how it can be anything other than the 
individual, in the case of honey bees, the queen who heads the super 
organism, and if the hive is succesfull, to a lesser extent the drone(s)

That's what I was saying. But the question really boils down to whether the workers are helping because of so-called altruism, or if they are co-erced, like slaves. 

The question of slaves is interesting. Presumably the keeping of slaves advances the lineage of the slave-owner, as he or she is able to accomplish much more as a result. 

But do slaves also gain fitness thereby, as a result of this relationship? Obviously, fighting against the system can get you killed, so being a compliant slave may be adaptive in this case. 

But then, if the relationship goes south, you have slave-holders that cannot survive without their slaves, and quite possibly slaves that cannot survive without their masters. 

I realize that a colony of bees is not the same as this, but I think it makes a useful point from which to imagine how the relationship of the queen and her workforce mimics slavery, especially inasmuch as neither can survive without the other. 

See: Huxley, Brave New World

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