There doesn't seem to be a lot of footage of William Hamilton out there. However, here is one video where he explains the basic concept of inclusive fitness:
The existence of altruism in nature can be explained by thinking about the replication of genes. We need to descend to the level of the gene, rather than the individual, in order to see that the gene exists surrounded by copies of identical genes that exist in all its relatives - in particular in its close relatives, its siblings, who have a half chance of carrying a copy of that particular gene, its offspring, which also have a half chance, parents: a half-chance, cousins: one eighth, etc. Seeing this swarm of genes that exists around a particular one, we can then ask what is the behavior caused by this gene that is most likely to cause the propagation of this set of copies in the relatives around it.
Inclusive fitness is a key idea in modern evolutionary biology - with substantial explanatory power. It is a general framework which covers both altruistic behaviour towards kin and what was once referred to as being group selection. However, so far only a few people have applied the idea to memes.
Perhaps people are thinking of memes as being "like viruses" - and viruses don't have much in the way of parental care and can't easily recognise their relatives.
However, as it turns out, this line of thinking is wrong. The exact same argument that Hamilton applies to genes in this video can also be applied to memes - with similar significance and far-reaching implications.
Just as individuals tend to behave altruistically to those with shared genes, so they tend to behave altruistically to those with shared memes. We see that in police, military and religious groups. Shared memes are also a big part of why your computer cooperates with your printer. Inclusive fitness is a general aspect of Darwinian evolution. It is not confined to genes - it also applies to memes - resulting in "memetic inclusive fitness". There is, in fact a considerable scientific literature on this topic - however, it doesn't tend to use the terms "cultural kin selection" or "cultural inclusive fitness" - and instead frames the issue in other ways.
Cultural kin selection is the primary explanation for cultural nepotism and cultural eusociality.
I plan to rectify the neglect of cultural kin selection (or "memetic kin selection" or "kith selection" as it is sometimes called) in my forthcoming book on memes - which will have a large section on the topic.
Hi! I'm Tim Tyler and this is a brief review of this book:
SuperCooperators - Altruism, Evolution, and Why We Need Each Other to Succeed by Martin Nowak and Roger Highfield.
Martin Nowak is a Harvard biologist. His book is about why humans
cooperate as much as they do - which has been a classic puzzle to
evolutionary biologists for decades.
Nowak's number one answer seems to be "game theory". Nowak has worked on this topic with others in the field - including William Hamilton and Ed Wilson. In combination with Karl Sigmund, he came up with some of the ideas behind what he calls "indirect reciprocity".
The book has a chatty style that tries to weave the stories of those involved into the science. That might make the book easier to read for some, but it also makes getting at the scientific content harder. I found this aspect of the book pretty irritating. The author also focusses a lot on their own work and does not pay sufficient heed to the work of others.
Despite the author's reported enthusiasm for mathematical models, the book is almost entirely devoid of mathematics. I don't think I found a single equation - though there were some "sigmas" lying around.
Now on to some of my more serious technical issues:
The book starts out with chapters covering the basic mechanisms that the author claims produce cooperation. These are: direct reciprocity, indirect reciprocity, spatialisation, group selection and kin selection.
Spatialisation relies on reciprocity and kin selection to operate - and so barely deserves its own category, in my opinion.
Indirect reciprocity is a bit of a dustbin category - that mixes together some very different kinds of effects. I think it is better to separate these out - into virtue signalling - which is responsible for reputations and other ways of signalling niceness and overgeneralisation.
The list fails to mention the possibility that cooperative behaviour may have been adaptive in the past - when humans lived in close-knit tribes where kin selection and reciprocity may have played a bigger role. That's a pretty important scenario, in my opinion.
The list also fails to mention manipulation. Manipulation is a key mechanism promoting cooperation. If an agent takes a hit to help another agent, they may simply be being manipulated. They could be being manipulated by the other party, one of the other party's friends or relatives, or by one of their own enemies.
The list also fails to mention the symbiosis theory of cooperation. Parasites and symbiotes may manipulate their hosts into friendly interactions - since they depend on interactions between their hosts to spread.
Lastly, the list also fails to make any mention of the role of culture in forging human cooperation. The author has some awareness of this issue - since he has a chapter on the importance of language. Also he writes:
I do not restrict the use of the term “natural selection” to genes alone. Depending on whether we talk about cells, animals, or people, reproduction can be genetic or cultural. In the former case, successful individuals leave more offspring and pass more genes on to future generations. In the latter, successful ideas, fashions, and strategies spread by imitation and learning: a fad is born.
However, the treatment of culture in the book is cursory. Nowak views culture as facilitating reciprocal relationships via gossip and helping to distribute reputations. This limited perspective is unfortunate - since cultural transmission explains why modern humans are more cooperative than beasts - and the effects of culture aren't just down to reciprocity. In fact, cultural transmission is really what brings the symbiosis theory of cooperation to life. Memes manipulate their hosts into coming into close contact with each other - in order to facilitate their own reproduction. There are also cultural versions of kin selection and group selection to consider - in which these phenomena act on related memes and groups of memes respectively.
So, Nowak's list of mechanisms of cooperation misses out a lot that is significant. This book needs complete rewrite to account for the forces that actually result in human cooperation.
Nowak became infamous recently - in a bust up over kin selection. He wrote an article with E. O. Wilson about kin selection and group selection, in which kin selection got bashed. The article was called "The evolution of eusociality" and it was published in Nature. It was poorly written and was widely ridiculed. However, the group selection controvery and the role of E. O. Wilson in it is turning out to be an interesting chapter in the history of biology - and Wilson is publishing his own book on the topic later this year. One of the reasons I decided to read SuperCooperators is to see what Nowak had to say for himself on the topic in his book.
My council for any others thinking of doing the same is: don't bother. The section in the book bashing kin selection is pretty dreadful. Nowak doesn't say anything particularly interesting or insightful in the chapter that isn't in the "eusociality" paper. Rather he reinforces the impression that he doesn't know what he is talking about. He belittles Price's equation as being "tautological" - which seems to be a particularly stupid criticism to me. The chapter on group selection isn't much better. Group selection is an interesting topic - but I don't recommend learning about it from Nowak. He mostly just talks about the group selection models he personally built - without giving enough information for anyone to actually reproduce them. That is not worth very much, in my opinion.
In the final chapter, Nowak proposes that "natural cooperation" be added to Darwin's "mutation" and "natural selection" principles. I think that this suggestion doesn't make sense. Instead: production, elimination, mutation, symbiosis, synergy, self-organisation, learning and engineering are a much better list of fundamental evolutionary principles.
In the final chapter, Nowak explains why a scientific understanding cooperation is important for building cooperative enterprises - and that mastering cooperation is an important key for humanity when facing challenges in the modern world.
That all seems to be correct - but we need much better science that what Nowak has to offer in order to properly understand why humans cooperate.
There are a few interesting bits in this book, but probably its most salient feature is the way that it misses out so much that is of vital importance to the subject it is discussing.
Hi! I'm Tim Tyler and this is a brief review of this book:
The Imitation Factor: Evolution Beyond The Gene by Lee Alan Dugatkin.
Dugatkin is a biologist and science writer. His specialist field seems to be the mating behaviour of guppies. Mating bahaviour is interesting from the perspective of cultural transmission - since chooseing a mate is a very important decision that it is difficult to get right - and many creatures may well use social cues to help them decide on the best mate.
In the book Dugatkin describes in considerable detail the evidence that shows that guppies do, in fact copy each other's mate choice. This involves experiments with mirrors, fake male guppies attached to sticks, and other paraphernalia.
The book is mainly concerned with the issues of animal culture and animal teaching. It came out in the year 2000 - and the subject has exploded since then, with many new publications. As fellow reviewer Herbert Gintis says, it is rather strange that Dugatkin makes no mention of: "The Evolution of Culture in Animals" by John Tyler Bonner from 1983 - which is an important previous landmark in this area.
There are some studies relating to human mate choice in the book as well, but that isn't the focus. The book is pretty well written and readable. However, the topic involves a lot of descriptions of scientific experiments - which is an intrinsically dry subject area for many. You have to be pretty interested in the topic to want to read the book. Fortunately the topic is a key issue in biology, so there will be some who are interested.
There's a chapter on defining culture - where Dugatkin endorses the definition of Boyd and Richerson from 1985 that confines culture to information transmitted by imitation and teaching - and excludes other forms of social learning. I don't really approve of defining culture that way - but defining culture remains a controversial issue.
The book has a whole chapter on memes, which serves to put the author's ideas in context. A few months before his book was published, Dugatkin published a "counterpoint" to Susan Blackmore's Scientific American article titled "The power of memes" which was called: "Animals Imitate, Too". One of the flaws of Susan's book was that it was very human-centric. It played down the abilities of animals to imitate - and positioned memes as the factor that had transformed beasts into men. Memes did do that - but it is important to note that animals have cultural transmission - and thus memes - too.
Here is what Dugatkin wrote in that response article:
In my work as a behavioral ecologist I have run across dozens of other examples of animal behavior that fit the definition of a meme, and I would not be surprised if the total number were quite large. Memes may be older and more fundamental to biological evolution than Blackmore or anyone else has argued to date. More specifically, the difference between animal and human memes may be quantitative rather than qualitative. Memeticists may well take hold of the idea that animal memes are real and use this to bolster the claim that memes truly are a universally important force in evolution. But if memes do not separate us from animals, as Blackmore suggests, then they alone cannot explain why human culture is uniquely advanced.
The chapter in the book on memes is pretty good. Dugatkin closes by quoting Richard Dawkins on the power of memes (the bit where Dawkins says that memes may leave genes panting behind) - and then he says:
Memes may have the power that this quotation implies, but even so, the more profound point is that they may be older and more fundamental than Dawkins, Blackmore or anyone else has tended to think.
That pretty neatly encaspulates the main theme of the book - that culture is an ancient and ubiquitous phenomenon in the animal kingdom. These days, there's even evidence for cultural transmission among small insects like ants and flies - which have pretty limited cognition.
I think it is fair to say that Dugatkin's ideas about memes have withstood the test of time. We can now see that memes are ubiquitous in the animal kingdom, just as Dugatkin claimed that they were.
I review "Companion Book Summary for The Origin of Cultures" by John Lin (2011).
This is a 120-page summary of the much larger book 'The Origin of Cultures'. The "Companion Book Summary" contains an introduction and a summary of each chapter.
How long can this go on for? Are memes a temporary fad, or a lasting phenomenon? Will we see peak meme - and if so: when?
Memes are probably not a fad - and peak meme may be unlikely to arrive for a long time - but we could see the rise of memes start to level off a little at some stage.
It is pretty hard to say what will happen. Memes do have tremendous significance in terms of the future of the evolutionary process. At some stage, we may see an extended explosion of memes - as cumulative cultural evolution itself snowballs. However, what we are really seeing now is just the term "meme" snowballing. The rest of culture is snowballing too - but not with quite the same rapid doubling time. So: the snowballing of the term "meme" may well slow down a little - as it gets closer to saturating the population - and before the rest of culture itself starts to snowball more rapidly. However, when these events will happen is not clear.
The "meme" terminology explosion could go on for a while yet - it seems.
Young minds are more vulnerable to memetic infections. They are programmed to nature to absorb memes less critically - and have had less time to build up a memetic immune system. This is a kind of double vulnerability. So, it is no great surprise to see the nation's youth gripped by meme fever. This is mostly a recent phenomenon, though. Major news outlets have coverage:
Memetics - The first legitimate field of study effectively obliterated by the internet's power of irrelevant content.
...and...
It seems like everyone on the Internet acts like a 14 year old troll. Can I haz my culture back? I don’t think that lolcats are funny. They might as well symbolize the death of memetics. The fittest memes are parody macros. Can we say cyber-idiocracy?
One benefit I expect is that we will see fewer critics doubting the existence of memes. As late as 2005 we had comments like this (from Maurice Bloch):
[Memeticists] will bring up the originality of thinking of the evolution of culture from 'the memes' point of view'. And, of course, they are right, because if they had been able to argue that there were such things as memes, this would have been a fascinating new perspective on human history. The point is, however, that they have not succeded in arguing convincingly - any more than the diffusionists had before them when talking of 'traits' that there are such things in the world as memes. And so, talk of invasion by the 'body snatchers', to use Dennett's delightful phrase, is an idea as intriguing, as frightening and as likely as invasion by little green men from Mars.
I think one of the positive effects of the modern internet meme explosion is that doubting whether memes exist will look increasingly stupid. Now that memes are a ubiquitous pop-culture phenomenon, the audience will increasingly just think: of course memes exist - duh! So: the frequency of this criticism seems likely to go down.
The modern meme explosion also seems bound to bring new blood and faces to memetics. Yes, it associates memetics with junk pop culture, but I think that is inevitable - and not to be shied away from too much. Marketers and advertisers will trace the term back to its roots, and get turned on to the topic that way.
The meme explosion may well eventually lead to a memetics explosion.
After all, there's got to be a scientific field that studies memes - right?
This critic says that he is "putting the nails in the coffin" of memetics.
However, his main complaint is that memetics isn't original. He says it is reinventing a basic wheel, and that memes are just "ideas", "words" or "concepts".
He has a couple of follow-up videos on the topic here and here.
I posted some messages in the comments to try and straighten things out - a few of which are below:
Comments
On the difference between memes and ideas:
"Ideas" and "concepts" are different from "memes" - at least as these terms are usually defined. Memes are usually defined as being socially-transmitted entities - while ideas and concepts can be private.
On the virtues of using the term "meme":
The other main virtue of using the term "meme" (rather than "word", "idea" or "concept") is that it explicitly suggests the link to genes, biology and evolution. That is good - if you want to discuss things in that context (e.g. memetic drift, memetic engineering, etc).
Modern usage of the term "meme" reflects the links to biology and epidemiology - in that highly spreadable aspects of culture are the ones that tend to get described as being "memes" most frequently.
On whether memetics really was obvious at the time:
It isn't obvious that culture evolves along Darwinian lines. Serious scientific study of Darwinian cultural evolution barely began until the 1970s. Apart from "The Selfish Gene" there weren't really any serious books on the topic until the 1980s (see the "Memetics Timeline" for more detials). It is easy to claim that memetics is obvious - with hindsight - but you do then need to explain why it was that scientists after Darwin didn't cotton on to the idea for a hundred years.
On the competition over the term "meme".
Also, perhaps check out the "meme synonyms" article online. There were many other previous attempts to find a word to refer to a small chunk of human culture. They underwent a process of cultural competition and selection. Now, the term "meme" now appears to have pretty comprehensively won. The idea was not a new one at the time, though.
On the idea that Nietzsche had similar ideas before memetics.
I see that there is a book titled: "Nietzsche's New Darwinism". One reviewer says: "Nietzsche's perspectival metaethics not so much as a rejection of natural selection, but an addition to natural selection of social selection and self-selection by those powerful enough to do it. In the former sense, Nietzsche may be a forerunner of modern notions of memetics or gene-culture coevolution". The knitting together of Nietzsche and memetics appears to have already begun.
On how the "cultural symbionts" of memetics differed from other historical concepts that were similar:
It is true that the idea of "symbiotic cultural organisms" is part of ancient folklore - in the form of posessions and exorcisms - but those folk didn't express their ideas scientifically, or generalize them, to all of human culture. It is also true that this idea is present in Cloak (1975) and Auger (1952) - but Dawkins (1976) popularised it.
On what new things memetics offered:
Memetics introduced novel (and useful) terminology, the idea of symbiotic cultural organisms (akin to viruses), the idea of "replicators" and it emphasised the applicability of Darwinian evolution to culture. The "replicator" idea has caused much confusion, but its other introductions remain true and valuable.
Concluding remarks
It's rather nice to have some modern critics saying that memetics is obvious. That's the kind of criticism I can get along with!
A more common complaint is that memetics is wrong. For instance, here is Maurice Bloch (2005):
[Memeticists] will bring up the originality of thinking of the evolution of culture from 'the memes' point of view'. And, of course, they are right, because if they had been able to argue that there were such things as memes, this would have been a fascinating new perspective on human history. The point is, however, that they have not succeded in arguing convincingly - any more than the diffusionists had before them when talking of 'traits' that there are such things in the world as memes. And so, talk of invasion by the 'body snatchers', to use Dennett's delightful phrase, is an idea as intriguing, as frightening and as likely as invasion by little green men from Mars.
Perhaps we can put the "memetics is wrong" folk and the "memetics is obvious" folk together in a field and let them duke it out among themselves for a while - as an inexpensive method of whittling down their numbers a little.
In a video interview between John Maynard Smith and Richard Dawkins, Maynard Smith suggests that the distribution of parthenogenesis on the twigs of the tree of life shows that sexual recombination is a trait which is often maintained by species-level selection. The video is called: Explaining the maintenance of sex.
Maynard Smith goes on to explains why there's no parthenogenesis in conifers and mammals in another video, called: Are mammals stuck with sex?
Are there other traits which appear to be maintained by species-level selection? John Maynard Smith suggests viviparity as another possibility - in a video called: Viviparity may be a big evolutionary mistake!
Small size is another obvious candidate. Meteorite strikes regularly kill all species with individuals larger than a mouse. So there is systematic species-level selection for being small. Living in the deep ocean is probably favoured by species-level selection on similar grounds. Of course, these are cases where species-level selection gets rapidly swamped by other forces.
The other candidates that spring to mind involve traits that result in speciation. Easily evolving incompatible songs, habitats, or genitalia are other possible candidates. A tendency to swim off to islands might produce new species.
Culture has species too - or at least it has groups of cultural entites that cross easily among themselves but harder with outsiders. Outcrossing is never entirely prohibited in the realm of culture, but things like Cobol programs or origami designs are pretty closely analogous to cultural species - in that they recombine with each other a lot more than with the rest of culture. Is there cultural species selection? So far, we don't seem to have a lot of data on the topic. However, because species boundaries are weaker, cultural species selection may be a less significant force than species selection is in the organic realm.
What I didn't know - until recently - is that David subsequently did a rapid turnaround on the topic - and documented it in a series of blog posts. Here they are:
It is interesting to see the gears turning. In one article he points how different genetic and memetic evolution are - because genes are adaptive for their hosts, but memes are not necessarily - but then it is as though the penny drops as he is writing - and he says that:
There might be a closer analogy between memes and viruses, which are essentially disembodied bits of DNA, free to skip from one body to another. Dawkins himself makes the comparison with viruses, but does not pursue its implications as far as I would wish.
That is almost correct - memes are a lot like the heritable information of viruses - at least in the case of memes that are deleterious to their hosts.
At the end he says:
After recent discussion of cultural evolution I realised that I didn’t know much about memes, so I set myself the penance of reading Susan Blackmore’s The Meme Machine.
...and...
Still, it’s an important book, and well worth reading. Overall, it left me with the feeling that memes do need to be taken seriously, but only as one aspect or dimension of cultural evolution.
Not too shabby an awakening, overall. I'll have to annotate future citations of his criticisms to say that he eventually came around.
References to 
Memes go "viral" in freshman minds, it seems.
Some question whether 
In a video interview between John Maynard Smith and Richard Dawkins, Maynard Smith suggests that the distribution of parthenogenesis on the twigs of the tree of life shows that sexual recombination is a trait which is often maintained by species-level selection. The video is called: 
I've previously cited 
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