think most practicing biologists, myself included, rarely

2:08

read the philosophy of biology literature, but

2:11

those are some of my favorite episodes

2:13

because they've made me stop and reflect

2:15

on ideas and concepts that I often

2:17

use but never really thought deeply about

2:20

their meanings. In a 2019

2:22

article in the Proceedings of the National

2:24

Academy of Sciences, Lucy Laplan and colleagues

2:26

argued that philosophy plays a critical role

2:28

in the sciences. An example they highlight

2:30

is a critique of the cell versus

2:33

non-self idea in immunology. Philosophers pointed out

2:35

that the idea of self breaks down

2:37

will be recognized that all organisms are

2:39

made up of communities of non-self symbionts

2:42

and the shift has real-life implications

2:44

for developing medical treatments. Thank you

2:46

philosophers. One active area

2:48

in the philosophy of biology is

2:50

evolution by natural selection. Concepts

2:53

like fitness, selection, adaptation, contingency,

2:55

and determinism pop up all

2:58

the time. But talk

3:00

to philosophers of biology and you find out that

3:02

their meanings are far less clear than you would

3:04

think. For example, think about the

3:06

phrase survival of the fittest. When

3:09

we define fitness as the ability to

3:11

survive and reproduce, the phrase becomes tautological

3:13

because we're saying essentially survival of those

3:16

who survive. This is why the philosopher

3:18

Eliot Sober has said we need a

3:20

better definition of fitness. Another topic we've

3:22

covered a lot recently on the show

3:24

and that also shows up in a

3:27

lot of philosophical analyses is

3:29

agency. Agency has been

3:31

defined very differently inside and outside of

3:33

biology. Agency also carries a lot of

3:36

baggage which puts a lot of biologists

3:38

on edge. When you say

3:40

that organisms have purposes and goals, it

3:43

opens the door to conscious intention

3:45

and even supernatural effects and forces.

3:47

Yet even if we recognize that

3:49

most of what organisms do to

3:51

survive and reproduce is done unconsciously,

3:53

we often use casual and

3:56

sometimes clumsy language. Think about the cuckoo,

3:58

a bird that lays eggs in

4:00

the nest of other species. The cuckoo egg

4:02

typically hatches first, and the cuckoo chick then

4:04

pushes the host eggs out of the nest.

4:07

We might casually say that the cuckoo chick

4:09

does this because it consciously wants all the

4:11

food for itself. We often use

4:13

this kind of language as shorthand to help

4:15

us explain why the chick behaves the way

4:17

it does, even though it

4:19

may not be making a conscious choice.

4:22

A more traditional explanation would say that

4:24

this behavior evolved by natural selection acting

4:26

on heritable variation in the context of

4:28

an evolutionary arms race with its host.

4:31

So, while there is a growing interest

4:33

in the notion of agency in evolutionary

4:35

biology, there's still a lot of confusion

4:37

and debate, even among us, about what

4:39

it means, as well as its relationship

4:41

to fitness, selection, and adaptation.

4:44

Our guest today is Samir Akasha, who's

4:46

a professor of philosophy of science at

4:48

the University of Bristol in the UK.

4:50

His 2006 book, Evolution and the Levels

4:52

of Selection, won the Lakatos Award. Today,

4:54

we talk with Samir about his 2018

4:56

book entitled Agents and

4:59

Goals in Evolution. In contrast to

5:02

our past discussions of agency, Samir takes

5:04

a different approach. He

5:06

asks in the book, quote, Does

5:08

thinking about organisms as agents have

5:10

a genuine scientific role to play in

5:12

our understanding of adaptive evolution, or is

5:14

it simply another application of anthropomorphic

5:16

language? In our chat, we

5:18

look critically at the different definitions of

5:21

agency in biology. How is it different

5:23

from phenotypic plasticity? And do

5:25

we separate agency and fitness? And

5:27

what about agency below the organismal level?

5:29

Could a gene be considered an agent

5:31

if it acts selfishly to increase copies

5:33

of itself, like in cases of intergenomic

5:35

conflict? And what about groups

5:38

of humans or even groups of different

5:40

species that exhibit cooperative, goal-directed behavior? Do

5:43

groups have agency distinct from individual

5:45

members? Samir argues that

5:47

although we gain insight by

5:49

anthropomorphizing some adaptive traits, there's

5:51

still value in understanding how agency

5:53

influences evolution, even if we don't

5:55

know about consciousness in that organism.

5:58

We also discuss whether the process of

6:00

evolution by natural selection represents a form

6:02

of agency. This might sound odd,

6:05

but recall that Darwin himself depicted natural

6:07

selection with a language of intent, as

6:09

a background process that's always scrutinizing

6:12

populations, keeping the good and eliminating

6:14

the bad. We debate

6:16

some really difficult problems like whether agency

6:18

can evolve, is there heritable variation for

6:20

it, and how would we measure that?

6:22

Something we've been thinking about a lot,

6:25

but I'm not sure we've totally resolved.

6:27

So stay tuned for a philosophical assessment

6:29

of agents, goals, and evolution. I'm Cameron

6:31

Gallenbore. I'm Art Woods. And I'm Marty

6:33

Martin, and you're listening to BIG Biology.

6:51

Samira Kasha, thanks so much for joining us on

6:53

BIG Biology today. It's my pleasure.

6:55

We're super excited to talk to you

6:57

today about your recent book, Agents and

7:00

Goals and Evolution. Maybe to

7:02

begin, could you start off by

7:04

defining what is a

7:06

biological agent? Well, that's a

7:08

complicated and difficult matter. Let

7:11

me give you my take on it. As

7:13

I see it, the place we need to start is

7:15

by asking pretty much the question you did. I mean,

7:18

what do we mean by the term agent? And

7:20

why do people think that this is a good concept

7:22

or term to use when thinking about

7:25

biology? And I think that's an important

7:27

question to ask, because there's this flurry of

7:29

interest in the notion of

7:32

agency in biology, as many of

7:34

your past contributors have discussed, and as you'll

7:36

be aware, and as a lot of philosophers

7:38

have joined this discussion too. But

7:40

on the face of it, I mean, that's a

7:43

bit odd, because if you open any biology textbook

7:45

in any branch of biology, I

7:47

very much doubt that the term agent is going

7:49

to appear in the index. So

7:52

that basically raises the question, well, why should

7:54

this notion be so important for sort of

7:56

philosophical reflections about biology if the science itself

7:58

is a good thing? doesn't seem to have

8:00

any use for it. So that's I think the first sort

8:03

of skeptical question I would ask and next

8:05

move I would make is then to say well if

8:07

people do want to talk about this let's do the

8:10

sort of standard thing we do in philosophy of saying

8:12

well what exactly do you mean? It's

8:14

always a good question to ask. And

8:16

as I see it there are I mean there are

8:19

a number of different motions of agents at work

8:21

both in different sciences and

8:23

in vernacular English. So

8:25

in my book I distinguish between

8:27

four different notions of

8:29

age and as a sort of prelude to

8:32

the discussion. The first is

8:34

what in in traditional philosophy we

8:36

would call an intentional agent. Someone

8:38

or thing that engages in reasoning

8:40

and in thinking and in deliberating

8:42

about what to do and choose

8:45

these courses of actions because of particular

8:47

goals that they want to achieve. Where

8:50

the goal is something that's mentally represented

8:52

by the agent. So that means

8:54

that at the very least you've got to have a mind

8:57

of some sort. Now what exactly that

8:59

I mean that's clearly intended to include

9:01

humans. Whether it includes what

9:03

else it includes you know whether

9:05

it includes all vertebrates or even

9:07

some invertebrates or even possibly microbes

9:10

it could be debated. But that as I see

9:12

it is the sort of traditional philosophical notion of

9:15

agent. An agent engages not

9:17

just in behavior but in action where

9:20

action means that they're deliberately trying to

9:22

achieve the goal. And that's what

9:25

I think of as the most

9:27

restrictive notion of agent. The literal

9:29

applicability of which is the least

9:31

if you see what I mean.

9:33

But then in different sciences we

9:35

find broader notions of agents. So

9:37

for example in AI one finds

9:39

frequent reference to talk about intelligent

9:41

agents. Where that basically means

9:43

anything you could be a control system

9:45

or piece of software or a thermostat

9:47

that doesn't always do the same thing.

9:50

That does different things depending on

9:52

the circumstance that it's in. So

9:54

typically we'll sense the environment somehow

9:57

and change its behavior depending on what

9:59

it says. I mean, so implement

10:01

stimulus response conditionals. It's an

10:03

extremely inclusive notion of agents,

10:06

that of course. Similarly,

10:08

but differently, one finds in the

10:10

economics literature talk about rational agents

10:12

and a lot of talk about

10:14

the rational agent model, where

10:17

essentially that means an

10:19

agent who engages in consistent

10:21

choice, so behaves as if

10:24

they're trying to maximize some utility

10:26

function or something like that, as

10:29

whose choice behavior is

10:31

consistent, satisfies conditions such

10:34

as transitivity, things like that. Again,

10:36

that's a broad notion of agents

10:38

in that there are no psychological

10:40

requirements. I mean, any organism

10:42

that engages in choice or behavior

10:45

of any sort could satisfy

10:47

or fail to satisfy those conditions on

10:49

being a rational agent as

10:52

understood in rational choice theory

10:54

and economics. And then

10:56

the final notion of agent that I operate with,

10:59

or that some people operate with that I distinguish,

11:01

is what I have called minimal agent.

11:04

And that just means something that

11:06

does something. So an agent

11:09

in this sense is just something that does

11:11

something rather than something to

11:13

which things happen, if you like. And

11:16

how exactly you cash that out, I

11:19

think is a difficult philosophical question. But

11:22

I mean, examples can make it clear. So to

11:24

borrow one from the philosopher Fred Dretsky that I

11:26

like to use, he said, look,

11:28

contrast these two things. A rat

11:30

moves its paw in

11:33

order to press a lever to get a pellet

11:35

of food. And an

11:38

experimental scientist puts the hand into the

11:40

cage and forcibly moves the rat's paw.

11:43

In the first case, the rat moved its

11:45

paw. In the second case, its paw was

11:47

moved. And

11:49

that he says illustrates the difference between

11:52

doing something and having something happen to

11:54

one. Sort of active versus passive. Yeah,

11:57

exactly. And then

11:59

obviously... the next thing to say about that is, well,

12:01

what, how do we cash that out precisely? What does that

12:03

really mean? And some philosophers have

12:05

thought that the answer simply lies in the

12:08

question of whether the cause was internal to

12:10

the agent or not. You

12:13

know, so if a bird flies, then

12:15

that's one thing, a hurricane, force, wind

12:18

blows the bird. That's another thing. In

12:20

one case, the cause of the, the

12:22

motor output is internal. In the second

12:24

case, there's an external cause. So

12:27

that, that's what I call the most minimal notion of

12:29

agents entirely. So those

12:31

are the four notions I operate

12:33

within some sort of rough hierarchy,

12:35

the most demanding, the intentional agent,

12:37

the least demanding, the minimal agent,

12:39

and then the intelligent agent and

12:42

the rational agent somewhere in between

12:44

those two extremes. And

12:46

then the next question to ask is,

12:48

well, which event, if any of those

12:50

notions applies to biological systems, to

12:53

which biological systems do they apply and

12:55

how do we know and why? But here

12:58

a complication comes in because I maintain, and

13:00

this is my, I guess my real interest

13:02

in my book, the much

13:04

talk of agency in the

13:06

biological realm is metaphorical rather

13:08

than literal. Particularly

13:10

in evolutionary thinking, I

13:13

claim that there's a lot of appeal

13:16

to the intentional notion of agent, the

13:18

most demanding of all, that

13:20

I think of as in many ways, the core notion,

13:23

where the agent must have a mental representation of

13:25

the goal. But the concept

13:27

is only applied metaphorically. When we say

13:29

things like the bacterium swims up an

13:31

oxygen gradient, because it wants to get to

13:33

an area of higher oxygen concentration or something like that. Yeah.

13:37

So let's, let's unpack that first one a little

13:39

bit, because I had a question lined up about

13:41

that very thing. So, you know, in the, in

13:43

the book, you make it clear that for this

13:45

more restrictive idea of agency, that it requires a

13:49

mind and a mental representation.

13:51

And at least like among

13:53

those of us, the hosts that are talking

13:56

about agency, Marty and I have sort of kicked this

13:58

back and forth. I think our concern of

14:00

agency is a little bit broader

14:02

and it encompasses basically any complex

14:04

system that's able to receive

14:07

information, process it in some way,

14:09

and make a decision. And that

14:12

doesn't invoke anything about nervous

14:14

systems or minds or mental representations. You

14:16

certainly could have biochemical networks that are

14:18

making decisions. You could make, you know,

14:21

have sort of genetic regulatory networks that

14:23

are making decisions. And so I think

14:25

that that immediately expands this

14:27

idea of agency down to very

14:30

simple organisms like an, you know, an E.

14:32

coli deciding whether or not to make the

14:34

machinery to metabolize lactose or

14:36

not. So in your conception, why do you

14:38

have to have a brain and a mental

14:40

map? Well, I don't say that

14:43

you do. I only say that

14:45

you do in order to be a

14:47

literal agent in one sense of the

14:49

term agent. But I entirely hear your

14:51

line of argument. And I think it's

14:53

one I'm sympathetic to that says, look,

14:56

there are, you know, other, so what

14:58

you might call hallmarks of agency are

15:00

found in pretty much all complex

15:02

systems, biological and maybe

15:04

even non-biological, certainly including

15:07

cells and possibly subcellular

15:09

constituents and molecular

15:11

pathways and things too. However,

15:14

I would say that that involves a somewhat

15:16

different notion of agents. So

15:18

I would actually say that's closer to the

15:21

intelligent agent notion of AI. So

15:23

basically, what we mean is that

15:25

we've got something that receives

15:28

information from the environment

15:30

and modifies its behavior

15:33

as a consequence. So in short, it doesn't always

15:35

do the same thing. What it does depends on,

15:37

on circumstance, if

15:40

you like. But then I mean, then

15:42

if that's right, then it raises the

15:44

question, do we just mean behavioral plasticity?

15:47

When we talk about agency in that sense? Exactly

15:50

what I wanted to ask. Is

15:53

this a continuous grade from intelligent

15:55

to intentional? I mean,

15:57

and is there some way to find a clear

16:00

demarcation between these kinds of

16:02

things? Well, presumably, I mean,

16:04

the intentional one is

16:07

a relatively late arrive on the

16:09

evolutionary scene. In that,

16:11

you know, however widely we think the

16:14

true intentional agency is found in

16:17

the animal world, and, you know, cognitive pathologists

16:20

have, you know, often debated, for

16:22

example, whether corvids are capable of genuine

16:25

intentional behavior or not, and opinions divide on

16:27

that issue. But what's very, very clear is

16:30

that, well, I would

16:32

say at least the vast majority of

16:34

organisms, including all invertebrates, are

16:36

not literally intentional agents. I mean, if we,

16:38

we might describe them as if they were,

16:40

when we say, you know, if the cell

16:42

wants to achieve such and such an end,

16:44

but that's, to deliberately

16:46

anthropomorphize, and maybe for good reason,

16:49

but it's not literally true. It's a

16:51

bit like saying the suitcase doesn't want to close, you

16:53

know, it might be useful thing to say, it's not

16:55

literally true. Right. What do you think

16:58

Lars Chitka might say about that? I'm not sure

17:00

if you're familiar with his work, but we spoke

17:02

to him quite a while ago. So

17:04

he's very interested in insect cognition. And

17:06

he's done really amazing work on all

17:08

sorts of bees in terms of their

17:11

capacity for solving puzzles. One

17:13

of the really neat stories that he tells is,

17:15

you know, how do bees decide,

17:17

decide to

17:19

build the nest that they do, and

17:22

the configurations that they can take, you know,

17:24

it's not like it's a sort of hardwired

17:26

always the same kind of

17:28

nest, it's very responsive to

17:31

the conditions of the environment. And he's

17:33

done incredibly sophisticated experiments to show the

17:36

amazingly elaborate kinds of outcomes from, you

17:38

know, presumably a simple cognitive

17:40

system. Yeah, no,

17:42

I mean, I think it's a genuine challenge

17:44

that I mean, at some level, it's got

17:46

to be an empirical question, I think. One

17:49

way to look at it would be to say,

17:51

well, what's to be gained, this is a formulation

17:53

inspired by Daniel Dennett, where he says something

17:56

like, heuristically, what would we gain, if

17:58

anything, by treating some

18:01

organism or collection of organisms in

18:04

the B case perhaps as an

18:06

intentional agent as if it really was trying

18:08

to achieve some goal and had

18:11

beliefs about the world and processing

18:13

information and so on. I

18:15

mean is that a useful perspective or not?

18:17

I mean the answer might very well be

18:19

yes in that sort of example but I

18:21

take it that there are examples where it

18:24

would not where we have a

18:26

purely mechanistic understanding of

18:28

the movement of some microbe

18:31

or something like that or the

18:33

growth pathway of some plant for

18:36

example. I mean it would be seem

18:38

superfluous to give a psychological explanation

18:41

of plant growth towards the

18:44

light or something then. I

18:46

mean we have a fairly

18:48

good mechanistic understanding of why the

18:51

plant responds to light

18:53

that way and if someone wanted to say well

18:56

no it's because it wants to get more sunlight

18:58

or something or wants

19:00

to grow taller than its neighbor. That would not be,

19:03

I take it scientifically useful. So one test

19:05

is just the scientific utility. Now

19:08

some philosophers and others would oppose that

19:10

methodologically by saying look the question of

19:12

whether something is useful is distinct from

19:14

the question of whether it's true and

19:16

that may be but nonetheless I

19:18

think if it isn't useful at all then

19:20

that's some indication that it's not literally true.

19:23

So as I see it I mean I wouldn't at

19:25

all insist that we know that it could only be

19:28

in vertebrates that we find

19:30

the hallmarks of true intentional agency. I

19:32

mean another point though to bear in

19:34

mind is that I mean

19:36

I started when I defined intentional agency by saying

19:39

that there really is a mental representation of the

19:41

goal. The organism really

19:43

it's not just as if it was trying

19:45

to achieve a goal it really is. But

19:48

you see if you talk instead about decision then

19:50

that's perhaps a less psychologically

19:52

demanding notion than having a

19:55

mental representation of the goal in that I

19:57

think it's perfectly sensible and plausible to say

20:00

you know, even of a developing organism that

20:02

it needs to decide which pathway to

20:04

go down. So developmental

20:07

regulatory networks can make decisions, chemical

20:10

networks can make decisions. Yeah, I mean,

20:12

you know, some people might insist that

20:15

the literal meaning of the word decision is a

20:17

conscious decision made by human. It's sort

20:20

of inherently mental. Yeah, right. Yeah, but

20:22

I don't see that. I don't see

20:24

that that's either the standard

20:26

way that the word is used in science these

20:28

days, or has anything

20:30

particular to recommend it sort

20:32

of theoretically. So I'm quite happy for

20:34

decisions to be a pretty broad thing. I'm

20:37

very happy with the idea then that, you

20:40

know, all organisms, including microbes

20:42

make decisions and maybe decisions

20:44

are made at the level

20:46

of regulatory networks and, you

20:48

know, subcellular levels too. But

20:51

if so, then I would say that

20:53

decision making and having a mental representation

20:55

of the goal come apart.

20:59

Even that the former is taxonomically widespread, the

21:01

latter isn't. So if

21:03

we can circle back a little bit

21:05

to the concept of plasticity and flexibility,

21:08

for I'd

21:10

say most practicing evolutionary

21:12

biologists, the capacity

21:15

for being

21:17

flexible and whether it's

21:19

at the level of development in response

21:22

to some environmental cues or in the

21:24

context of behavior changing, say in the

21:27

presence and absence of a predator, a

21:30

lot of those types of responses

21:32

are captured currently under the sort

21:34

of framework of phenotypic plasticity. And

21:36

so just based on your

21:39

description, it seems that being

21:41

plastic and flexible

21:44

is inherent to being

21:47

agential in your response. Yeah, I would

21:49

agree. In all of the senses of

21:52

agency, in all of the four senses

21:54

that I've operated, I've distinguished. So are

21:56

they so intertwined that they are essentially

21:59

capturing? the same phenomenon or can we

22:01

sort of decouple them from one another?

22:04

Yeah, I know. I mean, it's tricky. I

22:06

mean, I do think that, you

22:09

know, the notion of phenotypic plasticity and

22:11

behavioral plasticity, that being a subset of

22:14

phenotypic plasticity, you know, given that not

22:16

all phenotypes of behaviors, you

22:19

know, captures a lot of what people

22:21

who talk about agency are

22:23

trying to say, and probably in a

22:26

more sort of acceptable way

22:28

to many practicing scientists. And I think

22:30

that's exactly the source of some of

22:32

the opposition to the language of

22:35

agency that one finds these days

22:37

where people say, look, you're just using

22:39

this mystical-sounding term, which the

22:41

cash value of which is something that we could say

22:43

anyway. Right. Now, you see,

22:45

I think that's a valid criticism of some

22:47

projects that involve the notion of agency, but

22:50

not of mine. In

22:53

some theorists, including some of

22:55

my colleagues in philosophy of

22:57

biology, seem to have convinced

22:59

themselves that there's a deep

23:01

and interesting sense in which

23:04

some organisms or maybe all organisms

23:06

and maybe other biological entities

23:09

too, are agents. And

23:11

their task is to figure out what that sense is,

23:14

subject to the constraint that it

23:16

better be interesting, and better not

23:18

just reduce the behavioral plasticity. But

23:21

that seems to me an odd way to proceed. So

23:24

that's why I'm one of the people who's

23:26

somewhat skeptical about the notion, despite having worked

23:28

on it myself. But my project

23:30

is a somewhat different one in that

23:33

I was, in writing my books,

23:35

specifically interested in a mode

23:37

of thinking and speaking

23:40

that is peculiar to evolutionary

23:42

biology, I think, which

23:45

involves deliberately

23:47

anthropomorphizing for

23:49

the purposes of achieving evolutionary insights.

23:52

And one example of which is

23:54

that the practice of taking

23:57

the adaptive function

24:00

or rationale of some phenotype,

24:03

but particularly a behavior, but not

24:05

necessarily, in the sense that the

24:07

reason why it's evolved. And then

24:09

treating that as if it was the

24:11

organism's own goal, a

24:14

literal goal mentally represented, even when

24:16

it really isn't. And

24:18

I give many examples in my book

24:20

of this practice actually at work in

24:22

evolutionary biology, manifest by the use of

24:24

intentional language. So when we say things

24:26

like that the female rat kills its

24:28

injured young, because it knows that they

24:30

won't survive, it doesn't want to waste

24:32

resources on them, for example, there

24:35

we explicitly employ, you know,

24:37

vocabulary that has its natural home in

24:40

relation to conscious deliberate human behavior, by

24:43

way of giving an evolutionary explanation, or an

24:46

adaptive explanation of the infanticidal

24:48

behavior of the female rat.

24:51

And so that was the phenomenon I was

24:53

specifically interested in the use of intentional vocabulary

24:57

in an evolutionary context. And

25:00

so I took my project, I mean, I

25:02

approached it very much in the spirit of

25:04

a kind of anthropologist of science, you know,

25:07

my starting point was, look, there is this

25:09

pre existing practice out there in the scientific

25:11

literature, that's really unusual and

25:13

philosophically puzzling. And what

25:16

on earth could be the point of talking

25:18

that way, given that it seems to willfully

25:20

invite a confusion of proximate and ultimate? And

25:23

given that it seems to just be a

25:25

form of anthropomorphism, could it really be

25:27

doing any scientific work? That was

25:30

the question I started from. And

25:32

I think of that as a somewhat

25:34

different project, indeed, an orthogonal project, to

25:36

the one that many people, you know,

25:39

both biology and philosophy, who talk about

25:41

agency are ultimately engaged in. So

25:43

I want to push back a little bit, because you

25:46

made a great case, and this adaptationist

25:49

program has been and is pervasive in

25:52

biology. But there's this particular word that

25:54

I hear you using that I'd like

25:56

you to address, and it's mental, right?

25:59

So An important

26:01

qualifier of agency

26:03

that you write about is integration.

26:06

So it's not just the

26:09

potential to perceive the problem or to sort

26:11

of think about the problem, but then the

26:13

system does something about it. So can

26:16

you expand the definition of mental? And

26:19

more importantly, it seems to me to

26:21

be leaning on mental as

26:23

the qualification of agential and justification

26:25

of how to think about that.

26:27

That seems akin to the anthropomorphizing

26:30

and adaptationist perspective. It's a

26:32

move that allows you to

26:34

do other things, but I

26:37

don't know. I guess I'm just not on board that

26:39

mental is something special

26:42

because you have to have integration. And as Art

26:44

was saying a minute ago, there's a lot of

26:46

different systems and even elements of systems that integrate.

26:49

Yeah, no, I mean, I agree that

26:51

integration and mentality are not

26:53

exactly the same thing, certainly.

26:55

So I mean, what I'm getting

26:57

at when I use the term mental is

26:59

that just the assumption that

27:02

psychological descriptors like believes,

27:06

wants, tries, apply

27:09

literally in the human case and

27:12

quite possibly in other non-human

27:15

cases too. But

27:17

don't apply literally in all cases

27:19

where there are sort of hallmarks

27:21

of agency in the sense of

27:24

flexible behavior, plasticity,

27:26

decision making, information

27:28

integration, all of that. Now,

27:31

I mean, that move could be

27:34

contested, I agree. That's not self-evidently

27:37

true. But I think

27:39

that's the natural default assumption that this is

27:41

the home of this vocabulary. It's

27:44

really weird to my mind to think

27:46

that the psychological language of

27:49

that sort literally characterizes what an

27:51

earthworm does. I

27:53

just don't think it's true. And

27:55

that's fair, but I think that's where I'm

27:57

coming from. I mean, to just sort of…

28:00

distinguish these forms of agency using

28:02

human cognition as

28:05

sort of a frame for how we think about all

28:07

of these other things. I sort of approach it from

28:09

the other direction. Life as a

28:11

complex system, one of its manifestations ends up

28:13

being the human brain, but

28:15

even the vertebrate immune system can learn

28:17

in an exquisite way that

28:19

has nothing to do with how nerves

28:21

learn. I mean if we had a

28:23

better grasp on what consciousness was and

28:25

the interplay between consciousness and mental,

28:29

maybe I'd feel a little bit more comfortable, but because those

28:31

are such abstractions too, I don't know, it

28:33

feels like a strange place to start. Right,

28:36

I see. Yeah, I mean why

28:38

do I start in that place? You see, I

28:40

mean I entirely agree with you that, for example,

28:42

the adaptive immune system learns. Yeah,

28:45

that seems that seems absolutely clear. But

28:47

why do I start with the mental?

28:50

I mean I think it's because my

28:52

specific interest is in this anthropomorphizing tendency,

28:55

or at least in that book,

28:57

that's the thing I'm specifically interested

28:59

in, is this anthropomorphizing tendency within

29:01

evolutionary biology. And my overarching question

29:03

in that book was, look, is this tendency

29:06

to use the language of intentional

29:08

psychology in an evolutionary context

29:10

in order to cause adaptive

29:13

explanations, is there any

29:15

real good reason to do that? Or

29:18

is it just a sort of casual

29:20

anthropomorphic bias? Does it pick out some

29:23

genuine class of phenomena? Or is it,

29:25

does it really just more reflect the

29:28

fact that the investigators are humans

29:30

themselves and haven't entirely always managed

29:32

to shed themselves of their anthropomorphic

29:35

bias? But in terms

29:37

of thinking about the origins of agency

29:39

itself and the origins of intentionality

29:42

and consciousness, I mean, I

29:44

think I agree with you

29:46

that precursor forms of all

29:48

of these things are found

29:50

in a much simpler biological

29:52

systems and that somehow

29:54

or other, you know, the

29:56

evolutionary process gave rise to creatures

29:58

whose cognition was sophisticated. sophisticated enough,

30:01

for whatever reasons exactly, that

30:04

they could actually mentally represent

30:06

their goals. Yeah. So Samira,

30:08

I gathered from your book that you

30:11

sort of slot these different ideas about

30:14

sort of levels of agency into what you

30:16

call type one and type two. And

30:18

type one is about organismal agency

30:20

itself, which I think, you

30:22

know, now many biologists are starting to hear

30:24

about and having strong reactions to.

30:27

Type two is this idea

30:29

that the process of evolution

30:32

itself and natural selection can

30:34

be agential and or

30:36

that it may appear to be

30:38

agential because of the sort of

30:40

psychological and intentional language that's used.

30:43

And I would say like it feels like

30:46

your book is largely, you know, in

30:48

the end dismisses this idea that type

30:50

two agency is important and actually captures

30:52

something real that's happening in biology and

30:54

that that we're sort of, in

30:57

a sense, taken for a ride by

30:59

the language that we use to describe

31:02

type two agency. Yeah, no, that's

31:04

exactly right. So it was, yeah, my sort of

31:06

simple one line argument is that

31:08

type one good in certain cases,

31:11

scientifically justifiable, despite being

31:13

anthropomorphic, type two,

31:16

generally not good, anthropomorphic,

31:18

but bad. So type two in

31:20

at least the paradigm sort

31:22

I have in mind involves, you

31:24

know, thinking of the evolutionary process,

31:26

as you say, or the process

31:28

of natural selection as a form

31:31

of agency, if you like. And Darwin

31:33

himself was the first to do this

31:35

in a number of passages in the

31:37

origin of species, where he says natural

31:39

things like natural selection is daily and

31:42

out, he's scrutinizing the world, looking

31:44

for any advantage and preserving

31:47

what's good, eliminating what's bad. So

31:49

that is quite deliberately using this

31:52

intentional language to characterize the

31:54

process of natural selection itself.

31:57

Now, Darwin went on to say, look, these are

31:59

just metaphors, but don't worry about it. It's

32:02

not a big deal. And I think

32:04

in fact, he was cavalier on that point. Because,

32:06

I mean, one of the

32:08

things that people didn't realize in the didn't

32:10

understand in the earliest days

32:12

of post Darwin was that the

32:15

process is, of course, mechanistic and

32:17

doesn't have any foresight, there is

32:19

no goal sorts, which evolution is

32:22

striving or anything like that. And, you

32:25

know, any attempt to think otherwise

32:27

is really is to misunderstand Darwin's

32:30

achievement. But nonetheless, I do

32:32

think that sort of vestiges

32:34

of that type two form

32:36

of agential reasoning and thinking

32:39

are still alive today, in

32:41

fact, because you see, I think

32:44

that we very often treat the

32:46

simplest sort of natural selection as

32:48

if it was paradigmatic. And

32:51

so by I mean, the simplest sort of natural

32:53

selection is just one we've got a number of

32:55

variants in a population in a fixed environment, with

32:58

no frequency dependence. And

33:01

the evolutionary process when driven by

33:03

natural selection simply involves fixation of

33:05

the best variant and elimination of

33:08

the worst ones and the, you

33:10

know, monotonic increase of mean

33:13

population fitness until it reaches

33:15

a maximum so pushes the

33:17

population up the adaptive landscape, all

33:20

ideas that are, you know, have their place.

33:22

And are useful, but rest on, you know,

33:24

that's only the very simplest case. And,

33:26

you know, it's all the evolutionists know,

33:29

those things hold true under restrictive model

33:31

assumptions that almost never apply. And in

33:34

the real world. And so I think

33:36

the tendency to adopt that

33:38

language leads us to

33:40

think that the simplest form of

33:42

natural selection as a sort of

33:45

simple optimization process is

33:47

the paradigm example. And

33:49

obviously people who really know the science

33:51

know that that isn't true. But

33:54

nonetheless, I think a vast number of

33:56

people who sort of know a bit about evolution and

33:58

natural selection, don't understand. understand that that

34:00

isn't true. And the use

34:02

of this intentional language, thinking of

34:05

the evolutionary processes, you

34:07

know, either driven to achieve some

34:09

goal or in any case as directed

34:12

in some sort of way or as pushing

34:15

populations up landscapes by the steepest

34:17

path, that sort of thing. Those

34:20

mistaken notions go hand in hand, I

34:22

think, with the use of these

34:25

anthropomorphic intentional descriptors to characterize the

34:27

process of evolution by natural selection.

34:29

So that's why I say type

34:31

too bad. But I want

34:33

to say type one, which involves

34:35

thinking of an actual evolved entity,

34:38

typically an organism, but possibly a

34:40

group or maybe even a gene

34:43

as akin to an agent, and

34:45

as trying to achieve a goal can

34:47

be in certain cases, a

34:50

useful mode of description.

34:52

I try and make this argument

34:54

that although anthropomorphic, it does genuine

34:56

work in that it picks out

34:58

a real phenomenon that, you

35:00

know, certainly could be described otherwise, but that

35:03

there is a real reason to describe this

35:05

way. And that's when I start

35:07

talking about integration and

35:09

absence of internal conflict as

35:12

being a prerequisite for thinking of

35:14

an organism as agent

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36:01

Yeah, so this is a good place to

36:03

really just ask the question, are genes

36:06

agents? Especially in

36:08

the context of thinking about transposable

36:11

elements and gene

36:13

for altruism or other

36:16

kinds of inter-genomic conflict, it

36:18

seems like our language

36:21

that we would use and our

36:23

theory to describe why there would

36:25

be selection for or against the

36:27

spread of a gene within a

36:29

genome fits within this concept

36:31

of an agent. Yeah, no, I

36:33

agree with that. And I think, I mean,

36:35

the broad story I'm trying to tell about

36:37

agency and the circumstances in which it's a

36:40

valid concept or useful concept

36:42

at least to employ in an

36:44

evolutionary context includes the

36:47

famous selfish gene notion of

36:49

Dawkins. Although I do

36:51

think of the many different sort

36:54

of lines of rationales and lines of

36:56

reasoning that Dawkins used

36:58

to support the selfish gene notion, the

37:01

one that I'm interested in is the one you

37:03

allude to, that is where

37:05

there's inter-genomic conflict, that it

37:08

makes most sense to think of a gene

37:10

as trying to achieve an outcome by

37:12

means of doing something that in this case

37:15

harms other genes within the same

37:17

genome. By now, a familiar

37:20

point and one that I think

37:22

Dawkins himself made in his

37:24

later work that the sense in which the

37:26

gene is the unit of selection where

37:29

there's inter-genomic conflict is really rather different from

37:31

the sense in which the gene is always

37:33

the unit of selection. I mean,

37:35

the latter is true in a sense,

37:38

but it's in a different sense that

37:40

we talk about genes as

37:43

the entity is being selected where

37:45

there's conflict within a single genome.

37:48

So I sometimes capture that by contrasting

37:50

the process of gene level

37:53

selection with a genetic

37:55

perspective on selection process that take

37:57

place at a different level. So,

38:00

Mir, can I ask you to tie back

38:02

genes as agents to your four types

38:05

of agents that we started with? Into

38:07

which category would genes fit for you?

38:10

Well, I would say really,

38:12

literally speaking, none of

38:14

them, because, I mean, you know, DNA is a

38:17

pretty inert molecule. I mean, obviously,

38:19

the more we know about how genes

38:21

work and, you know, how gene expression

38:24

works, you know, the more the

38:26

situation is a little bit complicated. But

38:28

I would still say that if we're

38:30

asking which of those agent concepts literally

38:33

characterise genes, I would be inclined to

38:35

say none. But nonetheless,

38:38

it can be useful to use one

38:40

or more of those concepts metaphorically

38:42

to describe genes for

38:45

certain projects. And I

38:47

think that, you know, what Dawkins and

38:50

Dennett and fellow travellers are doing is

38:52

basically taking one of those notions of

38:54

agency, namely the intentional agency one, and

38:57

using that notion, applying

38:59

that notion to genes, but in

39:01

an overtly, self consciously

39:03

metaphorical sense. I've always

39:05

been puzzled why people are so horrified by that.

39:08

I think it makes extremely good

39:10

sense to me. So long

39:12

as it's used carefully. Yeah, it's funny, I

39:15

feel relatively comfortable with, you know, the idea

39:17

of genes as a level of selection.

39:21

It's hard for me to conceive, and you

39:23

sort of said this, but it's hard for

39:25

me to conceive of genes as agents, because,

39:28

you know, to me, agency itself comes from

39:30

networks, doing computations and

39:33

making decisions. And of course, genes

39:35

don't contain those networks, right? They

39:37

can be a component of another network.

39:41

But they're just a part that is

39:44

participating in a greater whole that maybe

39:46

has agency, but that don't

39:49

have agency themselves. What do you think of that? Yeah,

39:51

I mean, I think I would agree. But

39:54

again, that's because you're using the terms

39:56

in this literal way. I mean,

39:58

I think that what say that

40:00

genes don't have themselves any

40:02

of the characteristic hallmarks of

40:05

agency, I think is true, but

40:07

I also think that that's compatible

40:10

with heuristically treating them as if

40:12

they did in order to further

40:14

our understanding of the evolutionary

40:16

processes, of certain evolutionary processes.

40:20

And so, I mean, if you

40:22

think of how people struggle to

40:24

understand the phenomenon of intra-genomic conflict

40:26

before they have the selfish gene

40:28

paradigm, which was really invented

40:30

for other reasons, really. Dawkins was

40:33

originally thinking about things like altruistic behavior

40:35

and so on. But, I

40:37

mean, I think, once you

40:39

have that idea, then

40:41

you can immediately see how to

40:43

make sense of this otherwise extremely

40:46

puzzling phenomenon. I think it'd be

40:48

useful to to give an example of

40:50

this type of inter-genomic conflict. Is there a

40:53

particular one that you really like? No

40:55

particular one. I mean, there are thousands

40:58

of these examples, but say, for

41:00

example, the phenomenon of cytoplasmic male

41:02

sterility in flowering plants,

41:05

where you have a gene within

41:07

a mitochondrion that causes

41:10

its plants to cease making

41:13

pollen, which is a really odd

41:15

thing to do. You know, the

41:17

plant, in some cases, the plant actually grows

41:19

stamen, so all

41:21

intents and balances in order to make

41:23

pollen, but then those

41:25

mitochondrial genes frustrate the

41:29

stamen from actually doing the thing that they're meant to do.

41:32

I mean, that's from the phenotypic perspective of the

41:34

whole organism. You look at them, what on earth

41:36

is that about? What's the

41:38

rationale for that? Is this an

41:40

exception to the theory of evolution or

41:42

something like that? How could it be?

41:45

But it makes perfect sense when you

41:47

realize that the mitochondrial genes are only

41:49

transmitted, but only down the

41:51

female line, so investing in pollen is a waste.

41:53

Now, that's the sort of case I have

41:56

in mind, where I say that there's a threat to the

41:58

idea that the whole organism is a waste of pollen. is

42:00

agent like because it

42:02

doesn't have if you like the

42:04

sort of phenotypic integration the behavioral

42:07

integration that one would

42:09

need in order to apply

42:11

the psychological language metaphorically

42:13

to it. So what

42:15

is it that the plant is trying to achieve? You

42:18

know is it trying to self fertilize

42:20

or not? Well on the

42:22

one hand yes because you know a lot of its

42:24

apparatus is geared to that but on the

42:26

other hand no because they don't make any problems. Yeah

42:29

and this this might be a bizarre thing to

42:31

say Samir to the author of a book on

42:33

levels of selection but you know those

42:35

kinds of arguments in the way you make

42:37

a completely fantastic case and there's no question

42:39

that these conflicts are true but

42:42

because biology is organized across

42:44

levels to sort of use

42:46

these as ways to understand

42:48

organisms you know is

42:50

it possible that these are great examples of genes

42:53

as parasites or something

42:55

different and the rest are a large

42:57

fraction of the genome and all its

42:59

regulatory components they aren't so

43:01

much parasites as commensals or mutualists I mean

43:04

you know to use the parasitic

43:06

argument to be foundational for everything

43:09

organismal I have always viewed that as an overreach.

43:11

Yeah I see I mean I think I

43:14

think I'm on board with the idea that

43:17

we can have genomic parasites and

43:19

that that actually accurately captures quite

43:22

a lot of cases of intragenomic conflict so

43:24

what's the thing that you think is overreach?

43:27

You know your arguments about why not

43:29

make pollen I mean that's sort of framed

43:32

in an adaptive context at the

43:34

organismal level where another explanation is

43:36

some type of frozen accident happened

43:38

in the past that produces this

43:40

lineage that you know the system

43:43

it's still alive natural selection is still going to

43:45

operate as natural selection

43:47

does as life does it makes the best of

43:49

what it has but to use that to

43:52

sort of expand that out to a full

43:54

way to understand the evolutionary process that

43:56

seems to be ambitious. Well you see what

43:58

I'm trying to do is to make an argument that

44:01

says, look, it's because phenomena

44:03

like that are relatively rare,

44:06

that the language of agency has

44:08

any purchase at all in

44:11

an evolutionary context. In

44:14

that we have this sort

44:17

of general philosophical line of

44:19

argument which says something like, look,

44:22

talk of intention, goal,

44:24

belief and desire is

44:27

really only appropriate when there is

44:29

a kind of rational integration indicative

44:33

of the fact that in cases, say,

44:35

of multiple personality then we really

44:37

struggle to say what on earth it is that a

44:39

person, you know, whether there really is one

44:41

person there at all or what it is that they believe

44:43

or what it is they want. We can't say it. You

44:46

know, the psychological language just ceases to

44:48

apply in that sort of case.

44:50

And what I'm arguing is that we have

44:53

a kind of biological analog of that in

44:55

the sense that if there were

44:57

too much intragenomic conflict, the

45:00

phenotypic consequences of

45:02

that would make it impossible for

45:04

us to think of the organism

45:07

as trying to achieve anything and

45:09

it would be impossible to metaphorically

45:11

characterize it as, you know,

45:13

aiming at some goal or other,

45:16

if you like. So I guess

45:18

I'm suggesting that this psychological integration

45:20

and rational integration is a necessary

45:22

condition of applying psychological language literally.

45:25

And the biological phenotypic integration

45:28

that one gets when intragenomic conflict

45:30

is effectively suppressed, as in the

45:32

vast majority of extant

45:35

organisms, is the necessary condition

45:38

of applying that psychological

45:40

language metaphorically to organisms.

45:43

And we see those types of conflicts not

45:45

only inter-genomically, but we

45:48

also see those conflicts, for example, between

45:51

the goals of males versus females and

45:53

how those play out, you know, in

45:55

terms of toxic sperm that may help

45:57

an individual male against set of conditions.

46:00

competitors but actually does harm to

46:02

the female. And so it's

46:04

the same conceptual framework that we're applying

46:06

in those cases as well. But in

46:08

that case, the conflict is between genes

46:10

in different organisms, I guess, or between

46:13

two separate organisms. Yeah. Or

46:15

yeah, two sexes, I guess. So

46:17

for that very reason, I think it would

46:20

be a mistake to think that a mating

46:22

pair was agent-like, if

46:24

you see what I mean, or to talk about what

46:26

the mating pair is trying to achieve. Obviously,

46:29

their interaction is partially mutualistic.

46:31

You use the term integration,

46:33

but also, I think

46:35

along with that, you also use the

46:37

term unity of purpose. And

46:40

so I think that's

46:42

the outcome of the integration, that

46:44

these actions are towards the goal,

46:46

towards this purpose, and it requires

46:48

this integration or coordination to achieve

46:51

that goal. Absolutely. But

46:54

I mean, one thing I do stress is that it's a matter

46:56

of degree. So there

46:58

are local violations of this unity of

47:01

purpose do certainly exist, and there are

47:04

relatively common in intra-genomic conflict, as you

47:06

guys all know, has been found in

47:08

almost every tax of where it's been

47:10

looked for. But nonetheless,

47:12

I do think that those

47:15

violations are necessarily local, if you

47:17

like, in that if you had

47:19

too much breakdown of unity

47:21

of purpose, you probably wouldn't

47:23

actually have an organism at all. And

47:26

you certainly wouldn't be able to think of

47:29

it as akin to a rational agent trying

47:31

to achieve an end to which its various

47:34

phenotypes all contribute. Yeah.

47:37

So I have a related question here that

47:39

builds on what you just said about the

47:41

idea of whether a mating pair

47:43

has agency, and that's just to

47:46

expand, it's also kind of a

47:48

levels question, but going in the other direction,

47:51

and thinking about groups of organisms and

47:53

whether or not they show agency and

47:55

how that relates to their evolutionary processes.

47:57

And like at some level, It's

48:00

easy to think of groups

48:02

that almost certainly do have

48:05

agency like U.S. Social Insect

48:07

Colonies in which many of the individuals are

48:09

closely related to one another and they act

48:12

essentially like a super organism.

48:15

But what about, say, interspecific

48:17

groups which occur

48:19

all the time in the wild? And I'm

48:21

thinking of things like, so there's a

48:24

retired professor here at Montana, Eric

48:26

Green, whose work some on communication

48:29

among different bird species in between

48:31

mammals and birds in forests. And

48:34

you may be aware of some of this work,

48:37

but there are waves of communication about predators

48:40

that travel between the members

48:42

of these interspecies groups. And

48:45

that really define a lot about where

48:47

these individuals go and, you know, when

48:49

they're feeding and when they're being fearful

48:51

and paying attention to predators

48:54

that are coming by. So would you

48:56

say that a large multi-species

48:58

group like that also shows agency

49:00

in some way? I

49:02

guess I would hesitate to say that,

49:05

although it might be true. I mean, I don't

49:07

think you could rule it out, definitely. But I

49:09

think the reason I would hesitate to say it

49:12

is just for the same reasons

49:14

that people have always been

49:16

reluctant to think of such a

49:18

multi-species group as an adapted unit

49:20

itself. I mean, clearly

49:22

the organisms in it are adapted and

49:25

in part they're adapting themselves to

49:28

the social environment that

49:30

they find themselves in, which includes

49:32

living in these multi-species communities. But

49:34

that's still different from saying that,

49:37

you know, the multi-species group itself is adapted.

49:39

I mean, this goes back to the point

49:41

that George Williams made way back in 1966

49:43

in his book Adaptation and Natural

49:46

Selection, where he

49:48

wanted to say, look, distinguish between a

49:51

group adaptation and a fortuitous

49:53

group benefit. We

49:55

can't really easily conceive of

49:58

an evolutionary process by which species

50:00

groups could have come to be

50:02

the adapted units themselves. Although,

50:04

I mean, it's possible. Because their

50:07

evolutionary interests are not closely enough

50:09

aligned consistently, right? Right,

50:11

exactly. Yeah, because we would think of

50:13

the evolutionary process as taking

50:16

place on the separate lineages

50:18

within those multi-species groups. And

50:21

to think of, you know, the multi-species group

50:23

as really part of the environment relevant

50:25

to the evolution of the, of

50:28

the populations, the

50:30

con-specific populations that

50:32

make them up. So, I mean, I think

50:34

that's the reason that I would hesitate to

50:37

apply the language of agency here. But

50:39

I would want to say that it

50:41

is ultimately an empirical question that, I

50:43

mean, there are people who, who take

50:45

seriously the idea of community level, natural

50:47

selection, where the community can

50:49

be a multi-species aggregate

50:52

of some sorts. But

50:54

I think if we sort of relax

50:57

the definition of integration, and I think

50:59

another way to capture integration is, is

51:02

to call it some degree of cooperation,

51:04

whether it's at the

51:06

whole organism level, you have to

51:08

have these coordinated responses. I

51:11

could also see cooperation, you know,

51:13

obviously within a group, but if

51:15

we kind of relax a little

51:17

bit what we, what we mean

51:19

by cooperation, I could see it

51:22

also applying to the examples that Art was

51:24

giving of, you know, mixed species,

51:26

because ultimately then

51:28

through that cooperation, what

51:31

emerges is this unity of purpose.

51:33

They all have the same interests

51:35

in avoiding the predators. Yeah,

51:37

I mean, I think that that's possible,

51:40

presumably that, that overlap of evolutionary

51:43

interests will be kind of

51:45

contingent, if you like, and

51:48

could break down. Especially if some of

51:50

those bird species are

51:52

also competitors and don't

51:54

have, under other

51:56

contexts, aligned differences. So yeah,

51:59

I think that kind of context

52:01

dependency probably plays an important role.

52:04

Right. But and of course, even if

52:06

the groups were not conspecific, you know,

52:08

comprised of a single bird

52:10

species, then I mean, it would still

52:12

be problematic to think of the

52:14

group of such a group as, as

52:17

an adapted unit automatically. I mean,

52:20

because I mean, I again, I would be happy

52:22

with that if a demonstrable

52:24

process of group level selection

52:26

has shaped the evolution of

52:28

some phenotype. So then

52:30

we get back into this question of, you know,

52:32

what sort of selection process is necessary

52:34

in order to produce an

52:36

adapted group, as opposed to

52:38

a group of adapted individuals, which

52:41

I've always thought is a helpful

52:43

framing that distinction that the Williams

52:45

made. Yeah. Do you

52:48

think that groups of humans, given

52:50

what we were just talking about with individuals within

52:53

a bird population, would groups of humans count as

52:55

a gentle or do you have to have the

52:57

same qualifiers for us? I

53:00

mean, I think potentially. So, I

53:04

mean, if you think, for example, of not

53:06

a group of humans, like a tribe

53:08

or a, but something more smaller, like

53:11

say committee, or something like that,

53:13

many people in philosophy of social

53:15

science take very seriously the idea that groupings

53:18

like committees can quite literally, literally

53:21

be agent like, we can talk

53:23

literally about what the committee believes

53:26

and decides and did. And

53:28

that certainly is manifested in the way we talk

53:30

ordinarily, we say, yeah, you know, the

53:32

firm raised prices, the committee decided what

53:35

to do, we do use this

53:37

language of choice and decision and

53:39

belief. And there's

53:41

a standard psychological language to

53:45

groupings of humans in

53:47

certain contexts. And

53:49

so I'm happy with the idea that that

53:51

could indeed be literally true. Yeah. And certainly

53:54

that it could be heuristically useful to think

53:56

that way. But again, only I mean, only

53:58

of course, under certain and circumstances. I mean,

54:00

if the committee couldn't come to a decision

54:02

or if, you know, one person on

54:05

the committee said, said X and the other said

54:07

not X, then of course we

54:09

couldn't sensibly talk about what the committee think

54:12

about the topic. So

54:25

we want to transition more directly

54:27

into the interplay between agency, your

54:29

ideas about agency and classic evolutionary

54:31

theory. So to put

54:33

this in an explicitly evolutionary perspective, and

54:36

we could stick with humans or we

54:38

can go in any direction you'd like,

54:41

is there evidence or do you expect that

54:45

in nature there is heritable variation

54:47

for agency? I think as

54:49

you were talking about agency there, it wasn't obviously

54:52

in the adaptive, the classic evolutionary adaptive

54:54

sense, but in natural

54:56

populations, what do you think about heritable

54:58

variation for agency? Well, I

55:01

suppose I would say, you know,

55:03

agency depends on, in any

55:06

of the senses that we've been discussing, it depends on

55:08

the possession of

55:10

certain phenotypes, such

55:12

as, you know, information processing,

55:15

sensing the environment, cognitive

55:18

abilities of various sorts. And

55:20

there must of course have been heritable variation

55:23

in all of those for them to evolve,

55:26

whether there still is now, is perhaps

55:28

slightly different question. But I still think

55:30

of that, I think of the heritable

55:32

variation as being with respect to phenotypes

55:34

that prerequisites

55:36

of or building blocks of

55:39

agency, rather than thinking of

55:41

agency itself as being something that

55:44

is itself a phenotype. Huh,

55:47

that's interesting. So you

55:49

wouldn't call agency itself a trait for

55:52

which there would be heritable variation? No,

55:54

I think it would, I think it's

55:56

almost at the wrong level itself to

55:58

be a trait. But

56:01

that raises a sort of an

56:03

interesting problem for me because, you

56:05

know, if we go back to Fischer, he

56:08

talked a lot about, in his

56:10

fundamental theorem, he talked about essentially

56:13

heritability for fitness. And

56:15

yet fitness is also made

56:17

up of all of these other components.

56:21

And so I see a lot of parallels there,

56:23

but yet in our models we don't have a

56:25

problem thinking about, you know, as long

56:27

as there's additive genetic variance for fitness.

56:30

And maybe that's because we're thinking about

56:32

it at a population level as opposed

56:34

to individual level. Yeah,

56:36

no, I mean, I think that's

56:38

an absolutely fair point. Yeah, so

56:40

if we take it that, you

56:42

know, fitness ultimately depends on the

56:44

vital rates, birth and the

56:47

death rates, and that

56:49

those that contribute to many, many different

56:51

phenotypes, of course, will contribute to the

56:54

vital rates of an organism of

56:56

any sort. So how is it

56:59

then that we're happy to talk about

57:01

heritability of fitness? Yeah, I

57:03

mean, I see the point. You're

57:05

saying, well, why by parody of argument, why

57:07

can't we talk about agency

57:10

as a heritable trait or

57:12

something exhibiting heritable variation? I

57:14

mean, in part, maybe the difference is in

57:16

part just that it's not so easy how you would

57:18

measure it. Yeah, I agree. Right.

57:22

Yeah, I mean, so do we find

57:24

an organism that's got more of it than

57:26

one that's got less of it? I

57:29

mean, how would you? Yeah, right. What does that

57:31

mean to see a population in which there's a distribution

57:33

of agencies, right? Yeah, no, that's

57:35

right. But nonetheless, I mean,

57:37

if we take it that in contemporary

57:39

biological populations, then we do see agency.

57:43

And presumably, if we go far enough back

57:45

in their evolutionary ancestors, then we don't

57:48

find agency or at

57:50

least only precursor forms of it, then some

57:52

evolutionary process must have taken place to produce

57:55

it. So in that sense,

57:57

it's got to be an evolved attribute. But

58:00

I'm still a little bit uncomfortable with thinking

58:02

of it itself as a trait, in

58:05

part just because it seems hard

58:07

to measure and to depend

58:09

on too many other things that have a

58:11

sort of clearer status as traits, let's say.

58:14

I agree, and I think that

58:16

actually is a point of argument

58:18

between me and Marty and Art,

58:20

that I recognize that organisms

58:23

behave and act

58:25

agentially, but I see

58:28

it more as sort of an

58:30

epiphenomenon. It's an emergent property that

58:32

probably has evolved over time, but

58:34

as an emergent property, it's something

58:36

that in of itself is really

58:38

difficult to, as Art was

58:41

saying, like, you know, plot

58:43

a distribution of agency in

58:45

a population. Absolutely.

58:47

I mean, I suppose someone might

58:49

make the argument, well, you

58:51

know, maybe it's fixed within

58:53

every species of something, so

58:56

it doesn't show any distribution, you know,

58:59

within a human population, for example.

59:01

But then it wouldn't be able to evolve. No,

59:04

but maybe it's evolved already. Maybe it's, you know,

59:06

it's like bipedalism or something. Interesting.

59:09

Well, it's a good thing in evolutionary biology, we

59:11

don't have any other great big emergent words like

59:13

gene and fitness and things like that. So,

59:18

Samir, you spent considerable

59:21

time or space in the

59:23

book focusing on Alan Grafven's

59:25

formal Darwinism project. And I

59:28

think the goal of Grafven is to

59:30

move away from population

59:32

measures of fitness to thinking

59:34

about individual optimization, in

59:37

particular as it relates, you know, to this

59:39

idea of agency. It

59:41

seems like Grafven's goals are

59:44

to separate selection

59:47

from individual optimization. But

59:50

then I think you made this

59:52

point in the book that natural selection may

59:54

not always then lead to

59:57

adaptation. And this

59:59

seems like a very counter... counterintuitive conclusion,

1:00:01

could you help me understand how

1:00:04

those get separated from one another?

1:00:07

How natural selection and adaptation get

1:00:09

separated from one another. Yeah, and

1:00:12

this idea that selection alone

1:00:14

can, in some cases, not

1:00:16

lead to adaptation. Right, yeah.

1:00:19

No, I mean, what I was

1:00:21

getting at with that was just

1:00:23

drawing attention to some points that

1:00:25

are fairly well known among evolution

1:00:27

theorists. Phenomena,

1:00:30

for example, are cyclical dynamics,

1:00:32

right, where even in simple one

1:00:35

locus population genetics then we

1:00:37

can point to certain

1:00:39

circumstances under which equilibrium

1:00:42

will not be attained, but

1:00:44

rather gene frequencies will cycle indefinitely,

1:00:48

never settling down and attaining equilibrium,

1:00:51

even though natural selection is the only force

1:00:53

at work, is the only evolutionary force at

1:00:55

work and it's not a stochastic effect. It's

1:00:58

nothing like that. So I

1:01:00

think simple examples like that seem

1:01:02

to illustrate, you know, fairly clearly,

1:01:04

I think that although we usually think

1:01:06

of natural selection as an

1:01:09

engine for adaptation, as producing

1:01:11

organisms that behave as

1:01:13

if they're trying to achieve some end,

1:01:16

some fitness related end, there's no sort

1:01:18

of logical guarantee that that's got to

1:01:20

be the case. And I think one

1:01:22

sees the same in models of frequency dependent

1:01:24

selection. So if you look in the popular

1:01:27

adaptive dynamics framework for thinking about

1:01:29

that, although the same is true

1:01:31

in other formal frameworks as well

1:01:33

for thinking about frequency dependent selection,

1:01:35

it's not always the case that

1:01:37

natural selection will take the population

1:01:39

to an equilibrium at

1:01:42

which individuals are playing

1:01:44

the optimal strategy conditional on the rest of

1:01:46

the population. I mean, often it

1:01:48

will. And that's the nice case in

1:01:50

which we get an evolutionary

1:01:53

stable strategy as the outcome

1:01:55

of frequency dependent selection. But

1:01:57

unpleasant, nasty, nonlinear.

1:02:00

dynamics are a cyclical dynamics for example and

1:02:02

other things are possible too as people have

1:02:04

known for a long long

1:02:06

time even though natural selection is

1:02:08

the only force at work.

1:02:11

So I think of I mean what Grafon is

1:02:13

doing as I see it is in part a

1:02:15

response to that situation in

1:02:18

that he's trying to say something like well I

1:02:21

mean can we say something about when

1:02:23

natural selection will be to adaptation and

1:02:25

he thinks that there's got to be

1:02:27

an intimate and tight link between them

1:02:29

and I think intuitively that makes sense.

1:02:31

I mean the whole point of the

1:02:33

theory of natural selection was precisely to

1:02:36

explain to phenomena adaptation and

1:02:38

diversity but then we face

1:02:40

this fact that when explicit evolutionary models

1:02:42

are made you know what obviously went

1:02:45

beyond what Darwin himself was

1:02:47

thinking of we find that

1:02:49

natural selection sometimes seems to result in

1:02:51

something that we can't really call adaptation

1:02:54

in any clear sense. So

1:02:56

then we want to say well can we

1:02:58

point out conditions under which

1:03:00

it will and that I think

1:03:03

of as what Grafon is trying to do

1:03:06

in his project and I think it is very

1:03:08

useful I mean although I don't think all the

1:03:10

moves he makes are ultimately successful

1:03:12

I think it's a conception it's extremely useful

1:03:14

to say look on the one hand we

1:03:16

have the process of natural

1:03:18

selection and on the other hand

1:03:20

we have the phenomenon of

1:03:23

adaptation to the environment and the

1:03:25

one is meant to explain the

1:03:27

other and the fitness concept rather

1:03:30

oddly is sort of features

1:03:32

on both sides of that equation. My

1:03:34

follow-up to that is it seems like

1:03:37

there's another concept here that is somehow

1:03:39

lost in the arguments which

1:03:41

is variation and so I

1:03:43

completely agree in the simplistic models

1:03:46

you know we think about selection

1:03:48

leading to adaptation and

1:03:51

the fixation of a single sort

1:03:53

of variant but whether it's

1:03:56

frequency dependence or density dependence or

1:03:58

just if there's fluctuation

1:04:00

environments, it seems like

1:04:03

one outcome of that is that

1:04:05

no single variant will always be

1:04:08

optimal. And a byproduct of that

1:04:10

is that variation is maintained in

1:04:12

the system, in the population in

1:04:14

this case. And so

1:04:17

I'm just kind of curious where

1:04:19

variation among different

1:04:22

agents or different individuals in a

1:04:24

population falls within

1:04:27

this idea. We touched on

1:04:29

this a little bit that

1:04:31

maybe bipedalism evolves and

1:04:33

then becomes fixed. But I feel like

1:04:36

at some level we

1:04:38

have to incorporate variation into

1:04:40

these ideas. Yeah, no, I

1:04:42

mean, absolutely. I mean, I

1:04:44

think everybody would agree that you need variation in

1:04:47

order for there to be natural selection at all.

1:04:50

If all organisms are the same in

1:04:52

some respect then there's no way that

1:04:54

some variants can out proliferate others. Everyone

1:04:56

has the same variants, if you like.

1:04:58

So that I think is presumably common

1:05:00

ground that variation is necessary

1:05:02

for natural selection. But

1:05:04

I mean, there's also a different question of

1:05:07

whether the outcome of

1:05:09

natural selection will be to

1:05:11

preserve variation or

1:05:13

to eliminate it. Yeah, we think about it

1:05:15

eliminating it usually. Right, we do. We do.

1:05:17

But of course, as you rightly point out,

1:05:19

I mean, if there's,

1:05:22

you know, environmental heterogeneity, temporal

1:05:24

or spatial, or a

1:05:27

host of other complications, or

1:05:30

non additive genetics, for example,

1:05:32

then we can have scenarios where

1:05:35

variation, genetic and phenotypic is in

1:05:38

fact preserved by natural selection or

1:05:40

natural selection can lead to adaptive

1:05:42

diversification, for example, as well.

1:05:45

So I sort of starting,

1:05:48

you know, assumption that natural selection will

1:05:50

reduce variation, although, you know, true, usually

1:05:53

for most, for the simplest

1:05:55

sort of natural selection need not always be

1:05:58

right. So. One

1:06:00

of the issues you brought up is that there's lots

1:06:02

of variation in the environment. And so often

1:06:04

there's not going to be a single genotype

1:06:07

that's optimal. Is

1:06:10

it possible that things like plasticity

1:06:12

and then maybe on top of

1:06:14

that, agential behaviors sort of writ

1:06:16

large are a response

1:06:19

to that and evolved outcome to deal

1:06:22

with that very kind of uncertainty,

1:06:24

right? So there's these systems that

1:06:26

are much more flexible over much

1:06:28

shorter time frames than alleles

1:06:31

and genomes are that

1:06:33

allow organisms to deal with

1:06:35

that kind of uncertainty. To deal

1:06:37

with environmental uncertainty. Yeah. Yeah. Yeah.

1:06:41

So you're thinking of an environment

1:06:43

that changes sort of within

1:06:46

the lifetime of an organism.

1:06:48

Yeah, within or between. But I

1:06:50

mean, either way, the systems that

1:06:53

produce these organisms during development inherently

1:06:55

have this sort of flexibility and

1:06:57

plasticity and agency built into them

1:06:59

to sort of compensate for the

1:07:01

fact that, of course, they're not going to have the ideal

1:07:03

alleles at all, the loci. There's no way

1:07:05

they possibly could, right? No,

1:07:07

I mean, I think that that may well be

1:07:09

true. I mean, of course, I

1:07:11

guess it's not the only possible way that national

1:07:14

selection could adapt a species

1:07:17

to environmental uncertainty. So I

1:07:19

mean, I think that's absolutely

1:07:22

right. But I mean, environmental

1:07:24

fluctuations and heterogeneity and uncertainty

1:07:26

are presumably pervasive in

1:07:29

the living world. I mean, pretty much

1:07:31

all lineages must face those. So then

1:07:33

I guess the question is why some

1:07:35

lineages evolved these agent like

1:07:38

behaviours or capacities

1:07:40

for dealing with them, whereas others

1:07:42

didn't. You know, so

1:07:45

an alternative might be to engage in

1:07:47

some sort of debt

1:07:49

hedging or diversification, for

1:07:51

example, where a given genotype produces lots

1:07:53

of different phenotypes in order

1:07:56

to hedge against the environmental

1:07:58

uncertainty. you know, if

1:08:01

one of those feed types will germinate,

1:08:03

whatever the weather turns out to be.

1:08:05

That well-known idea. So for me, the

1:08:07

question is, can

1:08:09

anyone say something sort

1:08:11

of concrete about the particular

1:08:14

sorts of environments that led to

1:08:16

the elements of agency being the

1:08:18

adaptive response as opposed

1:08:21

to sort of sub-agential things

1:08:23

that one finds more widely? Yeah.

1:08:25

Well, Samir, we've really enjoyed the conversation.

1:08:27

I really appreciate your effort. This has been

1:08:30

wonderful. Oh, it's been great. I mean, I

1:08:33

could talk about this stuff all day, as I'm sure you

1:08:35

guys could too. I

1:08:38

think we all could as well. Excellent.

1:08:41

So we always like to wrap with sort of

1:08:43

giving you a blank page to say anything about

1:08:45

the book or future plans or anything else you'd

1:08:47

like to talk about. Is there anything else you

1:08:50

wanted to hit? Not particularly. I'm currently

1:08:52

writing a book about the concept of

1:08:55

fitness and why it's caused so much

1:08:57

controversy and difficulty and why

1:09:00

it's not as simple a matter as it should

1:09:02

be. Okay. The subject for our next conversation,

1:09:04

please let us know. When that

1:09:06

one comes out, that would be wonderful. Okay.

1:09:09

I will. Hopefully this next year. Yeah.

1:09:11

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

1:09:14

Well, thank you so much. Yeah. Thanks,

1:09:16

Samir. Thank you. My pleasure. Thank you

1:09:18

all. Thanks

1:09:36

for listening. If you like what you hear, let us know

1:09:38

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1:09:40

review wherever you get your podcast. And if

1:09:42

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1:09:45

Right to us at info at big biology.org.

1:09:47

Thanks to Steve Lane, who manages the website

1:09:49

and Molly McGinn for producing the episode. Thanks

1:09:52

also to Dana Della Cruz for her

1:09:54

amazing social media. Katie Shumary

1:09:56

produces our awesome cover art. Thanks

1:09:58

also to the college of public health. University of

1:10:00

South Florida and the National Science Foundation

1:10:02

for support. Music on the

1:10:04

episode is from Poddington Bear and Tierra in

1:10:07

Costova.