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Old 06-01-2021, 10:34 PM   #51 (permalink)
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It seems like you could

It does seem like ants can tell nestmates apart from non-nestmates (from another colony) based on smell, but it is unclear to me whether they are able to sniff out kin from non-kin.. which should be an important factor in how interactions evolve.
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Old 06-01-2021, 10:55 PM   #52 (permalink)
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Old 06-05-2021, 12:20 AM   #53 (permalink)
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Another source of conflict seem to be in ant species where the shapes (morphology) of ants are more similar. The typical queen is ideally suited for egg-laying while a worker is not, so this would reinforce how a worker should rather spread her own genes by using the queen as her sexual proxy. However, if queens and workers are physically more similar and more alike in capabilities, that seems to increase the level of violent conflict.
Yes, this actually is part of the logic behind Wilson's suggestion that some types of colonies could qualify as superorganisms.

There was a documentary I saw years ago by national geographic which highlighted the ways in which colonies exist along a certain sort of spectrum of eusociality. It was the same documentary I mentioned earlier which showed one of the workers in a colony start to try to go into the process of laying eggs and she was physically restrained and prevented from doing so by her sisters.

This was in a long legged ant colony, where the difference in morphology between the queen and the workers was not as drastically pronounced.

The workers weren't infertile, they just tended not to reproduce. In other colonies, however, the workers are born sterile and cannot reproduce. Their only reproductive outlet is through the queen which in essence means they are intrinsically tied to the well being of the colony.

That is how it is suggested that traits can be selected for on the basis of making the colony more efficient. To my mind it's not much different than traits being selected for that make your body more equipped to survive to reproduction.

If the genes of the colony flow strictly through the queen, then it seems like conceptualizing the colony as a sort of "body" with the queen as its reproductive organ is not all that different from the picture Dawkins paints of how genes eventually started coding to build robots (organisms) to carry the genes around and propagate them.

Maybe there is something I'm missing but I don't quite see the conflict there. Since it's genes that are being selected for, not individuals, why must we insist the individuals necessarily be self interested?. If a colony of individuals which are born into specific castes to work for the purpose of facilitating the queens reproduction is a more effective way of spreading said genes than allowing for more individualism then why wouldn't natural selection favor it?

BTW I read the selfish gene years ago... Could probably use a refresher but I'm familiar with it and as I noted above I don't really see a conflict between his thesis and the idea of a superorganism

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Old 06-05-2021, 12:54 AM   #54 (permalink)
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Also you asked am I in Wilson's camp... Not at all I'm only vaguely familiar with him tbh. I don't take any hard stances one way or the other but the way he articulates the superorganism seems to make sense to me... But wtf do I know?

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Old 06-05-2021, 12:21 PM   #55 (permalink)
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You may be missing something. The gene view / kin selection / inclusive fitness theory predicts normal one family/ monogamous queen ant colonies just fine because the ants are related and have the haplodiploid sex system.

However, the problem is the super colonies for aforementioned reasons. The ants are cooperating with ants to whom they are not (very) related, which seems like it would not evolve and also would be unstable.

Reiterating quickly a couple of things:
  • Altruism - as in a gene sacrificing it's fitness for the sake of other genes - is a losing strategy. The gene by definition doesn't take care of itself in a competitive way. Natural selection weeds such genes out.
  • But if you somehow get a group of altruists, then it is generally very good to be a selfish individual in that group (take from all, give nothing vack). Hence, selfish genes and strategies would be expected to invade a group of altruists

Wilson, as far as I can tell, says that groups of altruists outcompete groups of selfish individuals. But AFAIK his group selection idea doesn't explain how pressure on groups also turn into pressure on the individual. For example, queens and their genes compete against many other queens with their own genes. It's easy to imagine some of the selection pressure that would come from that. Queens and their genes make and shape colonies. Their everyday conflicts would shape those genes. While altruistic groups may be competitive, I still can't quite see what pressure there is on these genes to evolve 'nicely' on behalf of a largely unrelated group. What's the mechanism?

I currently don't feel like group selection is the answer, but I'll admit it's a conundrum. I suspect it's a varied explanation that has a lot to do with ant ecology coupled with possible constraints / limits. Like I wrote, if ants CAN'T tell which ants are kin and which are not (just nestmates from non-nestmates), then something as simple as that should be expected to turn them altruistic. The reason is their ancestral trait is that ALL of an anthill used to be kin. There was no other way, hence they didn't have to recognize kin. If they were friendly to nestmates, then they were nice to kin. Hence, that ancestral ant would be nice to a nestmate whether it was kin or not.

So maybe they could have carried such naivety with them from olden days? You wouldn't expect that because in a colony with several queens/families, selection would so favor any new mutation in ants that lets them recognize kin. But possibly, there could be some limiting factor to ants ability to evolve such a trait.

That's just my thinking so far and I expect it to change in the near future

Edit:

I'll check out Wilson's lecture when I find the time
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Old 06-05-2021, 03:19 PM   #56 (permalink)
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You may be missing something. The gene view / kin selection / inclusive fitness theory predicts normal one family/ monogamous queen ant colonies just fine because the ants are related and have the haplodiploid sex system.

However, the problem is the super colonies for aforementioned reasons. The ants are cooperating with ants to whom they are not (very) related, which seems like it would not evolve and also would be unstable.

Reiterating quickly a couple of things:
  • Altruism - as in a gene sacrificing it's fitness for the sake of other genes - is a losing strategy. The gene by definition doesn't take care of itself in a competitive way. Natural selection weeds such genes out.
  • But if you somehow get a group of altruists, then it is generally very good to be a selfish individual in that group (take from all, give nothing vack). Hence, selfish genes and strategies would be expected to invade a group of altruists

Wilson, as far as I can tell, says that groups of altruists outcompete groups of selfish individuals. But AFAIK his group selection idea doesn't explain how pressure on groups also turn into pressure on the individual. For example, queens and their genes compete against many other queens with their own genes. It's easy to imagine some of the selection pressure that would come from that. Queens and their genes make and shape colonies. Their everyday conflicts would shape those genes. While altruistic groups may be competitive, I still can't quite see what pressure there is on these genes to evolve 'nicely' on behalf of a largely unrelated group. What's the mechanism?

I currently don't feel like group selection is the answer, but I'll admit it's a conundrum. I suspect it's a varied explanation that has a lot to do with ant ecology coupled with possible constraints / limits. Like I wrote, if ants CAN'T tell which ants are kin and which are not (just nestmates from non-nestmates), then something as simple as that should be expected to turn them altruistic. The reason is their ancestral trait is that ALL of an anthill used to be kin. There was no other way, hence they didn't have to recognize kin. If they were friendly to nestmates, then they were nice to kin. Hence, that ancestral ant would be nice to a nestmate whether it was kin or not.

So maybe they could have carried such naivety with them from olden days? You wouldn't expect that because in a colony with several queens/families, selection would so favor any new mutation in ants that lets them recognize kin. But possibly, there could be some limiting factor to ants ability to evolve such a trait.

That's just my thinking so far and I expect it to change in the near future

Edit:

I'll check out Wilson's lecture when I find the time
it seems like the concept of a superorganism is not necessarily the same as a super colony. Not sure if I'm misunderstanding you but it sounds like you are conflating the two. A colony can be a super organism without needing multiple queens etc. If the ants are born into specific castes and more specifically are born without any way to reproduce other than through the queen, that sort of set up seems like one where increased coordination and cooperation between individuals is directly linked to fitness.

As for the super colonies, I can't speak generally but say the one in the bbc documentary that was farming aphids on the spruce trees, Attenborough does mention how the cooperation there is serving a practical benefit with regard to being granted access to the resources that can be marshalled by that big of a colony.
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Old 06-07-2021, 08:13 AM   #57 (permalink)
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it seems like the concept of a superorganism is not necessarily the same as a super colony. Not sure if I'm misunderstanding you but it sounds like you are conflating the two. A colony can be a super organism without needing multiple queens etc. If the ants are born into specific castes and more specifically are born without any way to reproduce other than through the queen, that sort of set up seems like one where increased coordination and cooperation between individuals is directly linked to fitness.

As for the super colonies, I can't speak generally but say the one in the bbc documentary that was farming aphids on the spruce trees, Attenborough does mention how the cooperation there is serving a practical benefit with regard to being granted access to the resources that can be marshalled by that big of a colony.
Yes, I did conflate the two. I've also since watched his lecture that you posted.

His lecture wasn't very illuminating. To my mind, he suggests that evolution happens at multiple levels. Through evolution at the gene/individual level, you can, by chance, evolve a set of important pre-conditions. Something he mentions is a defensible nest and the trait of not dispersing, but staying put. Wilson's thought is that if "normal" evolution leads to a species/colony attaining all these pre-conditions, they can flip over into an altruistic colony. That colony then has to make it in a competition with other non-altruistic colonies and has to be competitive. If it is, it will be selected for at the group level. It is my impression that he thinks group selection is what keeps these groups altruistic over time, yet he does not go into the supposed mechanisms by which that should work. He does mention that it can be shown in maths which I assume means you can make it work in a model. It's true, you can, but my current impression is that this is possible because the living conditions for organisms in models can become very weird, like they can have no parental investment in their young and possibly become adults once they're born, there may be no cost to simply living, etc.

Anyways, something he fails to bring up is whether or not the ants he talks about are kin. He says he distrusts kin selection as an explanation for superorganisms and so, if the individual ants were not kin, he could have capitalized and gotten support for his statement by mentioning that, but he didn't. Hence, I assume the ants in his examples were kin. As mentioned, kin selection theory has no trouble explaining the evolution of a superorganism and actually, solving that mystery was one of kin selection's big selling points in the 60s. Kin selection alone cannot explain a supercolony, but a superorganism is no trouble.. as long as the individuals that make up the superorganism are closely related.

There are other ways to evolve cooperation (like reciprocality), but kin selection is expected to lead to higher apparent "altruism" (altruism on the level of individuals, selfishness on the level of genes).

The word superorganism isn't one that I myself would use because it kinda hides the fact that ant hills are colonies occasionally marked by conflicts of interests between its inhabitants. Individuals don't always behave altruistic and even if they did, normal evolutionary theory (kin selection / inclusive fitness) can predict when that altruism should break down. The word is a heuristic device good for illustrating a point or to use in sentences like "evolution of the superorganism", but that's about it.

He's mostly in the world of ants, but when talking about superorganisms, it would be interesting to go into the world of colony animals, like siphonophores which is the group containing the famous portuguese man o war.



In their specific case, all the zooids (individuals) that make up the man o war are genetically identical and so kin selection would fully explain that.
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Old 06-07-2021, 08:36 AM   #58 (permalink)
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famous Portuguese man o war.
Washed up on a beach they look liked deflated balloons.

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Old 06-07-2021, 09:39 AM   #59 (permalink)
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Yes, I did conflate the two. I've also since watched his lecture that you posted.

His lecture wasn't very illuminating. To my mind, he suggests that evolution happens at multiple levels. Through evolution at the gene/individual level, you can, by chance, evolve a set of important pre-conditions. Something he mentions is a defensible nest and the trait of not dispersing, but staying put. Wilson's thought is that if "normal" evolution leads to a species/colony attaining all these pre-conditions, they can flip over into an altruistic colony. That colony then has to make it in a competition with other non-altruistic colonies and has to be competitive. If it is, it will be selected for at the group level. It is my impression that he thinks group selection is what keeps these groups altruistic over time, yet he does not go into the supposed mechanisms by which that should work. He does mention that it can be shown in maths which I assume means you can make it work in a model. It's true, you can, but my current impression is that this is possible because the living conditions for organisms in models can become very weird, like they can have no parental investment in their young and possibly become adults once they're born, there may be no cost to simply living, etc.

Anyways, something he fails to bring up is whether or not the ants he talks about are kin. He says he distrusts kin selection as an explanation for superorganisms and so, if the individual ants were not kin, he could have capitalized and gotten support for his statement by mentioning that, but he didn't. Hence, I assume the ants in his examples were kin. As mentioned, kin selection theory has no trouble explaining the evolution of a superorganism and actually, solving that mystery was one of kin selection's big selling points in the 60s. Kin selection alone cannot explain a supercolony, but a superorganism is no trouble.. as long as the individuals that make up the superorganism are closely related.

There are other ways to evolve cooperation (like reciprocality), but kin selection is expected to lead to higher apparent "altruism" (altruism on the level of individuals, selfishness on the level of genes).

The word superorganism isn't one that I myself would use because it kinda hides the fact that ant hills are colonies occasionally marked by conflicts of interests between its inhabitants. Individuals don't always behave altruistic and even if they did, normal evolutionary theory (kin selection / inclusive fitness) can predict when that altruism should break down. The word is a heuristic device good for illustrating a point or to use in sentences like "evolution of the superorganism", but that's about it.

He's mostly in the world of ants, but when talking about superorganisms, it would be interesting to go into the world of colony animals, like siphonophores which is the group containing the famous portuguese man o war.



In their specific case, all the zooids (individuals) that make up the man o war are genetically identical and so kin selection would fully explain that.
alright so basically if I understand you correctly you're saying kin selection doesn't explain super colonies yet you also acknowledge that super colonies exist. So apparently there is for the time being, a sort of wrinkle in that specific way of explaining all of evolution.

You don't think group selection is a good explaination. But you also don't have any idea what a good explaination would be. Is that more or less where we're at?
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Old 06-07-2021, 03:43 PM   #60 (permalink)
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alright so basically if I understand you correctly you're saying kin selection doesn't explain super colonies yet you also acknowledge that super colonies exist. So apparently there is for the time being, a sort of wrinkle in that specific way of explaining all of evolution.

You don't think group selection is a good explaination. But you also don't have any idea what a good explaination would be. Is that more or less where we're at?
This seems about right, except for the bolded part.

I have suggested a possible explanation. Ants being unable to recognize kin-nestmates from non-kin-nestmates could (should?) turn them altruistic. Generally, I would expect ants to evolve this ability, but there may be some constraint to its evolution.

However, not being an ant expert, I am unsure of how good that suggested explanation is. Perhaps we should just shoot an email to Bert Hölldobler and ask him what he thinks.
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