...Huh. That page is by Eliezer Yudkowsky, and apparently it was written back in June 2009. It seems that writing something up on the wiki is a very poor way to get it discussed.
The argument given seems to make sense, but with two noteworthy caveats. The first is that, while group selection doesn’t work, kin selection does, and in an organism that mostly interacts with close relatives, altruism only towards relatives may not be meaningfully distinct from altruism towards all others of its species. The second issue is that group selection can interact with the founder effect, which is where a very small group enters a new environment and founds a colony that turns into a large population. In that case, altruistic genes in the original founders affect the probability that the colony succeeds; so if an environment is particularly hard to colonize, and many attempts are made, it may be that only a group with some altruistic genes could succeed.
I am no biologist so I may be confused at some point here, but isn’t the point of this discussion article that research has demonstrated group selection in colonies of insects?
Is there some strong sense in which queen bees/ants constitute “founders” or make entire colonies “kin”? Or is there still conflict between the paper nhamann cites and the wiki?
Keep in mind that eusocial species are different in a way that strongly affects who is kin: members of the same colony share 75% genetic material because of how their reproduction works (compare with 50% for parent-child or between siblings—or to 100% for cells within the same organism).
So as a non-biologist, what I’m hearing from this and nhamann’s reply is:
Maybe/probably there is an effect in genetics that would cause the bees in the original study cited to be more closely related to one another in ways that would encourage specific group selection in ways that wouldn’t extend smoothly to other species. However, this effect is losing ground in the scientific community so it is worth reexamining the case.
It’s more complicated than that. It actually turns out that several mathematical analyses since the 80s have shown inclusive fitness theory and group selection theory to be mathematically equivalent*. In fact, both sides seem to readily acknowledge this fact, so the two groups’ appear to be arguing over semantics (where does selection “really” occur?)
This argument over semantics is best illustrated by the following quote at the end of the supplementary information for the paper in the OP:
Finally, we propose that kin selection among social insects is an apparent phenomenon which arises only when you put the worker into the center of evolutionary analysis. Kin selectionists have argued that a worker who behaves altruistically by raising the offspring of another individual, requires an explanation other than natural selection, and this other explanation is kin selection. We argue, however, that there exists a more convenient coordinate system.
* There is, of course, ongoing debate about this, and whether one theory is more general than the other.
Some animals (esp. social insects) behave “altruistically,” i.e. via raising others’ offspring.
Some people consider the explanation “an altruistic set of animals produces more offspring which inherit their altruism” sufficient, and some people require saying “animals that behave altruistically toward their kin produce more offspring etc.” and calling it “inclusive fitness.”
People who think the first way point out that people who think the second way are ignoring phenomena like domestication (i.e. behaving altruistically toward other species) and being too specific.
I’m throwing the example of domestication in there because it came to mind for me as a way in which the first statement seems more general than the other.
Mmm, I’m probably going to have to try to write a post explaining what I think I understand about the debate, because there’s a lot of other points.
Researchers in the behavioral economics literature have been conducting experiments with human subjects playing public goods games. Time and time again, it has been shown that cooperation can be sustained in groups when subjects are allowed the ability to punish defectors and reward cooperators. They argue, at the very least, that inclusive fitness theory cannot explain such behavior between strangers.
Nowak and others seem to be proposing an expansion of the biological theory of genetic evolution to a mathematical theory of evolution. He and others propose that evolution can be genetic or cultural or any other kind of evolutionary process you can dream up, as long as it fits the mathematical framework they’ve developed (which is based on evolutionary game theory).
The whole controversy seems silly, however. Group selection effects have been observed among insects in laboratory conditions. Inclusive fitness theorists argue that these conditions are rare, so they can be safely ignored for the genetic theory of evolution. The multilevel selection theorists argue that evolution is a mathematical process which can explain a variety of phenomena beyond just genetic evolution. Both sides are strictly correct, they’re simply talking past each other.
What you said is only true for organisms which happen to engage in haplodiploidy. From the paper by Nowak et al in the OP:
By the 1990s, however, the haplodiploid hypothesis began to fail. The termites had never fitted this model of explanation. Then more eusocial species were discovered that use diplodiploid rather than haplodiploid sex determination. They included a species of platypodid ambrosia beetles, several independent lines of Synalpheus sponge-dwelling shrimp (Fig. 2) and bathyergid mole rats. The association between haplodiploidy and eusociality fell below statistical significance. As a result the haplodiploid hypothesis was in time abandoned by researchers on social insects
...Huh. That page is by Eliezer Yudkowsky, and apparently it was written back in June 2009. It seems that writing something up on the wiki is a very poor way to get it discussed.
The argument given seems to make sense, but with two noteworthy caveats. The first is that, while group selection doesn’t work, kin selection does, and in an organism that mostly interacts with close relatives, altruism only towards relatives may not be meaningfully distinct from altruism towards all others of its species. The second issue is that group selection can interact with the founder effect, which is where a very small group enters a new environment and founds a colony that turns into a large population. In that case, altruistic genes in the original founders affect the probability that the colony succeeds; so if an environment is particularly hard to colonize, and many attempts are made, it may be that only a group with some altruistic genes could succeed.
I am no biologist so I may be confused at some point here, but isn’t the point of this discussion article that research has demonstrated group selection in colonies of insects?
Is there some strong sense in which queen bees/ants constitute “founders” or make entire colonies “kin”? Or is there still conflict between the paper nhamann cites and the wiki?
Keep in mind that eusocial species are different in a way that strongly affects who is kin: members of the same colony share 75% genetic material because of how their reproduction works (compare with 50% for parent-child or between siblings—or to 100% for cells within the same organism).
So as a non-biologist, what I’m hearing from this and nhamann’s reply is:
Maybe/probably there is an effect in genetics that would cause the bees in the original study cited to be more closely related to one another in ways that would encourage specific group selection in ways that wouldn’t extend smoothly to other species. However, this effect is losing ground in the scientific community so it is worth reexamining the case.
It’s more complicated than that. It actually turns out that several mathematical analyses since the 80s have shown inclusive fitness theory and group selection theory to be mathematically equivalent*. In fact, both sides seem to readily acknowledge this fact, so the two groups’ appear to be arguing over semantics (where does selection “really” occur?)
This argument over semantics is best illustrated by the following quote at the end of the supplementary information for the paper in the OP:
* There is, of course, ongoing debate about this, and whether one theory is more general than the other.
Okay so now what I’m getting is more like:
Some animals (esp. social insects) behave “altruistically,” i.e. via raising others’ offspring. Some people consider the explanation “an altruistic set of animals produces more offspring which inherit their altruism” sufficient, and some people require saying “animals that behave altruistically toward their kin produce more offspring etc.” and calling it “inclusive fitness.” People who think the first way point out that people who think the second way are ignoring phenomena like domestication (i.e. behaving altruistically toward other species) and being too specific.
I’m throwing the example of domestication in there because it came to mind for me as a way in which the first statement seems more general than the other.
Mmm, I’m probably going to have to try to write a post explaining what I think I understand about the debate, because there’s a lot of other points.
Researchers in the behavioral economics literature have been conducting experiments with human subjects playing public goods games. Time and time again, it has been shown that cooperation can be sustained in groups when subjects are allowed the ability to punish defectors and reward cooperators. They argue, at the very least, that inclusive fitness theory cannot explain such behavior between strangers.
Nowak and others seem to be proposing an expansion of the biological theory of genetic evolution to a mathematical theory of evolution. He and others propose that evolution can be genetic or cultural or any other kind of evolutionary process you can dream up, as long as it fits the mathematical framework they’ve developed (which is based on evolutionary game theory).
The whole controversy seems silly, however. Group selection effects have been observed among insects in laboratory conditions. Inclusive fitness theorists argue that these conditions are rare, so they can be safely ignored for the genetic theory of evolution. The multilevel selection theorists argue that evolution is a mathematical process which can explain a variety of phenomena beyond just genetic evolution. Both sides are strictly correct, they’re simply talking past each other.
Okay, so, again parsing this in layman’s terms:
-Some researchers say “let’s use group selection for studying evolution.”
-Some other researchers say “when we talk about genetic evolution we don’t need group selection.”
-The first researchers say “but it’s useful in other contexts, why ignore it?”
-The second group says “because we research genetic evolution.”
Is that about right?
I think that’s about right, but this is what I understand from at most a weekend of skimming journal articles, so I might be off here.
What you said is only true for organisms which happen to engage in haplodiploidy. From the paper by Nowak et al in the OP: