I recently came across a radio lecture given by Dr Lee Alan Dugatkin on 7.6.2007, titled "Is Goodness Natural?" It deserves comment. (An article on the same subject but with some differences in text was published at Huffington Post.)

The talk began well with some historical background describing the attempts by Charles Darwin, Thomas Huxley, Peter Kropotkin and W. D. Hamilton to explain the origin of goodness, (in the sense of being nice to one another,) in light of evolutionary theory. He concluded that the first three had failed to adequately explain goodness, (Kropotkin’s great work “Ethics” was obviously overlooked) but that Hamilton had solved the dilemma with a “simple but elegant mathematical equation.”

According to Dr Dugatkin, Hamilton’s equation, (c<r b, where c is the cost to the donor, b is the benefit to the recipient, and r is the degree of relatedness,) known henceforth as Hamilton’s Rule, shows that “if enough relatives receive benefits from altruism to outweigh the cost of altruism, then altruism spreads; otherwise, it does not.” (Keep in mind that altruism in the definition used by biologists refers to an act that benefits the recipient’s capacity to survive or reproduce while adversely affecting the altruist’s capacity to survive or reproduce. It should also be noted that Dugatkin did not explain that distinction to his audience, and made matters worse by equating altruism with any act of goodness.)

There’s a problem with Hamilton’s Rule that Dr Dugatkin has perpetuated with this explanation. It assumes that altruism is restricted to acts between relatives. This is not the case, no matter which definition is used. It also assumes that altruism is spread genetically and therefore has a deterministic aspect. This is obviously incorrect. Humans in particular can be influenced by example, that is, by receiving from others or by observing altruistic acts. He continued; "Phrased in the cold hard language of natural selection, blood relatives are worth helping in direct proportion to their genetic (blood) relatedness, weighted by how great a benefit they received."
Here we are again, dealing with that concept from another universe, proportional altruism. It is no doubt true that from a gene’s point of view the statement is correct, but this exposes the folly of the gene-centric view of evolution. Genes do not have a point of view! To look at this from the gene level is ludicrous. Genes are not giving and receiving. Organisms are the participants, and organisms do not recognize proportional altruism as described.

Of course proportional altruism does have some credence, but it’s not related to kinship. In the normal course of events we are generally altruistic by nature. We would rather be nice than nasty. It’s easier to smile than snarl. Who can we most easily be nice to? Those closest to us. Who’s closest to us? Our relatives. So altruism is directly proportional to physical distance, and kin, being closest, are the major beneficiaries. This also explains why we are occasionally helpful to strangers. It’s not rocket science. Is this the full story of altruism? Probably not, but at least it’s grounded in reality.

Then came the highlight of the presentation.  “…Literally thousands of experiments with both nonhumans and humans show the power of Hamilton's Rule.” (Wow, I’d like to see that data!) Now hold on to your seat. He continued “This little equation is evolutionary biology's version of e = mc2.” (emphasis added.)
Please excuse me for a moment while I regain my composure.

There is no justification for this sort of exuberance in a science program. If the good doctor wants to be a journalist that’s fine, but if he wishes to be regarded as a scientist of merit he needs to conform to certain standards. In a presentation to a mass audience generalizations can be acceptable, but propaganda cannot.
Even a non-mathematician can see that these equations have nothing in common.
First, Einstein’s equation has a constant as its base; it has a foundation while Hamilton’s is a collection of variables. Second, Einstein’s equation has a symbol for equality, so precise values can be derived from it. Hamilton’s has no such symbol, so nothing certain can be derived from it.
Then there’s the problem of process. “Einstein's theories sprang from a ground of ideas prepared by decades of experiments.” (from The Einstein Exhibit, Centre for History of Physics, American Institute of Physics) Hamilton on the other hand, tells us that when he began his biology studies he was dismayed to find group selection was the dominant theme of evolutionary studies, he apparently saw selfishness or individualism as the dominant force in the natural world. In other words, Einstein’s theory grew out of the scientific method, while Hamilton’s equation grew out of a preconceived notion the origin of which is debatable, the validity of which is untestable, and for which data was sought after the event. This sorry tale is evidence, if any was needed, that there is not only no comparison between the two equations, but if this is the best that biology can come up with then there is also no comparison between physics and biology. 

Secure in the knowledge that he had satisfactorily explained most of the altruism “problem,” Dr Dugatkin rounded the total picture off with a brief explanation of “reciprocal altruism.” “Hamilton's Rule, of course, does not explain all altruism. Another large chunk of goodness falls under the category of "reciprocity". Individuals are sometimes willing to be altruistic to someone now in the expectation that they will, in turn, be helped when they need it.”
Unfortunately no further details were given as to the workings of reciprocal altruism but the audience was assured that “Following up on work done by Trivers in the early 1970s, in 1981, Axelrod and Hamilton used the mathematics of game theory to predict when so-called "reciprocal altruism" should evolve. Again, scores of empirical studies have followed up the model. Reciprocity can be complex, but an evolutionary perspective has cleared the path to understanding, just the same way it did in the case of blood kinship and altruism.”
Should we accept that at face value?

Let’s look at an online lecture course (from W.W. Norton) that deals with the matter:
   Kin selection is not the only reason why cooperative and seemingly altruistic behavior may have evolved. Reciprocal altruism - the continued, mutually beneficial interaction between individuals over time - is another behavioral strategy that has been tested in a simulated evolutionary forum. For reciprocal altruism to work as a strategy, several conditions are required:
* frequent interaction
* recognizing individuals
* remembering past interactions with individuals
* assisting only those who provided past assistance

There are some problems with this outline, as with Dr Dugatkin’s.
It’s assumed, without justification, that a reciprocal altruism system exists, and that the system is based on selfishness, as the outlined conditions clearly show. The conditions flow into each other with ongoing personal benefit being the overarching condition. If you cannot recognize individuals, cannot remember who helped you in the past, the system breaks down. Remember that this hypothesis has been put forward to explain an uncomfortable (for some) reality; the existence of altruism in animal societies. For the hypothesis to be accepted it must be shown that it occurs regularly in nature. All of the conditions outlined do occur in social interaction. We do remember acts of kindness and acts of greed, and we act accordingly in the future. But does that constitute a system or a strategy? Not at all. These processes go on at the margins. They are the mopping up that goes on at the edges of the main game, and the main game is mutual aid.

How does mutual aid differ from reciprocal altruism? For reciprocal altruism to be consistent with the accepted definition of biological altruism it must involve a serious loss to the altruist. But this rarely occurs in nature. A child rarely receives from its parents to such an extent that the ability of the parents to survive and reproduce more children is impaired. A donor to charities rarely gives to the extent that the capacity to make a living is impaired. Altruists give that which they have in excess. They give that which does not affect their biological fitness. Altruists cannot be exclusively altruistic as this would be suicidal, so what they engage in is mutualism. Reciprocal altruism, “you scratch my back and I’ll scratch yours,” seized by selfish gene theorists and redefined as way of getting around the “problem” of altruism, is no more than old-fashioned mutual aid. Most of the reciprocity that occurs in nature and in society does not involve a loss to one party; it involves a benefit to both parties. That benefit might not always be equal, it probably rarely is, but the benefit still exists for both. The process of mutual aid can take various forms, the most common of which, and therefore least noticed, is known as division of labour. 
Division of labour (I’ll give you a side of bacon, you give me a bag of flour,) is a form of reciprocity that has had a most profound effect on evolution. It has allowed for inequalities in abilities to contribute to the social good, i.e., the biological fitness of the group, and can be observed not only in human societies, but also as an example, in the hunting techniques of lions. It is at the edges of that framework, and of other forms of altruism, that the conditions wrongly attributed to reciprocal altruism in the above quote come into play. Reciprocal altruism as portrayed by selfish gene theorists is no more than a concoction designed to divert attention from the cooperation that underpins life itself. And because Hamilton’s Rule is also based on this self-sacrifice definition of altruism that exists principally in theory and rarely in practice, it falls into the same category. Non-scientific, pretentious, inconsequential and irrelevant.

It’s worth noting at this point that Dr Dugatkin’s claim that genetic relatedness is a pre-requisite for altruism is not merely a personal idiosyncrasy on his part, it’s a widely held view (see for example Foster et al 2006?) that came directly from Hamilton. The Antarctic penguins that mass together in thousands for protection from icy storms not only put a dampener on this but also on Hamilton’s other great misapprehension, his treatment of so-called “selfish herds.” According to Hamilton and all who followed blindly, herding occurs for purely selfish ends, so there should be jostling among the penguins for a position inside the group away from the cold, with the weakest being forced to the perimeter. Instead, there is an orderly rotation to the outside of the group, with all taking their turn in protecting the rest without regard to relatedness. 

How did Hamilton get it so wrong? The answer to that can be found in a description he gave in 1988 for the development of his 1964 paper The Genetical Evolution of Social Behaviour, in which his equation first appeared. He described the influence of R. A. Fisher on his thinking and made the comment; “Clearly it was Fisher who had thought out his Darwinism properly; where interpretations differed, therefore, he must be right, but were the others always wrong? I started on what seemed the key theme in this puzzle – altruism. Did it exist? Could one evolve it in a model?”
The final two sentences are wonderfully revealing of how Hamilton’s mind worked. He is talking of thoughts that went through his mind when in his late twenties. Hamilton had reached that age and did not know if altruism existed? Let’s cut him a little slack and assume that what he meant was; did it exist in the generally accepted form. To put it another way, does a form of altruism exist in nature that has gone unnoticed by naturalists. At that point Hamilton should have made the decision to go bush for a year or two to find the answer. (Even the Antarctic would have done!) Instead, he looked for the answer in mathematics, pressed on to a conclusion, submitted his paper and the rest, unfortunately, is history.

To summarize, Hamilton’s Rule fails at the first hurdle. There’s no need to even discuss his preposterous degree of relatedness and the alleged effect of that on altruism, because the definition of altruism on which it’s based does not occur often enough in nature to be regarded as a regularity. It’s an equation about nothing. The overwhelmingly dominant form of reciprocity in nature is seen in mutual aid, so logic tells us that if we wish to reach conclusions about the impact on evolution of altruism or goodness or cooperation, then it’s to that area we must look. 

So, is goodness natural?
It certainly is, but that’s a story for next time.