I know the TSG gets lots of attention and lots of people seem to take it as the fundamental word on how evolution works. This is simply not true. TSG presents the rather limited view of a geneticist thinking about genes and is directly at odds with other understandings of how evolution works such as genes as followers [0]. I often have a visceral reaction to such individualistic and incredibly western views of how evolution proceeds because they seem to be wilfully blind to the fact that for 99+% of the time evolution does not operate at the level of individuals, in fact, if you are in one of those rare events where individuals DO matter your likelihood of extinction is incredibly high and as a species you really don't want to be there. If the perspective offered in TSG has a place anywhere in evolution it is long before there were multicellular organisms or perhaps even cells.

I was inspired to this rant the other day when watching a video of tortoises flipping each other over [1]. Early in tortoise evolution there are HUGE benefits to a male that doesn't flip his buddies over and gets all the ladies as a result. No amount of kin selection can prevent this if there is not already a significant latent potential in the organism's behavioral repertoire to cooperate (flip). The notion that a single gene is somehow in control is a complete and utter fiction, SNPs are not robust ways to build behaviors or even to create phenotypic diversity. The variation that leads one tortoise to flip the other over is likely the product of thousands of genes and is far more continuous than a simple 'flipping gene.' In fact, if you see flipping behavior at all it is extremely likely that the developmental program and the genome of the organism as a whole is structured such that the behavior will arise repeatedly in different individuals as a result of a number of different minor changes in gene expression. If a flower grows, its is more likely to be the case that the ground is good for growing flowers than that the flower suddenly innovated.

tl;dr If you want your tortoises to flip, you're going to need more than one gene and you can't think about evolution on an individual basis.

0. http://www.ncbi.nlm.nih.gov/pubmed/21257223, https://www.researchgate.net/profile/Eva_Jablonka2/publicati... 1. https://www.youtube.com/results?search_query=tortoise+flip