Talking to Professor Samir Okasha of Bristol University about altruism and evolution:
Save this mp3 file to your computer
Save this mp3 file to your computer
Text version
Paul Craze: One of the good things about a subject like evolutionary biology is just how it can bring in so many ideas from lots of different disciplines and lots of different areas of science and investigation of human knowledge. And one of those areas is philosophy, and that has a lot to say about exactly how natural selection operates on organisms and, in particular, the level at which selection acts. Does it act on genes, we’ve heard a lot about the selfish gene, does it act at the level of a gene or at individual organisms or populations, this so-called levels of selection debate that’s been going on for sometime, and I’m here today with one of the people who’s done a lot of work on this, Samir Okasha, from the University of Bristol, the Department of Philosophy at the University of Bristol. So this levels of selection debate, Samir, what is all that about?
Samir Okasha: It’s a foundational problem in evolutionary biology that goes back to an issue that was discussed by Darwin himself, so for the most part Darwin thought of natural selection as operating at the individual level. So what that means is that it’s individual organisms that survive, compete and reproduce in Darwin’s scheme, leading to traits which enhance their potential to do that. So the Darwinian view of things, in the first instance, applied at the individual level, leads us to expect that individual animals and plants will possess traits that enhance their own chances of survival and reproduction that give them a competitive advantage vis-à-vis other individuals. However, we indeed do find that process in operation throughout the living world. However, it’s also the case that we sometimes encounter traits, and Darwin was aware of this point, which seem not to serve any particular advantage to the individual organism that bears or expresses the trait. So, for example, in many social animals, we find animals engaging in so-called cooperative or altruistic behaviours which can in fact impose a cost to themselves and yet benefit other or other members of their local group or population or species.
Paul: So you say a social species, would that be humans or some other species?
Samir: Certainly, we find this in humans, but we find it much more generally. I mean, a striking example of this sort of cooperative or altruistic behaviour are traits that we find in insects; in particular in so-called eusocial insects which live in, often in very complex colonies. So ants, wasps, bees and termites, all contain species which live in extremely elaborate colonies, where the vast majority of members of the colony do not in fact reproduce at all but devote their whole lives towards indirectly assisting the reproduction of the queen. So they’ll devote their whole lives to building the nests, foraging for food, guarding the nest, tending the queen’s eggs, tending to every need of the queen and, in fact, forego reproduction themselves.
Paul: And would this explain something like the probably apocryphal stories of bees killing themselves and pulling the sting out of them and…?
Samir: That would be another example.
Paul: Yes, but they obviously can’t reproduce then?
Samir: Yeah, I mean any animal that engages in self-sacrifice, in fact self-sacrifice isn’t particularly common but it does happen, that clearly is not doing anything to enhance its personal fitness. I mean if you engage in a behaviour that reduces the number of offspring that you leave, and in some cases reduces it even to zero, by altruistically sacrificing yourself for the good of your group then clearly that’s doing nothing to enhance your individual fitness. And on the face of it such behaviours, which are quite common through out the living world, pose a puzzle, pose a challenge to the theory of evolution by natural selection, because that theory leads us to expect that animals, individuals will behave in a way that enhances their own prospects for survival and reproduction, not those of others, so why is it that we find examples of animal behaviours, particularly in social species including humans, where the behaviours seem detrimental to the individual fitness of the organism? Now, there are very many possible explanations you can give of such traits but one quite standard way of thinking about them is to argue that they are evidence or they reflect the operation of natural selection at a higher level than that of the individual organisms.
Paul: So this is why it is called the levels of selection?
Samir: Precisely. So in the levels of selection type problem we typically see a conflict between what’s individually advantageous for an organism and what’s advantageous for the whole group or species of which the organism is a part. So this problem goes, sometimes it’s called the tragedy of the commons, and also it’s called, in economics it goes under the name of public goods problem, where we find a conflict between individual self-interest and group welfare. And often in such circumstances, then individual self-interest wins out and we find individuals behaving in ways that don’t enhance their group welfare. As for example, when individual nations all pollute the natural environment to such an extent that it becomes detrimental for everybody - the problem there is that there's no individual incentive to cease to do so.
Paul: Yes. It’s usually in their economic interests to pollute as much as possible.
Samir: To continue their selfish polluting behaviour, that’s right. But interestingly, in the evolutionary sphere we also encounter a whole range of cases where natural selection doesn’t seem to have produced the purely individual self-interested solution that’s detrimental for group welfare but, on the contrary, seems to have moulded individuals, in some cases, to behave in ways that are in the interests of the larger community, one of the larger groupings to which they belong. So, social insect colonies are one such example, other possible examples come from human species, and from any group living species, and indeed the concepts can apply more generally. So you can think of a multi-celled organism, like you or I, or any multi-celled organism, be it plant or animal, as a cooperative grouping of cells. So as we know multicellularity is obviously not the ancestral state. I mean before multicellularity evolved, however many millions or billions of years ago, then life existed in the single cell stage for a long, long time, and the transition to multicellularity arose independently I think perhaps some fourteen different times, so clearly there was an advantage to it. But it’s striking that a multi-celled organism can be thought of in many ways as analogous to a cooperative group, all of the cells working together to enhance the welfare of the whole, and sometimes of course that cooperation breaks down, and that’s when you find things like cancerous cell lineages and the development of tumours that are detrimental for the organism as a whole.
Paul: Right, so this is cells going off on their own, really.
Samir: That’s right, that’s right. So this is the conflict between individual and group played out at a different level, where the individual is the cell and the group is the whole collective of cells, and indeed at a lower level still, within a cell, then you can find conflict between the different genes in the cell. So, for example, there can be conflict between the nuclear genes and the mitochondrial genes, even within a single cell, which arises because their mode of transmission to the next generation can be different.
Paul: So these are genes which are in the nucleus of the cell and then there are some which are in the other parts of the cell?
Samir: Yeah, that’s right, which are found in little intracellular organelles called mitochondria, and other organelles, which exist in the cell but outside the nucleus. Then you can even have conflict between genes within the nucleus, between nuclear genes, within the same cell, within the same organism, can be in evolutionary conflict with each other, and that arises basically because reproduction isn’t clonal, so the genome in a sexual species isn’t passed on intact to the offspring generation but is broken up and recombined, and that can create interesting situations where the different genes, within the same genome, are in a sense in a conflict with each other, have different interests. So, one very nice example of this arises in virtue of the fact that genes in the mitochondria are only transmitted down the maternal line, so in a…
Paul: So you just inherit those from your mother?
Samir: That’s right. You only inherit those from your mother.
Paul: It’s not because they’re in the, they’re outside of the nucleus?
Samir: That’s right, that’s right, so they’re not contained.
Paul: You just get that from the egg?
Samir: Yeah, so the sperm doesn’t contain any mitochondria, and the same applies in plants. So in hermaphroditic plants, that’s plants which contain both males and females, which devote resources to both male and female sexual function, you can get an interesting conflict where the mitochondrial genes have an interest in the plant only producing females, but the other, the nuclear genes have an interest in devoting resources to producing males as well, and that conflict can lead to striking evolutionary phenomena. So that’s just one more example of this more general point that in any multilevel scenario, we get potential conflict between what’s optimal at one level and what’s optimal at a higher or a lower level, which is the source of the sort of levels of selection problem.
Paul: And it’s fascinating that that’s such a, we’re really talking about something very fundamental here, the transition between single-celled life and multi-celled life and…
Samir: We are. We are. We’re dealing with a phenomenon that’s not at all peripheral, I mean, and the history of this debate is striking in that originally in the 1960s there was a large discussion of the concept of group selection, and many authors came to the conclusion, rightly or wrongly, that it was a relatively peripheral and minor aspect of evolutionary biology, not something you needed to worry about too much, but we’ve since come to realise that the idea of group selection and more generally multilevel selection is in fact fundamental to the evolution of life. And so the very things that we call individuals, organisms, multi-celled organisms, like you and I, from another perspective can in fact be regarded as groups, and the fact that individuals function so well and so cohesively most of the time is testimony to the power of selection at the group level to shape these entities which then become so cohesive that we cease really to think of them as groups and think of them as individuals themselves. So, in short, the levels of selection discussion and controversy is by no means a minor and peripheral one, as it was sometimes thought to be, but in fact permeates the biology of pretty much all life forms on Earth.
Paul: It’s certainly fascinating to think of yourself as this collection of cells. We think of ourselves as just the individual things but as a collection of cells…
Samir: That’s right. It is fascinating, and the phenomenon of carcinogenesis, of cancerous tumour development, does indeed illustrate this because I mean what’s happening in a case of cancer in effect is some rogue cell lines ceasing to behave in the interests of the whole organism; replicating themselves very fast to gain a short term replicative advantage that’s then detrimental to everybody in the group because it unfortunately often kills the host organism, as we know.
Paul: I think there, if anyone was looking for a reason for studying evolution and biology, you know, who didn’t just think that it was interesting, I mean there is an obvious example.
Samir: There would be potential practical applications.
Paul: Definitely, well Samir, thanks very much indeed.
Samir: Thank you.