Nichola Raihani is a professor of evolution and behaviour at University College London. Her research focuses on the evolution of punishment and paranoia. In this conversation, we talk about the fieldwork she did for her PhD in the Kalahari desert, the evolution of punishment, proximate and ultimate explanations, cleaner fish, and Nichola's book The Social Instinct.
BJKS Podcast is a podcast about neuroscience, psychology, and anything vaguely related, hosted by Benjamin James Kuper-Smith. New conversations every other Friday. You can find the podcast on all podcasting platforms (e.g., Spotify, Apple/Google Podcasts, etc.).
Timestamps
00:05: Surnames in science
03:33: Behavioural ecology or psychology?
13:37: What's it like to do fieldwork in the Kalahari desert, habituating birds to humans?
20:41: The evolution of punishment
29:51: Proximate and ultimate explanations in evolution
46:05: What can we learn about human cooperation by studying cleaner fish?
Podcast links
Website: https://bjks.buzzsprout.com/
Twitter: https://twitter.com/BjksPodcast
Nichola's links
Website: http://www.seb-lab.org/
Google Scholar: https://scholar.google.de/citations?user=u6_SEO4AAAAJ
Twitter: https://twitter.com/nicholaraihani
Ben's links
Website: www.bjks.page/
Google Scholar: https://scholar.google.co.uk/citations?user=-nWNfvcAAAAJ
Twitter: https://twitter.com/bjks_tweets
References
Andreoni, J. (1990). Impure altruism and donations to public goods: A theory of warm-glow giving. The economic journal.
Clutton-Brock, T. H., & Parker, G. A. (1995). Punishment in animal societies. Nature.
Laland, K., Uller, T., Feldman, M., Sterelny, K., Müller, G. B., Moczek, A., ... & Strassmann, J. E. (2014). Does evolutionary theory need a rethink?. Nature News.
Laland, K. N., Sterelny, K., Odling-Smee, J., Hoppitt, W., & Uller, T. (2011). Cause and effect in biology revisited: is Mayr’s proximate-ultimate dichotomy still useful? Science.
Raihani, N.J. (2021). The Social Instinct. Penguin.
Raihani, N. J., & Bshary, R. (2019). Punishment: one tool, many uses. Evolutionary Human Sciences.
Raihani, N. J., Thornton, A., & Bshary, R. (2012). Punishment and cooperation in nature. Trends in ecology & evolution.
Raihani, N. J., & McAuliffe, K. (2012). Human punishment is motivated by inequity aversion, not a desire for reciprocity. Biology letters.
Raihani, N. J., McAuliffe, K., Brosnan, S. F., & Bshary, R. (2012). Are cleaner fish, Labroides dimidiatus, inequity averse?. Animal Behaviour.
Raihani, N. J., Grutter, A. S., & Bshary, R. (2010). Punishers benefit from third-party punishment in fish. Science.
Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift für Tierpsychologie.
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[This is an automated transcript with many errors]
Benjamin James Kuper-Smith: [00:00:00] So I think I'm gonna start off with something really random, which is I was really, so yesterday evening I was really, I was trying to figure out where your surname is from. Oh, right. Because it, it really confused me because I thought like, it doesn't sound English right, and it's, it doesn't sound Germanic either.
Then I thought, okay, it's. It doesn't strike me like, it's not Slavic or anything either. Right. And it's not like from the romance languages and, uh, I thought maybe Italian, but then they wouldn't call a daughter Nicola. Um, so I was really kind of confused where it came from and then I was just like thinking for probably like 20 minutes or something.
And I, I read, I, you know, kind of excluded different areas and countries and then I eventually reached the conclusion that it's probably from the kind of. Arab world, roughly speaking. And [00:01:00] then I wanted to ask, because my prediction was gonna be, for some reason I knew some football players from Algeria and from Iran who had roughly similar surnames.
Nichola Raihani: Mm-hmm.
Benjamin James Kuper-Smith: So I'm gonna guess those, but then today I found on a Wikipedia article that your dad's from Iraq.
Nichola Raihani: Yeah, that's right. Yeah. I was gonna say. But it's quite weird actually because, um, I think Hanney is quite a common surname in Malaysia, which is also, it's because of the, um, Arab speaking
Benjamin James Kuper-Smith: Yeah,
Nichola Raihani: yeah.
Population there as well. But also, um, I remember when I first did my, when I did my, well, before I did my PhD, actually, I worked as a field assistant for. Mandy Ridley on the Baler project, which I ended up doing my PhD on, and she's from New Zealand. And she told me that when I applied for the position, she thought I was actually gonna be Maori because apparently it sounded to her, it sounded like a really Maori surname.
So she was quite surprised when I turned up and I wasn't, you know, didn't, [00:02:00]
Benjamin James Kuper-Smith: yeah.
Nichola Raihani: Wasn't from New Zealand and. Didn't, you know, wasn't Maori. And so I think it is one of those weird surnames that sort of pops up in different places,
Benjamin James Kuper-Smith: huh? Yeah. I mean it, because that's the weird thing, like it never, you know, it's not like sometimes you see a surname and it just sounds really odd and you think, what does that surname?
But in this case, it always sounded completely normal until I somehow start, then started thinking about it, and then suddenly it just got like, it's not from there. It's not from there. It's not from there.
Nichola Raihani: In academia. It's good actually because, um, I never changed it even when I got married, partly because I kind of didn't see why I should change my name, but also my husband's name is much more common and, um, not to disparate his name on me.
Yeah. But his name is perfectly fine. My children have it. But, um, for, I just thought, okay, it is actually quite good being one of the only hanney because there's just no ambiguity about if it's you or someone else.
Benjamin James Kuper-Smith: Yeah, exactly. I mean, I had, um, so I was at. You know, as I said before we started recording, I did my masters at UCL and I had, [00:03:00] um, one lecture by Jar Reese and he said the exact same thing when he was like, it's really easy to find like Jarre.
There just aren't many people with that name. It's a really good thing to have a like slightly unusual name. Mm-hmm.
Nichola Raihani: Yeah, definitely.
Benjamin James Kuper-Smith: And not be, you know, I mean for me, like my surname is Coopersmith, which is half German, half English. If I was just called Benjamin Smith. Yeah, that would be a nightmare to, to find anywhere on the internet.
Anyway. As I said, I'd like to talk more about like evolution of corporation, uh, specifically punishment. Um, and maybe can we start with briefly about your background? So you did a, your degree in natural sciences, right? From what I understand, that's a very. You can kind of choose your, your specialties and all that kind of stuff from that.
Um, so what did you, what did you exactly do as part of that degree?
Nichola Raihani: Yeah, so I did my undergraduate degree at Cambridge, and in hindsight, [00:04:00] I was really lucky to have done my degree there because one feature of the natural sciences degree at Cambridge is that you start very broad and you're only forced to specialize on.
Um, a topic in the final year really. Um, and it, for me, that was really good because it turned out that what I thought I wanted to study and what I actually wanted to study were two different things. So I knew that I'd always been quite interested in behavior. Why animals did the things they did, and I thought that, I kind of thought at the time that that meant that I was gonna end up specializing in psychology.
But as it turned out, the thing that I actually was really interested in, although I didn't know the name for this field at that point, was behavioral ecology. And I really remember having a moment sitting in undergraduate lectures, we were being lectured by. A professor called Nick Davies, who is one of the founders of the field of Behavioral Ecology, who is [00:05:00] still at Cambridge now.
And he was, I can't remember what lecture was, it was would've been something on optimal foraging. 'cause that used to be what he, he used to give us lectures on these kinds of animal decision rules. And I just remember thinking, oh, this is the thing that I'm, this is the thing I'm really interested in, but I didn't know that this wasn't psychology.
I didn't know that this had a, this. Understanding animal behavior was, had this whole different name and it's called Behavioral ecology, and that's what I ended up specializing in in my third year.
Benjamin James Kuper-Smith: Okay. That also, I was always curious like how you. Um, when I look, just look at UCV, I assumed it was, you know, natural sciences to me sounds more like biology than psychology.
And then you did your PhD in biology and you really worked with animals for the first whatever decade or something done. Exactly. But you know, for the first few years of your, of your career, and I was always curious like why exactly you are at a psychology department now and why you then started working with [00:06:00]humans also a few years ago.
Um, but I guess that that interest was, uh, was, was rooted a lot earlier than I assumed just by kind of scanning your cv.
Nichola Raihani: One thing I always say when people ask me that question is that I really feel like I became a psychologist by accident, and it was really driven by the fact that I. Started working on human behavior.
You asking pretty much the same questions that I'd always asked when I was working on non-humans. But it just seemed to me that once I started applying the same sort of logic to try to understand human behavior and everybody else's eyes, that suddenly became, you know, if it's, if you're looking at a cleaner fish or a babbler, fine, that's biology.
Or if you're, it's a human. Oh no, now you're a psychologist. And so, right. In some ways it just became. It, it just became easier to go with the flow and just say, okay, fine. It's psychology. And, [00:07:00] but I've never really, I would, I mean, even now it, I struggle a little bit if someone says, what do you do? What, how do you describe the field you're in?
I, I don't have a particularly good. Answer to the question. I mean, I don't think I'm really an evolutionary biologist anymore, but I also really don't feel like I'm a true psychologist. I mean, I really have no background in psychology at all. And so, yeah, I don't know. I'm a human behavioral scientist, I suppose so.
Benjamin James Kuper-Smith: So you're more a behavioral ecologist who, one of the species you study is humans. Then
Nichola Raihani: Yeah, I, yeah, something like that. Although, I mean, even now that gets confusing because actually I do, you know, a lot of the work I do now is really on the proximate psychological basis of behavior. And so yeah, it, I mean, in a way I am kind of a psychologist, but I, I, I mean.
For example, I'm not, I tried to join the British Psychological Society at one point, and I wasn't allowed to do it because I didn't have an undergrad degree in psychology. So, you know, [00:08:00] I'm, I'm clearly not a psychologist for some people, but I dunno, really what I'm,
Benjamin James Kuper-Smith: oh God, I, sorry, I didn't want to open up a identity question here.
Uh, become more existential. But yeah, I mean, I guess like that's also part of the field, right? I feel like, I think probably anyone who studies. Game theory or something related to it also, it, it just immediately becomes very interdisciplinary. And
Nichola Raihani: yeah,
Benjamin James Kuper-Smith: I mean, like part of the reason also why I, you know, asked you whether, uh, you wanted to be a guest on the podcast is because for, for me, as someone who's, who's.
Also quite intentionally interested in all these, or explicitly interested in these different areas that relate to game theory. It's, it's very difficult to, you know, understand exactly what the evolutionary biologists are trying to do, what the economists are trying to do and that kind of thing. So I think, um, I think, yeah, it probably applies to most people in this field who, yeah, I mean, like I, you know, I, I'm in the, well, 30 of my PhD, but we spent one year doing COVID research, but.[00:09:00]
I feel like I've pretty much read papers from most academic disciplines by now, just by trying to regain theory. Right. Um, but did you, I mean, were you, how did you end up doing your PhD with Tim Clutton Brock? Because as far as I understand, he was one of the first people to really write about punishment in animals and that kind of stuff.
I mean, he had this one famous review paper right. So it seems like you started your PhD, uh, or rather you are, you're still doing a lot of research about punishment and you started it off with one of the, not first people necessarily in the field, but it seems like a great place to do your PhD on that topic.
Nichola Raihani: Yeah, the, that's true. Although, funnily enough, my, my own interest in punishment was sparked. After I'd already finished my PhD and when I started to work with, um, red one, BA on Cleaner Fish. So that was kind of a, almost like a coincidence in some respects, like you could say. [00:10:00]
Benjamin James Kuper-Smith: Mm-hmm. Okay.
Nichola Raihani: Um, yeah, I started working with Tim because, um.
I'd finished my undergraduate degree and I was a bit unsure what I wanted to do and whether I wanted to go and get what my, all my family approvingly called a real job or whether I was gonna, um, you know, maybe do something like different and stay on and do some more studies. And, um, I saw. An advert in the department, somebody recruiting for a field assistant, and it was gonna be, it said it, the position was in the Kalahari, and it would be studying this bird that I'd never heard of.
A pi babbler. I mean, I wasn't particularly like. Interested in birds in any more than any other, the species. I mean, I wasn't an ornithologist or, uh, it, see, I mean my family thought it was a bit of a strange move and I can sort of have some sympathy with that. But anyway, I went to the interview I met who, Mandy Ridley, who had started the Babbler project in the Kalahari and she took me on.[00:11:00]
Um, and so I went out to the Kalahari and helped her to establish this habituated population of PI BAAs that we were able to. Work very closely with, in the sense that we could get very close to the birds without scaring them. That was our main aim of habituating them and that meant we could understand and observe their behavior in a lot more detail than if they're always just flying away 'cause they're scared.
Anyway, it just so happened that the Babbler project was on the same field site as the already established and much larger. Um, project of cooperative breeding in meccas, which was Tim's project in the Kalahari. And so, um. And Mandy was Tim's PhD student actually at the time. So when Tim came out to visit the project, as he usually did twice a year, I met him at that point and I think he was quite keen to keep the baler project [00:12:00] active.
And I suppose I was just, was lucky that I was in the right place at the right time. And that, given that I. You know, I'd done my PhD, uh, sorry. I'd done my undergraduate studies in a relevant field and I was there and I was working on the babblers and I understood the system. Uh, and in those days it was one, it was much easier for a supervisor to basically.
Just offer you a PhD more or less. I mean, there were, I had to still have interviews and things like that, but it is quite different to how the system is now. Where a su, like for example, I don't really have any authority to give anybody a PhD. Uh, they have to, it is much more like in the way it works nowadays is the student gets the student.
Kind of get selected onto a PhD program and then they decide who they want to work with. Whereas this was in the days of when supervisors got given, you know, it was their turn to have a PhD and then they could pick the student they wanted, essentially. And yeah, luckily for me, um, I just was, you know, Tim wanted a baler student.
I was there, [00:13:00] I was working on babblers and. It is kind of embarrassing to say it really, but I just kind of fell into this career, really. I mean, it was not like a big strategic plan that I had that I was gonna work on cooperative breeding and work on baler and everything like that. But it's just, you know, sometimes you make decisions in life and it really affects your whole trajectory then going forward.
And I think that was definitely one of those decisions that it just sort of set me on a different path.
Benjamin James Kuper-Smith: It, it looks a lot more goal directed on paper than I guess it was in reality. So as a, you know, as someone, yeah, I did psychologist. So I have no experience in this kind of, uh, field work or what you, what was the.
Term.
Nichola Raihani: Yeah. Field work. Yeah.
Benjamin James Kuper-Smith: Um, so what, what does it mean to say you were in the Cal Desert habituate, these birds, like practically, what does it mean? Like, first you were, were you there all the time or you had to like come back and forth or, I mean, were you in the desert or like [00:14:00] outside of it, or
Nichola Raihani: No, really.
Yeah. Like what exactly does this mean? In the desert? It means Yes, we were there all the time. So I, my PhD took me three years and. Well, it took me about three and a half years and I would say for a solid maybe 18 months of that I was, um, in the field. I can't quite remember now actually. So I did, when I worked for Mandy, I think I spent, that first field season was nine months, and then I would come and then I think I came back for a few months and then I went back out again for another nine months and came back for a few months and went back out again.
So, tended to be fairly long stints and in a way you, it has to be like that because. And first of all, you are, you're just traipsing around after these animals that are doing, you know, doing their normal life and don't, you know, they're not putting on any special kind of show because you are there. So if you're just trying to get a sense of what this animal does in its day-to-day life and what [00:15:00] kind of events happen and what are the big sort of what we call life history events that can happen in, in, in these animals lives.
You sort of have to stick around a little bit because. Most of what any wild animal does is, is focusing on getting enough food and not getting eaten by something. And you know, it's all kind of quite mundane housekeeping stuff. And so it's only every now and then that you really start to see sort of really.
Not interesting behaviors, but the, the more unusual behaviors, by definition, you just don't see them all that often. So if you want to really get to know a species, you have to spend a lot of time with that, with that species. And that was really the ethos of the work that we did on the Babbler. Um, and in particular, the reason we did it, a lot of the studies we did were observational, although we, we could do some experiments, but we were quite limited in the experiments we could do because.
Part of, part of what we did with the BAAs and also what, what [00:16:00] Tim had done with the Mekas just prior to us starting our project was, um, in the beginning we, we made a huge time investment to habituate the BAAs to our close presence. Um, and that basically means just spending a lot of time. Sitting down not being scary, trying to get them to come closer to you without getting scared.
And um, basically just getting them used to the fact that you are there and you are not a threatening presence. And that just takes a lot of time. Uh, and once you've achieved that, you have to be really careful not to, there are just some experiments you just can't do because it damages. It has the potential to damage that trust between this wild animal that doesn't have to hang out with you if it doesn't want to.
And, uh, the observer, so most of the experiments we ever did were what were experiments that the baler liked. So like, we could do a [00:17:00] feeding experiment because Babbler love, you know, they would love it if we suddenly start, you know, that thing suddenly throwing mealworms every five seconds. That, that's a great experiment.
But experiments involving, like trapping a bird or. Anything that they would perceive as being sort of scary? We didn't really do because it just, the, the cost of it in terms of the, the ability to continue to collect high quality data was too high. But it's, it's actually amazing. Like how, I mean, this was, Mandy was the one who had worked previously on Arabian Babbler with a monster Harvey in the Negev Desert in Israel.
And Ammas had actually trained the Arabian baler to fly over to him from anywhere in their territory. He could get them to come to him by giving this very specific whistle so he could walk around their territory and whistle and all the birds will fly to him. And then also you can weigh the birds by, um, putting a small cup of, uh.
The lid of your water [00:18:00] bottle. If you put some water in there and put it on a, a portable balance, you can tempt the birds to get weighed so you can get a measure of condition and who's doing well and who's not doing so well. And we basically replicated those methods with the pi babbler. So we were able to walk around the desert and whistle in the territory of one group.
That group would come to us and in exchange they'd get a meal worm each, and maybe a small sip of water in. It's actually quite, you know, they're, they're almost better trained than a lot of dogs. I see. In some respects.
Benjamin James Kuper-Smith: I, I always find it fascinating when you hear about these kind of field work studies.
Part of it sounds really interesting and, you know, you're like, it's, it's just also so different than, you know, sitting at a desk all day at programming or experiment or whatever. But at the, but the other half of it also sounds incredibly dull or I dunno what exactly you were doing when you were.
Habituating them and trying not to be threat. I mean, were you reading something or like what were you, were you sitting around and [00:19:00] observing for like five hours all day or?
Nichola Raihani: It is boring and I always say that it's sort of like, I dunno if this is unique to doing field biology, but you really have to be able to have two.
What seemed like completely opposite personality traits, which is you have to be, have a really inquiring mind and want to understand things about how the world works and you know, find out answers to these big questions. But you have to have a really, really high boredom threshold to be able to just trudge around after these animals day after day after day, and not much is happening.
And you know that when I did my PhD, it wasn't. Mobile phones weren't really like a thing then. So it wasn't like you'd just be sat in the, you know, sprawling on Facebook or Twitter or doing, there was, we didn't do anything like that. We were just, I suppose it was like extreme mindfulness. We were just in the desert, sat quietly watching these birds and hoping, you know, they would not be scared of us.[00:20:00]
Mostly as well. A lot of it I remember was trying to make sure you were not sat too near an an ant nest or something like that, because if you sat near an ant nest, it was just a real disaster because you, it was really hard to move because the, it would always make the birds a bit frightened if you started moving around, but you would just start getting attacked by these ants if you sit too near to their nest.
And so most of it was just trying to avoid being sort of bitten by hostile insects.
Benjamin James Kuper-Smith: I, I think I'll remember that whenever I complain about my, me having to do something boring or whatever, I think I'll, I'll bear this in mind, at least I'm not getting attacked by ants. Maybe let's start directly with evolution of punishment then.
Maybe can you first give the kind of precise definition you use of what punishment means in game theory in particular? Um, so we're not confusing it with the everyday term of, I dunno, being mean to people or whatever. [00:21:00]
Nichola Raihani: Is actually, um, a much harder question for me to answer than it sounds because until.
Uh, 2019, I probably would've given you a quite straightforward answer to the question, but in 2019, I published, I basically brought together a lot of different data from different papers, um, in collaboration with Redwan Bahari, who I've worked with quite a bit. And our main conclusion is that punishment is a kind of catchall term that has been used to apply to many different.
Um, behaviors that we see in, in particularly in humans. Uh, and so it, it sort of made us really rethink whether the classic definition actually. Still held or not, but I'll give you the classic definition that we were operating under until we confused ourselves. Um, which is the one which was, um, as he [00:22:00] mentioned, is the one which Tim, uh, sort of proposed in his paper in 1995 with Jeff Parker.
And so, and under that view, punishment happens when one individual gains a benefit by cheating at the expense of a partner. Then the partner who's the Punisher pays a short term cost to inflict. A reciprocal and usually larger cost on this cheating partner. And then the question obviously is if you're paying a short-term cost to do something, well, for this to be under positive selection, we have to ask how that cost is somehow repaid, or how do you get recoup that that cost in future and the one way.
That you could, in theory, recoup that cost in future is if the target of punishment subsequently behaves more cooperatively with the Punisher in future interactions. And so [00:23:00] you have this sort of three step, uh, three steps to the, to an interaction that you would classify as where punishment has occurred.
And I think that quite, uh, I think not everyone, but I think that is a reasonably widely used sort of. Framework for thinking about punishment.
Benjamin James Kuper-Smith: I think the, the crucial isn't the crucial point also that it has to inflict a cost on the Punisher, right?
Nichola Raihani: Yes. Yeah. So that, yeah, so that, that's the, that's the part about the Punisher pays a short term cost to I to impose.
What is usually assumed to be a larger cost on the target. But yes, that point about punishment involving an investment is really, I import or has been really important in thinking about the evolution of punishment. I mean, some people have argued in the past, and I, I also, I think this is fine. Like I'm not, I wouldn't get drawn into semantics on this, [00:24:00] but some people have argued that, um, for some kind of punishment you might not need to.
Specify that there's a cost to the Punisher. And I think that, um, in some ways I agree with that and it, and particularly since we wrote this paper in 2019, I agree with that. I, I agree with that, but I think the reason why people have focused on cases where punishment does involve a cost to the Punisher is that those are just the more.
Difficult cases to explain from an evolutionary perspective. And so there's really no puzzle with explaining something that's immediately beneficial to the actor. And so, you know, for obvious reasons, people maybe haven't spent as much time to try to understand how this could have evolved. And it's, it's cases where.
You do see investments being made, just like in the case of helping behavior, that those are the ones that are kind of, are a bit more puzzling from an evolutionary perspective and do warrant some kind of [00:25:00]explanation.
Benjamin James Kuper-Smith: So does it depend then, does the defini, I mean, does it make sense then to have different definitions between different fields?
So if your evolutionary biologist, uh, may be having the one. You know, narrowing it down to the definition where you, it has to involve a cost for the punisher or investment or whatever you, however you want to call it. Uh,
Nichola Raihani: um, to be honest, I'm not really like the semantics police. Like, I don't care. It, it doesn't, if people wanna call something punishment, um, um, it doesn't fit that definition.
I, it's not, I, I kind of think it's unrealistic to expect that everyone is gonna use exactly the same terminology. To describe a phenomenon, like we can't, people can't even agree on what they mean by cooperation and you know, let alone punishment. So in some respects, I feel like while that might be a nice goal to aim for it sort of also, it might be a bit unrealistic and it doesn't really matter.
I think the more important [00:26:00] thing is just to, or what I always try to do at least, is just be really clear. Like when, if you are talking about something, what do you mean by it? And then people can either, you know, use it in that way or not, or you know, just being clear about what you mean I think is maybe more important.
Benjamin James Kuper-Smith: Yeah. Yeah. I think that's a. I mean, yeah, that's always probably a good point. Yeah. But I think it's a particularly important point if you have like here a term that might be used differently. Um, but the, I mean, you kind of alluded to this earlier, but what exactly is the question or the problem we're trying to solve when we study the evolution of punishment?
Like, why is this something worth studying?
Nichola Raihani: Well, in the context of two player games, it seems it, I mean, at least in theory, it seems reasonably straightforward, how you might explain how punishment could be under positive selection. I mean. I say in theory because in practice it seems like the empirical data don't necessarily support the theory all that well, [00:27:00] but in principle, uh, wanting what a punisher can pay a short term cost to harm a cheating partner, and that cost is repaid if the partner or a bystander is more cooperative with them in future as a result.
I think the bigger puzzle or the bigger question that's, that's a bit more difficult to explain is how punishment evolves in. Groups of more than two players, where you can get one individual who potentially pays the cost associated with punishing, and then any benefits that arise due to increased cooperation are shared among all players in the group, regardless of whether they were they punished or not.
Because what can happen in that situation is that you get what is called second order free riders who. Don't punish, don't pay to punish cheaters, but they nevertheless enjoy the benefits that punishment [00:28:00]brings because they enjoy, they're in the group and they can enjoy any benefits of increased within group cooperation that result from punishment.
And so there, then you're starting to get into the territory of how do you understand, um, the evolution of a costly trait like, like punishment. In these large groups where, where the bene, where the costs are personalized, but the benefits are shared,
Benjamin James Kuper-Smith: is that a different problem from the general problem of cooperation though?
I mean, so for example, you know, you call it second order free writing. Is it in principle or different problem than first order free writing where some people just don't contribute to the common. Good, but others do. Um, it just, it seems to me that it's kind of the same problem almost.
Nichola Raihani: It, it pretty much is.
I mean, it is been described as the same sort of problem. I mean, it is been described as basically putting a second public goods problem on top of a first public goods problem. [00:29:00] And I think so in some ways it is similar, but I think that, um. Understanding the evolution of punishment because, because punishment is by definition, a harmful act that imposes costs on other individuals, understanding how it evolves will not be the same as understanding how our cooperation evolves because they just fundamentally differ in that.
Cooperation is defined as being an act which generates benefits and punishment is defined as being an act which harms. Other individuals. And so, um, they won't be subject to the same evolutionary dynamics because in one case you're generating wealth or whatever you wanna call it, and in another case you're destroying it,
Benjamin James Kuper-Smith: but only in the short term.
Right. I mean,
Nichola Raihani: well,
Benjamin James Kuper-Smith: so as soon as, it seems to me like as soon as you have like an organism that has some sort of foresight, doesn't [00:30:00] that almost eliminate that difference then? Well, if you assume it has to, I guess, increase cooperation. To work that way.
Nichola Raihani: I think, I mean that sort of positing, I think if you were to say that you can solve the evolutionary question of how punishment evolves, I don't think you can just offer this, that kind of foresight as a, as a solution because it is kind of offering approximate.
Mechanism as to an answer to an ultimate question. So it would be a little bit like saying, um, oh, why, why does punishment evolve? Well, if we just assume that individuals just have a punitive sentiment, then it will evolve. But you kind of, you are not explaining where that you, it kind of doesn't really get us to the deeper question of like, how do the costs and benefits get squared off essentially?
Benjamin James Kuper-Smith: Yeah. Maybe can you, so is the, the, [00:31:00] was it proximal and um, the other explanation mechanisms?
Nichola Raihani: Proximal and ultimate. Yeah. I can talk about
Benjamin James Kuper-Smith: is the tin bag thing.
Nichola Raihani: Yeah, basically. Yeah.
Benjamin James Kuper-Smith: Yeah. Can you elaborate a bit on that? Because I think, actually my last interview, someone told me I should read it and now you're the second person.
Nichola Raihani: I think it's, yeah, I think it's actually like one of the least well understood concepts that that could be of the most practical use to people that are interested in psychology. But it is, I, my impression is it's just not that well widely known in psychology. Um. So basically Bergen was a nephrologist who worked on animal behavior and he was famous for his, what was called Bergen's four Ys.
So the four kinds of y question you can ask if you want to know YY an organism does behavior X. You can offer these different kinds of explanation, which are not mutually exclusive, um, but which just. Answer different parts of [00:32:00] that why question. And broadly speaking, you can collapse the four why's into two big buckets.
One is the proximate level explanations, and the other is the ultimate level explanations. And I always find it more helpful to think about things with an example. So let's say you are on safari and you see a lion s. Chasing, um, a springbok and you say, oh, why, why is the lioness chasing the springbok? You might have a why question, right?
Um, so that you could offer approximate level explanation, which would be something like, she's hungry or she has some cups to feed, or she hasn't, you know, maybe she hasn't had food for three days and she needs to get something to eat. So all of those approximate level explanations. Um, and you could also have things like, oh, she's got a high cortisol level because she hasn't eaten for, you know, you could have hormonal explanations that could come under this proximate level explanation.
None of those things get you closer to the ultimate level [00:33:00] explanation, however, of, of, of, on average, how does getting off your button if you're a lion s and chasing a spring block, contribute to your fitness, to your survival and reproductive success? And there you might say, well, on average. Lioness that do hunt Spring book, instead of just sitting under a tree and not bothering to do it, tend to have more offspring and tend to survive better than the ones that don't do it.
And so, um, the ultimate level explanation is more concerned with adaptive significance of the behavior. Like what is, what does the behavior do and why does that help the organism? Um, and then the proximate level. Explanations are more about like, how is the behavior achieved or what prompts the behavior, what and which motives govern or bring about the behavior.
So, and usually evolutionary biologists try not to offer proximate level answers to [00:34:00] ultimate questions. And so, to go back to the punishment question, if you wanna know why an individual, why punishment is favored in. Within groups, like how could a, how could a punishing trait ever be favored within a group of non punishers?
For example, positing something like, oh, you could give, uh, an agent a punitive sentiment, or you could say, oh, the agent has foresight, or something like that. Those would both be examples of offering proximate level. Answers to what's actually an ultimate level question. And because in a way they, those kinds of answers, although they could be perfectly valid, they still, they still allow you to ask another question.
They still allow you to ask, well, um. In the case of they have a punitive sentiment. For example, you can say, oh, well, why is a punitive sentiment under positive selection? Why would agents just not have it? Why don't they just not have a brain that works like that? You know, why don't they have a brain that instead [00:35:00]has a non-punitive sentiment instead of a punitive one?
So anytime you can sort of counter with a, with a counter argument like that, you, you often in the territory of a approximate level explanation.
Benjamin James Kuper-Smith: Okay. Yeah, that's, that's really interesting. And I see what you mean, but in some degree, so I had a, another question I wrote down that kind of relates to this.
Um, and the question is kind of more generally about, um, to what extent it makes sense to see if I can formulate this properly, to what extent it makes sense to do studies in human behavior cognition. If we want to understand evolution because, um, I guess what you kind of just said is that I took a proximal, um, is it approximate or proximal?
Sorry?
Nichola Raihani: Usually approximate, but yeah,
Benjamin James Kuper-Smith: approximate. Okay. Then I'll [00:36:00] use that one. Um, so that I used approximate explanation for an ultimate question, but it seems to me that. So it, it's clear to me how it is an ultimate question if you have an organism that's, let's say non-cognitive, um, so it's like these, like evolutionary simulations and these kind of things that, for example, Martin Novak did for that kinda stuff.
I completely get it. And, um, I have no, um, disagreement there that it seems to me that for humans may be the. You know, if I want to, maybe I'll finish my sentence. It seems to me that for humans, the proximate explanation maybe is also the ultimate explanation, almost in the sense that if I, you know, if, if I am, let's say I'm in a group of people and I want to, I don't know, uh, you know, make money, whatever it is, right?
Um, and I have a specific goal or whatever, if. I somehow struggle to see how the proximate explanation isn't the actual [00:37:00] explanation here. Yeah.
Nichola Raihani: Um, so I can to I totally understand the sort of difficulty actually. And, um, you might, it might be sort of. Heartening for you to know that you are not the only person that has sort of raised this issue.
So, um, and actually this is something which I'm not like massively, um, up to date on, but that Kevin Leland and. Who else? There's a bunch of other people basically who have argued, I can't remember when these papers came out now, but it was quite a while ago, like 10 years ago, I think, um, who basically argued that the prox, that the proximate ultimate distinction doesn't always apply.
And that's, and in particular in the cases of species like humans that are able to construct their niches, um, often the pro, often approximate pro, often approximate. Level mechanisms shape ultimate level, [00:38:00]uh, outcomes. And so I kind of, I understand your consternation and it, I think it's more like a useful heuristic rather than an iron rule.
But I think, I think by and large, for me at least, splitting things up into approximate and ultimate level. Explanations is helpful and, and, and helps you to just be really clear about what exactly is it that you're trying to explain. So, um, for example, another example I use sometimes is sex is sex. So like, why do we, why do humans have sex?
Or why does any, you know, let's think about humans, not just any species, but. We have a lot of sex, we have more of it than we need to, and we seem to really enjoy it. So maybe, you know, on approximate level, you could say, the reason we have sex is that we really enjoy it. But you have to, but then you have to also ask a sort of, um.[00:39:00]
Then you can ask a deeper question of like, well, why has our brain been wired in such a way to make us enjoy doing something that actually, um, you know, takes time for a lot of species? It might be a moment when they're quite vulnerable to predators. Um, they, they, they, if they're having sex, they can't do other productive things like foraging or, um, I dunno, feeding their offspring or something like that.
So. Yes, it could be subjectively enjoyable, but you still have to figure out, well, why has your brain, why is your brain wired that way? Like, why is your brain wired in a way to make you enjoy doing something, um, that you know, actually isn't just cost free? And obviously the answer is that, um, quite, you know, sex has the potential to lead pretty directly to reproduction and to fitness, right?
And so, and every time. That humans for, if we are just talking about humans, that's not to say that every time we have sex, we are doing it with the aim of like, I'm doing this because I'm gonna [00:40:00] want to have a baby or something. Like, obviously that's not the case, and so you don't often, you don't even have to be aware of the ultimate level significance of a behavior.
That in order for that to be there and, and, and often it's not the ultimate level significance of the thing is not the thing that's driving why you're doing that in this, in this immediate moment, and I think it's kind of useful to. I find those examples helpful because when it comes to thinking about cooperation and punishment, we kind of see similar things where it's like if you put people in a brain scanner and you allow them to punish, um, their coplay in a social dilemma game.
You see the areas of the brain reward system that are typically associated with doing rewarding things, those same areas are active when you allow people to punish one another. So, um, you could maybe make the inference that punishment is subjectively rewarding, but that doesn't tell us. Why that tells us why people punish on a [00:41:00] motivational level, but it doesn't tell us why punishment exists and what its ultimate function and benefits might be.
And the same goes for cooperation. So if you, again, if you scan people and have them do cooperative things like give money to charity, and even if you don't scan them, you just ask them how they felt after they did it. People like helping other people. There's, that's the thing. Economists call the warm glow of giving and, um, it's subjectively rewarding.
And so on one hand you could say, you know, some people get a bit annoyed when you try to explain cooperation from an evolutionary perspective because they think that you are, the implication is that if there's an ultimate benefit to the cooperative act, that that is calling into question their approximate.
Motive for doing it. So quite often people will say, well, I like to get help other people because I think it's the right thing to do. Or I don't do this because I'm thinking about myself. I'm just thinking about the [00:42:00]other person. And that's a perfectly reasonable thing to say and it's not quite often is true, but that doesn't, that also doesn't get us away from the kind of Thor question of, well.
Why is your brain, or why is anyone's brain designed in such a way that makes us enjoy doing this thing that is actually costly? And you know, it, you pay cost and you, you benefit somebody else. And, and, and so we need to think about, you know, if we want to understand why the behavior exists and not just what the proximate motive is, that's when you have to think about functional explanations.
Benjamin James Kuper-Smith: I see. Yeah. I mean, I think then maybe part of the differences also just yeah, what your question is, what you're interested in. Mm-hmm. Maybe because I see myself, I don't really see myself as a psychologist, but I think those are the kind of questions I'm more maybe just interested in what are the actual proximate things.
Although I really like when you have a kind of ultimate explanation, it always seems much [00:43:00] grander and unified and everything. But, uh, so actually two points. The first is just a very brief one, so I always put like the references of papers we mentioned in the description of the podcast. Then, so we're talking about like, you know, why like the ultimate costs and why.
You know, would humans or any other species have evolved a certain kind of behavior, but isn't, isn't the answer always to increase your fitness? I'm wondering right now, like, you know, every time you ask that question, I've thought of an answer and always was like, oh yes. To increase your reproductive fitness.
Um,
Nichola Raihani: well, I suppose, yeah, the main question, I mean, that's the what I suppose that's the, ultimately that is what we think, you know, happens in, not in every case. I mean, that's like a really strong adaptation perspective, right? But like the aim of evolutionary biology is to explain behaviors. In terms of their function.
And I think it's not just to basically say, oh, that happens because it increases fitness. But the main thing is [00:44:00] to understand how it increases fitness and because, you know, especially when we think about social behavior, it's just not that straightforward to only consider the actions of one individual in.
Isolation because the minute you have social behavior, your actions can affect the actions of other parties. And so straight away you're into like a whole world of complexity and trying to figure out how, if you do X, how does that affect the behavior of all the other individuals that you. That you either interact with or in your group or in your population.
Benjamin James Kuper-Smith: So is the ultimate question then, not exactly. Why would a certain, okay. So why would a behavior have evolved? Meaning how would that behavior increase the fitness of what, I guess rich tokens would say? The genes rather than the species, but [00:45:00]
Nichola Raihani: yeah. Yeah.
Benjamin James Kuper-Smith: I dunno whether that's still the. Current, I guess that book came out like 30 years ago or something, so I dunno whether that's still the, the current position, but, or whether it ever was.
Nichola Raihani: I think it's just really hard to give broad brush answers to, to those sorts of questions because now, I mean, it really depends on the perspective that, that, that people are taking. And so there. Some people still really still really adhere to the kind of gene eyes, the gene's eye view of evolution. And there's other people that take a much more explicitly multi-level selection perspective.
And I mean, it's really wading into a sort of quite complicated territory that we can go into it, but I think it is, I don't know to what extent we would, um. Maybe get a bit off topic.
Benjamin James Kuper-Smith: Yeah. It's also something I know even less about, about punishments. I'm not [00:46:00] sure. That's also, we only have like 50 minutes left, so I don't think we can start like a whole new thing now.
So maybe as, maybe like a last kind of topic is, I mean, I mentioned I wanted to talk a little bit about cleaner fish, I guess. So maybe because we're already like at these kind of bigger questions rather than like specific experiments or that kind of thing. One question I wrote down is, what can we learn about human cooperation from studying cleaner fish?
Does that question make any sense or is it, am I trying too much to learn? Yeah, because I'm interested in humans. I'm trying too much to incorporate knowledge from everywhere, just because I can. Or is there other specific things we can actually learn from Cleaner fish?
Nichola Raihani: So I'll answer a slightly different question, which is, how can studying cleaner fish help to illustrate some of the general mechanisms that might support cooperation among unrelated partners, including humans?
And so. Yeah, I [00:47:00] mean, on face value, a small fish that lives on a tropical coral reef doesn't seem to have much in common with our own species, and yet from an ecological perspective, some of the features of a cleaner fish's life do seem to have quite a bit in common with the features of a human life in the sense that cleaner fish have.
Thousands of interactions per day with non-relatives. Just like humans have many interactions per day with non-relatives, and in fact, cleaner fish have, um, interactions with complete strangers and they're fish of a completely different species, uh, and fish that they haven't met before and that they might never meet again.
And of course that characterizes large scale human societies to an extent as well. And so one of the questions in the cleaner fish world is what do these interactions look like and how is cooperation maintained? [00:48:00]So it might be. Um, helpful for me to just explain a bit about what it means to be a cleaner fish.
So basically, cleaner fish live on coral reefs, and they, they live on these small territories called cleaning stations, um, which is like, I I say, they're a bit like the hairdressers of coral reefs. They have their little place where they are and. The other fish that live on the reef, which we call the clients, visit the cleaner fish's territory to receive a cleaning service whereby they have act parasites and other nasty stuff on the surface of their skin removed by the cleaner fish.
So the cleaner fish provides a service to the clients. Um, but there's a conflict of interest between cleaners and clients because cleaners actually prefer to eat mucus and living tissue and don't act Parasites is their less preferred thing to eat. And so there's this conflict between cleaners and clients, which is clients would like the [00:49:00] cleanest to just remove it.
Parasites and cleaners attempted to cheat by eating the living tissues. And in that sense, the interaction structure or the incentive structure does have something in common with many of the kind of social interactions that humans have with strangers on a reasonably regular basis. Where mutual cooperation is the kind of.
Best outcome on a mutual level, but potentially an individual can gain a higher short-term benefit by exploiting their interaction partner. And then we have to ask, well, how is cooperation maintained in the face of the temptation to cheat? And it seems like in very, in a very broad sense, the cleaner fish client mutualism.
Uh, the cooperation in this mutualism is stabilized by quite similar mechanisms to those which we see as being important in human societies. Um, and so namely the things that seem to stabilize cooperation in the clean, [00:50:00] efficient client mutualism is punishment and a form of reputation based partner choice.
And we know that both of those things also are important for helping us to understand large scale cooperation in humans as well. So for the cleaner fish punishment works, um, in a couple of ways. So a client that gets a bad service might punish a cleaner fish, and then by aggressively chasing it, and then the cleaner fish will give a better service in the next interaction.
There's also a scenario whereby a pair of cleaner fish sometimes work together to clean a joint, one client between them. And if one of the cleaner fish cheats, if specifically, if the female cheats and causes the client to swim away, then the male will actually punish the clean, the the female, the his female partner for making their shared client disappear.
And that's sort of superficially similar to this, what we call third party punishment in humans, which is where. A Punisher [00:51:00] intervenes on behalf of a victim to punish someone who cheated. So in this case, it's the male cleaner fish intervening on behalf of the client, the victim, to punish the A cheating female.
So we see also this kind of rudimentary third party punishment in cleaner fish as well.
Benjamin James Kuper-Smith: So I kept imagining this as being a hairdresser now and like someone chasing after the hairdresser because they didn't do a good job. Yeah, which actually could happen. But yeah, I mean, I said like when, you know, when I asked like, what can we learn, uh, about humans from clean fish?
Obviously there's a, you know, a value in studying them by itself, and you don't have to be interested in humans, et cetera, et cetera. But, so it's more to, yeah, I mean, I guess you did those studies also. I mean, the, the one you mentioned. Where you studied this kind of third party punishment, I guess third parties always slightly tricky because I guess they have an interest in it working that way, but, um
Nichola Raihani: mm-hmm.
Mm-hmm. Yeah, that's one of the sort of, um, critiques in a way of [00:52:00] the, I mean, we called it third party punishment, and I think people reasonably sort of disagree that it was a case of truth, third party punishment, because you can argue that the male is actually. Involved because they, the food, it's their food source that disappears as well.
Um, we've actually now extended that, that approach to show that a male will punish a, a cheating female even when he isn't involved in the interaction. And so when, uh, okay. Um, he, it's not his food source that swims away and they still, you know, that we still find that the males will do this. So. I think it's one of the key things actually, and I talk about this in my book, um, which is coming out, uh, next well on June the third, or will have come out by the time this podcast is aired.
Um, one of the key things that I think is important to bear in mind about when we see similar looking, [00:53:00] uh, behaviors in humans and in other species, is that you can have something that looks like superficially. Really similar in terms of like the behavioral outcome or the, the behavioral destination you get to, so for example, you can have something which in cleaner fish it looks, you know, you can have something which you can call third party punishment.
And maybe we, we have this other thing in humans that we call third party punishment. But the, the cognitive journey that, that something like a cleaner takes to get to third party punishment compared to the cognitive journey that. A human takes to get to third party punishment are completely different.
And so I think there is this really important thing to bear in mind about just because you see something like reputation based partner choice in the cleaner fish client system or something like third party punishment, that doesn't mean that that, that they're using exactly the same cognitive mechanisms that humans use to achieve those [00:54:00] same.
And I think that is sometimes where there can be a point of confusion is like sort of implying that cleaner fish are able to mind read or you know, have advanced sociocognitive abilities and they really don't. I mean, cleaner fish learn everything probably through trial and error essentially, but because they have like 2000 interactions a day, learning things through trial and error is reasonably sort of ecologically valid for them.
Benjamin James Kuper-Smith: Yeah, I mean, so I've, you know, never done studied in animals. Is it, how difficult is it to stop a kind of anthropomorphizing of the thought process of the fish? I mean, is it with fish so simple that, you know, they're not thinking this through, it's just tried and error? Or do you also have to really kind of, I dunno, do you always have to like, think, like, what's the simplest way.
That this can happen cognitively or
Nichola Raihani: I, I think there's just really no good evidence for advanced sociocognitive abilities in terms of really [00:55:00] sophisticated mind reading capacities that might underpin something like a concern for reputation in humans. There, there just isn't that good evidence for those kinds of mental faculties in any species, let alone a cleaner fish.
Uh, and the same goes for something like punishment where, I mean, like increasingly now I have, um, I'm increasingly convinced that a lot of punishment in humans is approximately motivated by. Fairness concerns and by relative status concerns by looking at what you have relative to what, what another person has, and trying to change that differential in some way.
So sometimes it can be, um, to try to make things fair. Sometimes it can also be people use punish spite punishment spitefully to increase that status over another person. I think when it comes to cleaner fish, there's just no evidence that that cleaner fish have this kind of [00:56:00] awareness of what's fair or a sensitivity to relative payoffs.
And in fact, we know that they haven't got that because we try experimentally to see whether cleaner fish show this sensitivity to relative payoffs by doing inequity aversion like experiments with them and they really don't. Care, like, I just think it's, it's computationally quite complex to compare your own payoffs with the payoffs of, of another individual and to have a kind of fairness concern as so I think, whereas that might motivate some punishment in humans, there's just no evidence that, and that sort of fairness concerns or relative status concerns might motivate punishment in cleaner fish.
But again. It doesn't have to, like, you can get to the same, you can get to a similar behavioral destination via different cognitive roots. And that's, um, that sort of theme in a [00:57:00] way is one that I talk about quite a bit in, in, in my book and, and thinking about how you can see quite sophisticated behavior, even in species that we don't necessarily think of as having particularly sophisticated cognition.
Benjamin James Kuper-Smith: I think that's something I find really interesting is kind of how can you have this kind of seemingly really complex things without, seemingly, without having the apparatus to actually do those kind of computations. Um, but I think that's gonna be a topic for another discussion.