I have read Selfish Gene (now an outdated book that even Dawkins admits has many flaws...)
Fair enough, but I'm interesting in knowing if you read the original version or the one with "Nice guys finish first" chapter.
and do my best to stay on top of the literature of evolutionary biology.
The simple fact is cases of "reciprocal altruism" are very rare, and not a single scientist in the field suggests tick removal is an example.
You really want to kill my tick removal example I see, laugh
. I said long ago that it simply seemed to be an apt example, but it's clearly not the only one.
You do a fine job of babbling on but can never answer the hard questions. For the trait to qualify it must have an "evolutionary cost" to the organism carrying it out. There is no "evolutionary cost" to taking ticks off another animal. What is the evolutionary cost you suggest occurs from tick removal?
It's time spent on a creature other then the one doing the picking. Creatures only have so much time to do what they need to do to survive and procreate. The reason that I believe it would qualify as reciprocal altruism is that although a monkey picking ticks off of his friends back may not benefit him immediately, it -would- benefit him if his friend returned the favour later. This is why Dawkins' "Grudgers" are so important; essentially, in general, we are more generous to those who are more generous with us; evolutionarily speaking, this makes sense if helping each other out is more then a zero sum game.
Scientists are desperate for examples of it, and none of them mention tick removal.
Dawkins doesn't seem to hold your point of view, what with his belief
that "many wild animals and plants are
engaged in ceaseless games of Prisoner's Dilemma, played out in evolutionary time." Dawkins' himself mentions tick removal in one of his examples that I get to further below.
2) how do genes communicate their desires to the organism or;
Dawkins' has never argued that they do. You may want to take a look at his book
The Blind Watchmaker.
3) how does the environment act on genes, so that genes become the level of selection?
I would say that the environment is crucial in gene selection. However, I'd also like to point out that, with humans and other fairly intelligent animals, I don't believe that genes are at the forefront anymore; rather, I believe the memes, a term that Dawkins' coined, are the real masters now. Memes are essentially ideas. Ideas can take many forms, from religion to politics, but they can be quite removed from genes, which is how genetically suicidal ideas like a celibate priesthood could exist.
Even though you may want to remain stuck 30 years in the past, here is a much more recent peice in the field on the subject (which discusses the naming problem) that might help you understand why you are so far off the mark with the tick removal.
http://plato.stanford.edu/entries/altruism-biological/
Despite the attention paid to reciprocal altruism by theoreticians, clear-cut empirical examples are relatively few (Hammerstein 2003, Sachs et al. 2004). This is probably because the pre-conditions for reciprocal altruism to evolve- multiple encounters and individual recognition - are not especially common. However, one possible example is provided by blood-sharing in vampire bats (Wilkinson 1984, 1990). It is quite common for a vampire bat to fail to feed on a given night. This is potentially fatal, for bats die if they go without food for more than a couple of days. On any given night, bats donate blood (by regurgitation) to other members of their group who have failed to feed, thus saving them from starvation. Since vampire bats live in small groups and associate with each other over long periods of time, the preconditions for reciprocal altruism are likely to be met. Wilkinson's study showed that bats tended to share food with their close associates, and were more likely to share with others that had recently shared with them. These findings accord with reciprocal altruism theory.
Trivers (1985) describes an apparent case of reciprocal altruism between non con-specifics. On tropical coral reefs, various species of small fish act as ‘cleaners’ for large fish, removing parasites from their mouths and gills. The interaction is mutually beneficial — the large fish gets cleaned and the cleaner gets fed. However, Trivers notes that the large fish sometimes appear to behave altruistically towards the cleaners. If a large fish is attacked by a predator while it has a cleaner in its mouth, then it waits for the cleaner to leave before fleeing the predator, rather than swallowing the cleaner and fleeing immediately. Trivers explains the larger fish's behaviour in terms of reciprocal altruism. Since the large fish often returns to the same cleaner many times over, it pays to look after the cleaner's welfare, i.e., not to swallow it, even if this increases the chance of being wounded by a predator. So the larger fish allows the cleaner to escape, because there is an expectation of return benefit — getting cleaned again in the future. As in the case of the vampire bats, it is because the large fish and the cleaner interact more than once that the behaviour can evolve.
Yes, I'd heard of both of those examples before. I imagine I read them in one of Dawkins' books.
While the Prisoner's Dilemna makes for an interesting mathematical analysis, it does not provide any data to actually solve the problem.
That statement really makes me wonder if you read Richard Dawkins' updated version of The Selfish Gene. Let me quote a section of that chapter that's a little further in:
****************
The birds in Chapter 10 who removed ticks from each other's
feathers were playing an iterated Prisoner's Dilemma game. How is
this so? It is important, you remember, for a bird to pull off his own
ticks, but he cannot reach the top of his own head and needs a
companion to do that for him. It would seem only fair that he should
return the favour later. But this service costs a bird time and energy,
albeit not much. If a bird can get away with cheating—with having
his own ticks removed but then refusing to reciprocate—he gains all
the benefits without paying the costs. Rank the outcomes, and you'll
find that indeed we have a true game of Prisoner's Dilemma. Both
cooperating (pulling each other's ticks off) is pretty good, but there
is still a temptation to do even better by refusing to pay the costs of
reciprocating. Both defecting (refusing to pull ticks off) is pretty bad,
but not so bad as putting effort into pulling another's ticks off and
still ending up infested with ticks oneself. The payoff matrix is
Figure B.
But this is only one example. The more you think about it, the
more you realize that life is riddled with Iterated Prisoner's Dilemma
games, not just human life but animal and plant life too. Plant life?
Yes, why not? Remember that we are not talking about conscious
strategies (though at times we might be), but about strategies in the
'Maynard Smithian' sense, strategies of the kind that genes might
preprogram. Later we shall meet plants, various animals and even
bacteria, all playing the game of Iterated Prisoner's Dilemma.
Meanwhile, let's explore more fully what is so important about
iteration.
Unlike the simple game, which is rather predictable in that
DEFECT is the only rational strategy, the iterated version offers plenty
of strategic scope. In the simple game there are only two possible
strategies, COOPERATE and DEFECT. Iteration, however, allows lots of
conceivable strategies, and it is by no means obvious which one is
best. The following, for instance, is just one among thousands:
'cooperate most of the time, but on a random 10 per cent of rounds
throw in a defect'. Or strategies might be conditional upon the past
history of the game. My 'Grudger' is an example of this; it has a good
memory for faces, and although fundamentally cooperative it defects
if the other player has ever defected before. Other strategies might
be more forgiving and have shorter memories.
Clearly the strategies available in the iterated game are limited
only by our ingenuity. Can we work out which is best? This was
the task that Axelrod set himself. He had the entertaining idea of
running a competition, and he advertised for experts in games
theory to submit strategies. Strategies, in this sense, are
preprogrammed rules for action, so it was appropriate for contestants
to send in their entries in computer language. Fourteen
strategies were submitted. For good measure Axelrod added a
fifteenth, called Random, which simply played COOPERATE and
DEFECT randomly, and served as a kind of baseline 'nonstrategy',
if a strategy can't do better than Random, it must be
pretty bad.
****************
I found that the results from Axelrod's competition, as well as his followup competition, were quite illuminating.