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I'm trying to understand what problem led to a development of kin selection.

The explanations I've found consist roughly of the following:

The existence of some altruistic and cooperative behaviors in social insects like honey bees, the phenomenon of sterile castes formed by non-reproductive individuals that leave no offspring and the suicidal role of these insects in defending the hive, were traits that natural selection could not favor. Kin Selection | Nectunt

What I do not understand: Why could natural selection not favor the phenomenon of sterile castes?

From the perspective of the sterile worker: Since it can't reproduce, no strategy will affect the number of offspring it has (will always be 0). So there is no conflict with natural selection.

From the perspective of the queen bee: Her offspring consist of sterile worker bees and a new fertile queen bee. The worker bees improve the viability of the next queen bee. This is consistent with natural selection.

Therefore, I can't see the relevance of the worker bee sharing lots of genes with its sister queen bee. This seems to be nothing but an ancillary effect of the mechanisms of evolutionary change. Imagine, the queen bee could produce worker bees without a genome (like a robot), or a genome that is not related to its sister queen bee, those worker bees would still show altruistic behavior.

What am I missing?

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Why natural selection leads to selfishness

Imagine a population of individuals. At a given locus, some individual carry the A allele and some others can the S allele (A for altruistic and S for Selfish). For simplicity, we will assume organisms are haploid. Individuals carrying the S allele are selfish and will take care of themselves only. The A allele are altruistic meaning that they produce behaviours (and non-behavioural phenotypes) that will reduce their fitness and increase the fitness of other individuals in the population.

As the altruistic behave as to reduce their fitness, they will be at a selective disadvantage and therefore, over time, only selfish individuals will remain.

How can altruism evolve then?

The study of the the evolution (and evolutionary consequences) of behaviour (and non-behavioural phenotypes) that affect the fitness of other individuals in the population is called social evolution. Social evolution is a wide field of knowledge and I will here attempt a very short summary of it so many concepts will be left out!

There are two main (non-exclusive) ways by which altruism can evolve:

  • Repeated interactions (False altruism)
  • Kin interactions (True altruism)

Repeated interactions

The concept is grounded within game theory. A good introduction to game theory is definitely out of the scope of this question but I just want to give a little bit of an intuition. I will over-simplified the concepts of game theory but you should definitely have a look at it. Those concepts are very enlightening for questions of evolutionary biology, economy, psychology, sociology and even philosophy.

If for example, the altruistic individuals are actually only altruistic to those that have shown altruism in the past (via a Tit-for-Tat strategy for example or through the use of reputation), then altruistic individuals may end up have a higher fitness than the selfish individuals. Imagine for example that selfish individuals give nothing and receive nothing. Let's assume that all altruistic individuals, all make one good deed and receive one good deed. Giving a good deed comes at a cost of having, say 0.8 fewer offsprings on average. Receiving a good deed comes at a benefit of 1.2 extra offsprings. Overall and on average, each altruistic individual will produce 0.4 more offsprings than the selfish individuals and therefore, over time the whole population will become altruistic.

We often refer to these concepts as false altruism because altruistic individuals actually has a fitness increase over their life time. Let's now talk about true altruism where altruistic individuals really just give without receiving but yet, the allele coding for altruism thrives.

Kin interactions

Note first that any kin interaction follows within a give game theory scenario. As such kin interaction is just an add-on on what I discussed earlier. For those used to Hamilton's rule, please note that Hamilton's rule apply to Prisoner's dilemna (a type of game theory scenario) only.

So, imagine now that altruistic individuals are being altruistic with their family members (their kins) only (either because they direct their altruism or because there is a strong population structure). Note that such altruistic behaviour can pretty indirect as helping your partner (who we will assume is not part of your close family) will have for consequence that your partner may offer better parental care and therefore by being altruistic to your partner, you are actually being altruistic to you offsprings (which are your kins).

Because the kins of altruistic individuals probably carry the A allele, eve if the altruistic individual is experiencing a lifetime cost in fitness, the A allele may still thrive and the recipients of this altruism are likely also carrying the A allele. In other words, a truly altruistic individual is an individual that sacrifice itself for helping the allele A present in its kins to thrive. In yet other words, true altruism is when an allele is ensuring that copies of itself present in other individual will manage to get passed on.

And this is called Kin Selection. I can now clearly answer your question

Your question

What problem does kin selection solve?

Kin selection explains "true altruism"!

Only "true altruism" can yield to such separation of the reproductive task as that some individuals are perfectly steriles. In your post, you say

From the perspective of the sterile worker: Since it can't reproduce, no strategy will affect the number of offspring it has (will always be 0)

What is missing from this logic is that, 1) at a time workers could still reproduce and 'gave it up' and 2) in many (maybe most?) social species (honeybees, termites, ants and also in non-hymenopteran eusocial species), workers are still able to sometimes reproduce and are also able to overthrow the queen.

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  • $\begingroup$ Found this: rspb.royalsocietypublishing.org/content/282/1803/20142886 "helpers do not necessarily improve their inclusive fitness by foregoing reproduction. These findings suggest the earliest evolutionary steps towards eusociality are most favored if mediated through maternal manipulation.“ That’s what I mean, if this can be achieved my maternal manipulation, then it seems to me to follow, that relatedness isn’t absolutely necessary. But I’m starting to realize that I asked too broadly and should first read some papers. If I then still don’t get it, I'll post a follow up question. $\endgroup$ – Mathias Jan 6 '18 at 18:09
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Kin selection explains a violation of the assumption that individuals will always selfishly promote their own reproduction. (Lemmings jumping into the sea is a myth that does not happen). Every individual is supposed to work toward sending its own genes into the next generation. In your example, you say that the queen should just pump out workers to serve herself. And she does. But the workers are individuals too, with their own genes, and theoretically, should be selfish. However, in this case, due to honeybee reproduction and haplodiploidy, the workers are more than 50% related to their siblings. The math works out such that working toward the success of the colony is a more efficient strategy for promoting worker genes than if the workers were to try to reproduce on their own. Remember that one point in evolutionary time, these workers were reproductively capable, and gave up their own reproduction to serve the colony. So I think that what you are missing is that at some point in time, workers were once capable of reproduction, and gave it up. They were not always cogs in a machine.

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  • $\begingroup$ "But the workers are individuals too, with their own genes, and theoretically, should be selfish." -> I addressed that: They are sterile, so the cost of their "sacrifice" is 0, which is consistent with selfishness. $\endgroup$ – Mathias Jan 6 '18 at 12:41
  • $\begingroup$ "workers were once capable of reproduction, and gave it up." From the little I know, it doesn't like they were the ones who gave it up. It's more like it was taken from them. $\endgroup$ – Mathias Jan 6 '18 at 12:42
  • $\begingroup$ They were not always sterile. They evolved from ancestors that were fertile. This is what breaks the rule. $\endgroup$ – Karl Kjer Jan 6 '18 at 12:43
  • $\begingroup$ It was not a conscious decision. Kin selection explains the math, and how such behavior could evolve without breaking assumptions about selfish reproduction. $\endgroup$ – Karl Kjer Jan 6 '18 at 12:44
  • $\begingroup$ And no, reproduction was not taken from them. Their genes still go forward into the next generation more efficiently than if they were laying their own eggs. It is a kind of decoupling of the individual from reproduction. $\endgroup$ – Karl Kjer Jan 6 '18 at 12:46

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