In my 2011 video/essay title " Dawkins Dangerous Idea", I approvingly quoted Paul McFedries as saying:
Richard Dawkins became famous in the 1970s for his concept of the selfish gene, and he has become infamous in recent years for his unyielding atheism. But I predict that Dawkins will be known, a hundred years hence, not for these contributions to science and culture but for the concept of the meme. Feel free to spread that idea around.
Now it appears that genetics blogger Razib Khan has come around to much the same idea, writing an article titled:
In 2546 Richard Dawkins Will Be Remembered For “Memes”
I still think that this is right. What is Dawkins second-biggest scientific idea? Probably the extended phenotype. That seems rather insignificant compared to memes and memetics.
 Here's the late Marvin Minsky (1994) in " Will Robots Inherit the Earth?":
In the past, we have tended to see ourselves as a final product of
evolution - but our evolution has not ceased. Indeed, we are now
evolving more rapidly - although not in the familiar, slow Darwinian way.
It is time that we started to think about our new emerging identities. We
now can design systems based on new kinds of "unnatural selection" that
can exploit explicit plans and goals, and can also exploit the inheritance of
acquired characteristics. It took a century for evolutionists to train
themselves to avoid such ideas - biologists call them 'teleological' and
Lamarckian' - but now we may have to change those rules!
That's more or less what I have been trying to do over the last decade:
drag the theory of evolution into the 21st century by incorporating
intelligent design, Lamarckain inheritance, directed mutations,
evaluation under simulation and so on.
One of the things I have found is that these things are often not quite
as novel an Minsky implies. Organisms have been "inheriting acquired
characteristics" for at least as long as dogs have been passing their
fleas on to their puppies. Plans and goals are not exactly new either.
The first mammals were making plans - and these went on to influence
their evolution via sexual selection and in other ways. The picture
of these new capabilities arising with human engineering design is
not really correct - many of them have much older roots.
IMO, this is interesting because it makes the old school evolutionary biologists
and their textbooks wrong in their own terms, not just because of
human beings, genetic enginnering, etc.
Here is S. J. Gould in 1996 on why memes won't work (35 minutes in):
I think Brenda put her
finger on exactly why the meme concept
is bankrupt and I don't think it's going to
get very far although try it by all
means [...] I don't believe I'm a
First Amendment absolutist in U.S. terms:
pursue whatever you want [...]
but it's so central in science to
distinguish between metaphor and
mechanism. Metaphors are not useless, the
Gaia metaphor is a non mechanistic
statement that has some utility. To me
memes are nothing but a metaphor and
they're a metaphor based on a
fundamentally false view of consciousness
and I think that's why it isn't going to
work. You see it's ultimate Western
reductionism to have a notion of the
meme you have to be able - as you can for
genes because they are physical entities [...]
you have to be able to cash
out the notion of "meme" to divide the
enormous complexity of human thinking
into items, items that have a certain
hardness, that have a certain
transmissibility, but human thought is
not that, it's not breakable up into tiny
little hard units - everything interpenetrates.
The only thing memiec analysis has ever
been any good for are things that are trivial like changes
in hairstyles and skirt lengths because
those are things. The other thing is
you'll never be able to work a Darwinian
metaphor because the Darwinian mechanism
requires random variation. Memic
variation is not random there's no way on
earth it is ever going to be that's why
every attempt - and memic thinking is not
the first, there's a whole history of this - every
attempt at so-called evolutionary epistemology -
that is: to make a Darwinian
evolutionary epistemology - has failed because
you will never get the fundamental
characteristics of the Darwinian
mechanism: random variation and the
natural selection of random variants.
Mind directs its items, and there are
no items! As soon as you have the
impossibility of breaking down... it's
hard enough for genes - that's why
sociobiology failed because my thumb
length isn't the gene and aggression
isn't the gene and homosexuality isn't a
gene they are complex genetic and
environmental components you can't do it
for human culture, it won't work.
This is the same interview where Gould describes genes as being a "meaningless metaphor" (13:45).
This is mostly of historical interest now, but I think it illuminates some of Gould's confusion about cultural evolution.
IMO, these days, people are less likely to argue that human culture can't be usefully broken down into small units. The internet has comprehensively demonstrated that a very wide variety of types of human culture can in fact be broken down into bits: digital, discrete 1s and 0s.
 It's apparently difficult for people to publicly admit that they were wrong. While cultural evolution has seen its share of criticisms over the years I can think of very few critics who have publicly come around.
One such critic is John Maynard Smith. He wrote a number of somewhat critical reviews of books dealing with cultural evolution.
However, in 1999 he wrote:
I used to regard the meme as a fun idea - helpful in explaining to students that there can be more than one kind of replicator, and that all replicators evolve by natural selection - but not as an idea which could be used to do much serious work. Genes have clear rules of transmission (in sexual organisms, Mendel’s laws) whereas you can learn memes not only from parents, but from friends, books, films and so on. Consequently population genetics can generate precise, testable predictions, whereas it seemed to me difficult to make such predictions about memes. Susan Blackmore’s book, The Meme Machine, has gone some way to changing my mind. Perhaps we can make the meme idea do some work.
Another critic-turned-enthusiast was David Burbridge. I've documented his change of heart in an article titled
David Burbridges meme turnaround.
When I got involved in popularizing memes and cultural evolution I made a confession video available with transcript here: My Memetic Misunderstandings. However such articles seem rare.
This essay starts out with the hypothesis that it is difficult for people to publicly admit that they were wrong. A more sinister explanation for the missing turnarounds on the topic is also possible: people don't change their minds on this issue and take their delusions to their grave with them. Some dead critics confirm that this happens some of the time: Steven J Gould apparently took his delusions about the topic with him when he departed from the world. I hope that this explanation is wrong. Scientists are supposed to be responsive in the face of evidence, not dogmatically attached to their previous views. "I was wrong" is something that scientists ought to be able to take pride in saying.
Before the discovery of the importance of symbiosis, evolutionary theory worked with a few
fundamental operations - prominently including splitting and mutation. The history of life
was viewed as a branching tree with branches diverging, but not converging again, at least
not once they had fully divided. Life had an associated family tree.
Symbiosis represented a significant revolution in evolutionary theory because it introduced a
new operation into the fundamentals of evolutionary theory: merging or joining. If splitting looks like this:
...then joining looks like the time-reversed operation - like this:
Before the 1900s, merging and joining operations were not completely foreign to evolutionary theory.
The fusion of male and female gametes was a find of merging operation, though one that was confined by
species boundaries. It was also known that some creatures lived inside other ones - such as
gut bacteria - and that parasites regularly forged new associations with their hosts.
However such intimate relationships were not widely thought to involve permanent fusion
between previously unrelated organisms. Instead they were mere ecological associations.
The pioneers of symbiosis were mostly Russian. Though they discovered symbiosis early in the
20th century, their theories were ignored in the west until the 1960s, when it was discovered
that mitochondria contained their own DNA lineages. Over the next 30 years, evidence accumulated
that unrelated organisms could fuse together permanently, generating new kinds of recombination
operation distinct from sex. Gaining or losing symbionts could be profound and dramatic
evolutionary events - sometimes triggering speciation. The phenomenon was not confined to
bacteria, but affected organisms of all sizes. We now know that eucaryotic cells are
unions of many free living ancestors, that viruses carry DNA between all kinds of different
species, and that around 10% of the human genome shows signs of origins outside our own
species. Horizontal gene transfer has turned the tree of life into a web. Multi-cellular
organisms are now widely viewed as being menageries.
In modern times, the evolutionary significance of symbiotic unions is familiar to us,
so it is sometimes hard to appreciate the scale of the revolution the idea represented,
and the resistance that it faced at the time. Evolution was generally though of as being
gradual - yet in symbiogenesis, entire genomes could join forces in an evolutionary
instant. This idea violated established dogma and was widely rejected and ridiculed
for over 50 years. An avalanche of facts and data eventually vindicated the idea.
At the time, this was the the biggest revolution in evolutionary theory since Darwin -
eclipsing the discovery of DNA, kin selection and Mendelian inheritance, in my humble
opinion.
In cultural evolution, the story went rather differently. In 1975, Ted Cloak pioneered a symbiosis-based version of cultural evolution, in
which human hosts cultivated cultural symbionts inside their brains. Richard Dawkins went on to popularize the idea
in the 1976 book The Selfish Gene. These cultural "memes" were sometimes characterized as being "viruses of the mind".
However, in academia,
symbiosis was very slow to catch on. Many academics didn't seem to cotton on
to the fundamental concept of a cultural organism - instead opting to
model culture as an aspect of the host's phenotype. This was a bit like
modeling smallpox without the concept of the smallpox virus - and instead talking
about horizontal and oblique transmission of the smallpox rash phenotype. Many
of the researchers in the field are still laboring under the resulting hangover.
Symbiosis enthusiasts were one tribe of biologists and cultural evolution enthusiasts
were another. The intersection included Ted Cloak, Richard Dawkins and Ben Cullen -
but only a few others. What this meant in practice was that concepts such as parasitism,
mutualism, domestication, parental care and arms races tended to get neglected by most
students of cultural evolution.
The slow uptake of symbiology among cultural evolution researchers is an aspect of
cultural
evolution's scientific lag. Most anthropologists long ago demonized evolution
and expelled it from their halls as a dastardly idea that they wanted nothing to
do with. The important topic of cultural evolution subsequently suffered from lack
of attention and funding - and progress has been slow as a result. It wasn't until
the 1980s that symbiology gained acceptance among mainstream evolutionary biologists.
There's clearly still a long way to go before most students of cultural evolution start
applying the idea properly.
 Leigh M. Van Valen (of Red Queen fame) came up with an early expression of Darwinan physics in 1989. He pointed out that natural selection is common in the inorganic world. Here's what he said in Three Paradigms of Evolution:
Then again, look at the rock called granite. It is composed mostly of grains of feldspars and quartz, with some mica and other minerals inserted among them. When granite weathers, the feldspars and micas become clays but nothing much happens to the quartz grains. They are most resistant and get transported down streams or along shores. Thus most beaches are the result of differentially eroded granite. This is an example of natural selection in the nonliving world. Quartz grains survive longer than feldspar grains, and there is a progressive increase in the average resistance to weathering, of the set of grains that have still survived. This action of natural selection is even creative as we see by the formation of a beach. The lack of reproduction imposes constraints on the flexibility of evolution here, but one shouldn't confuse that with the selection itself. We do have here a common sort of evolution by natural selection and there are many other non-living examples.
I've made much the same point in my universal selection essay.
Of course, the case for Darwinan physics is quite a bit stronger than this passage implies - because copying and reproduction are also common in the inorganic realm. Rocks split into smaller rocks, streams split into smaller streams, and so on. There's also evidence of family trees - as seen in diffusion limited aggregation, and optimization and exploring a search space - as when a lightning strike finds the highest point in a landscape. However, Van Valen had some of the important ideas quite early on in the history of the field.
Domestication is surely an important concept for students of cultural evolution. Unfortunately, it first requires the concept of a cultural organism, something that academics seem to have difficulty in swallowing.
Daniel Cloud has written extensively on the domestication of words and language. Cloud credits Dennett with the idea that language could be domesticated - though he argues that Dennett didn't take the idea far enough. The earliest reference to domesticated memes from Dennett I can find is in his 1998 essays Memes: Myths, Misunderstandings and Misgivings AND Snowmobiles, horses, rats, and memes.
Dennett goes on to discuss the idea of domesticated memes some more in Breaking the Spell (2006), writing:
What I now want to suggest is that, alongside the domestication of animals and plants, there was a gradual process in which the wild (self-sustaining) memes of folk religion became thoroughly domesticated. They acquired stewards. Memes that are fortunate enough to have stewards, people who will work hard and use their intelligence to foster their propagation and protect them from their enemies, are relieved of much of the burden of keeping their own lineages going. In extreme cases, they no longer need to be particularly catchy, or appeal to our sensual instincts at all. The multiplication-table memes, for instance, to say nothing of the calculus memes, are hardly crowd-pleasers, and yet they are duly propagated by hardworking teachers — meme shepherds — whose responsibility it is to keep these lineages strong. The wild memes of language and folk religion, in other words, are like rats and squirrels, pigeons and cold viruses — magnificently adapted to living with us and exploiting us whether we like them or not. The domesticated memes, in contrast, depend on help from human guardians to keep going.
However, I notice that Adam Westoby seems to have written extensively on domesticated memes in 1994. He has the idea that memes domesticate humans as well as the idea that humans domesticate memes. Here's his 1994 manuscript. To quote from it:
The memes of theoretical natural science, as Wolpert (1992) points out, are highly "unnatural" memes, remote from "common sense". Like cattle or sheep, they have been bred for generations into the forms preferred by their domesticators (of whom some of the most important are other memes). Testability, generality, uniform vocabulary, unambiguous meaning, internal consistency, and so on - even taken singly such traits are rare memes, and to assemble them all requires long intentional selection. The domesticated memes of theoretical natural science, having embodied such significant adaptations to artificial circumstances, could no longer survive reintroduction to the wild. They can live and breed only with the aid of rather complex arrangements to sustain them. The cultivation of theoretical science (like keeping sheep) has come to rely on auxiliary breeds, such as scientists - rather like sheepdogs, who keep the flock together and bark at intruders. By comparison, much social science consists of more "common sense" memes, less "deformed" by domestic breeding. They more resemble semi-domesticated breeds which forage freely on the mountain slopes in summertime, but are herded in for the winter.
Westoby is the earliest reference to the idea of domesticated memes I have found so far. Is this the true origin story for the idea that memes could be domesticated? Did anyone else come up with this idea earlier? Please let me know if there's an earlier reference that I'm currently missing.
The importance of domestication in cultural evolution is apparently an illustration of the superiority of memetics in this area - compared to other strains of cultural evolution. It looks as though meme enthusiasts got to this idea first - because they have a symbiosis-aware version of cultural evolution. Academics are now picking up the idea (for example, Joseph Henrich's latest book has culture domesticating humans in its subtitle) but they appear to be playing catch-up.
I've seen a fair number of new popular articles on memetics (not just memes) as a result of the internet meme explosion. Here's one of the latest ones, titled memetics and the science of going viral. It's a reasonable article - though academic students of cultural evolution don't even get a mention - and instead we get some links to the author's own content from the field of law. The article notes that even the US president has referenced internet memes - as a testament to their popularity.
I noticed one mistake: he article says that the term “memetics” was first proposed by evolutionary biologist Richard Dawkins in his popular 1976 book “The Selfish Gene”. Chapter 11 of that book does use the term "population memeticist" - but the term "memetics" is usually attributed to Ariel Lucas - following Douglas Hofstadter's attribution in his 1983 book, Metamagical Themas. I'm often surprised how many people go from meme to memetics, completely bypassing the academic literature on cultural evolution, much of which systematically ignores memetics.
 The study of cultural evolution has lagged far behind the study of organic evolution - much to the distress
of its enthusiasts. One question is why there is such a lag. One answer is that the delay was caused, in part, by the modern evolutionay synthesis, which failed to include cultural evolution and is incompatible with it.
The story is that, for one reason or another, the architects of the modern synthesis had little or no understanding of cultural evolution. They failed to incorporate it into their synthesis. Then their synthesis became stagnant dogma, stifling innovation in the field.
I think that this story has considerable truth content, though obviously it isn't the whole story. A possible piece of evidence against the claim that the architects of the modern synthesis had little or no understanding of cultural evolution could potentially be found in the following books:
I haven't properly reviewed this evidence yet. Both books came out significantly after the modern synthesis crystallized, though.
Another problem with the thesis is that it doesn't offer an explanation for why the the architects of the modern synthesis lacked meme literacy in the first place. Maybe the explanation for that is sufficient to account for the lag, without the modern synthesis being to blame. I can't rule that out - but the modern synthesis was influential; it probably didn't help.
Looking at the memetics timeline the first half of the 20th century was an uneventful time for cultural evolution. In fact there are no entries at all from 1915 to 1945 - which is more-or-less when the modern synthesis came together. The founders of the modern synthesis were writing during a bleak time for cultural evolution. Perhaps the world wars were a factor here.
 The term "refactoring" commonly refers to a set of useful computer programming techniques which involve making structural changes to a program without altering its function. Refactoring is often done before making functional changes. If you launch into making changes without some preliminary refactoring, you are often more likely to make mistakes. Refactoring frequently allows you to make changes without breaking tests - so you can verify that your changes don't break anything. If you need to make a bunch of changes, it sometimes helps to make ones that you can verify do no damage first.
I argued in refactoring science that science can benefit from refactoring too. I agree with the many folk who think that Darwinism is in need of some changes. This raises a number of issues about what changes need making. One such issue is where to start from. Darwinism has had two main identifiable releases: Darwin's Darwinism and the Modern Synthesis. Many seem to assume that we should build on the most recent release of the theory, the Modern Synthesis. For example, here's Massimo Pigliucci:
It makes just as little sense to talk of "Darwinism" in modern science as it does to talk about Mendelism or Newtonianism. The current theory of biological evolution is the Modern Synthesis, and if one wants to make the point that cultural evolution works in the same way, one needs to take on board the most refined version available of the theory, not its earliest draft.
Incidentally, I think that's why we should avoid talk of Lamarckism as well as Darwinism altogether: they refer to murky (in the first instance) or outdated (in the second) ways of thinking about biology, and it doesn't help to resurrect them as if the last two centuries of science hadn't happened.
This rather assumes that the Modern Synthesis was an improvement over Darwinism. However, some have argued that the Modern Synthesis consists of a bunch of overstatements. For example, Mesoudi (2011) has argued that cultural evolution is Darwinian, but not Neo-Darwinian. The modern synthesis is often billed as uniting Darwin's idea of natural selection and Mendelian genetics. The problem is that Mendelian genetics only applies to inheritance via DNA. As a result, other forms of inheritance were sidelined by the synthesis. From such a perspective, the Modern Synthesis looks like a bit of a broken version of Darwinism. Failing to encompass cultural evolution is a pretty serious flaw. In this case, maybe we should start from an earlier revision of Darwinism - one without so many problems.
Starting with Darwin's theory should allow cherry picking any useful bits from the Modern Synthesis. Starting from the Modern Synthesis would probably be followed by bunch of 'revert' operations - to get rid of the dogmatic and mistaken bits. The former operation looks easier to me than the latter one.
I have some other concerns about starting from the Modern Synthesis. It is vague. Of course Darwin changed his mind about some things too - and the various editions of "The Origin" describe rather different theories. However, it seems to me that Darwinism is more clearly specified than the Modern Synthesis was. The Modern Synthesis can be difficult for critics to criticize - since there's no canonical version. In science, that's not a good thing.
In some respects, the Modern Synthesis illustrates how not to produce an evolutionary synthesis. It subsequently became a bit of a straitjacket for researchers. Inevitably any new theory of evolution will also be incomplete - but there's no need for it to be dogmatic and mistaken.
Modern definitions of "natural selection" include both differences in survival and differences in reproductive success. E.g. see here.
For Darwin, natural selection was all about the survival and preservation of individuals. Some quotes from Darwin should illustrate the point.
In The Origin of Species, Chapter 4, Darwin wrote:
If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and of procreating their kind? On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed. This preservation of favourable variations and the rejection of injurious variations, I call Natural Selection.
In The Origin of Species, Chapter 3, Darwin wrote:
Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection.
In The Origin of Species, Chapter 4, Darwin wrote:
Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure.
Darwin repeatedly uses the term 'preservation' and repeatedly says he is talking about individuals (rather than traits). In The Descent of Man, Darwin wrote:
The survival or preservation of certain favoured words in the struggle for existence is natural selection.
In a letter to Charles Lyell dated 1860, Darwin regretted the use of the term "Natural Selection," preferring the term "Natural Preservation". He wrote:
Talking of “Natural Selection”, if I had to commence de novo, I would have used "natural preservation".
Of course, Darwin did also refer to differences in reproductive success. He called these "sexual selection". Darwin wrote:
this leads me to say a few words on what I call Sexual Selection. This depends, not on a struggle for existence, but on a struggle between the males for possession of the females; the result is not death to the unsuccessful competitor, but few or no offspring.
For Darwin, natural selection and sexual selection were separate. Most modern authors would say that natural selection includes sexual selection.
Darwin's distinction between "natural selection" and "sexual selection" corresponds broadly to my terms "natural elimination" and "natural production". Darwin's term "sexual selection" is totally unusable in this context - since both sexual and asexual reproduction can result in different numbers of offspring.
In the 1930s the term "natural selection" was hijacked by other evolutionary biologists and redefined away from what Darwin originally meant by it. Here's Satoshi Kanazawa (2003):
In the 1930s, however, biologists redefined natural selection to subsume sexual selection and began to contend that differential reproductive success was the currency of natural selection.
Here is Geoffrey Miller in The Mating Mind, 2000, p.8:
In the 1930s, biologists redefined natural selection to include
sexual selection, because they did not think sexual selection was
very important. Following their precedent, modern biology textbooks
define natural selection to include every process that leads
some genes to out-compete other genes by virtue of their survival
or reproductive benefits. When one biologist says "evolution
through natural selection," other biologists hear "evolution for
survival or reproductive advantage." But non-biologists, including
many other scientists, still hear "survival of the fittest." Many
evolutionary psychologists, who should know better, even ask what
possible "survival value" could explain some trait under
discussion. This causes enormous confusion, and ensures that
sexual selection continues to be neglected in discussions of human
evolution.
I share the concern that this redefinition can be a source of confusion. Darwin is widely credited with the idea of natural selection. I wonder how many students realize that the natural selection they are taught is quite different from the natural selection that Darwin came up with.
Most of the sources I can see that mention this redefinition argue that its consequences were negative, and we should go back to Darwin's conceptual split between destruction and reproduction. That's essentially the split which I argued for in my essay natural production and natural elimination.
Satoshi Kanazawa (2003) says:
I concur with Miller (2000, pp. 8–12), Campbell (2002, pp. 34–35), and others in the current
generation of evolutionary psychologists and believe that we should return to Darwin’s original
definitions and treat natural and sexual selection as two distinct processes. I am fully aware that
this view is still controversial and in the minority, but I firmly believe that the conceptual separation
of natural and sexual selection will bring theoretical clarity to evolutionary biology and psychology.
Geoffrey Miller in The Mating Mind, 2000, p.8-9 says:
In this book I shall use the terms "natural selection" and
"sexual selection" as Darwin did: natural selection arising
through competition for survival, and sexual selection arising
through competition for reproduction. I am perfectly aware that
this is not the way professional biologists currently use these
terms. But I think it is more important, especially for nonbiologist
readers, to appreciate that selection for survival and
selection for attracting sexual partners are distinct processes that
tend to produce quite different kinds of biological traits. Terms
should be the servants of theories, not the masters. By reviving
Darwin's distinction between natural selection for survival and
sexual selection for reproduction, we can talk more easily about
their differences.
 I have long had an entry for "semes" on my meme synonyms page. However, I only recently read up about them. Barry Hewlett, a colleague of Cavalli-Sforza seems to have been one of the main proponents. Here's how one of Barry's papers introduces them:
Following Cavalli-Sforza, we call these units “semes” rather than “memes” (Dawkins 1976, Durham 1991, Boyd and Richerson 1985) because “seme” comes from “sign” and emphasizes the symbolic nature of culture.
Semes were apparently first mentioned in a 1970 book by Roland Barthes called "S/Z" - a three hundred page dissection of a short story written in French.
However, they were subsequently adopted by Cavalli-Sforza, Marc Feldman and others. Feldman explains the term in a 2007 presentation here. He presents "semes" as the cultural equivalent of "genes".
Marcus Feldman said in 2014 (16 minutes in):
I call them semes because I don't like memes, memes gives some impression of biology which I do not want to do and semes comes from semiotics.
Of course, culture is part of biology, so biologically-inspired terminology is completely appropriate.
I think "semes" are more-or-less dead terminology now. Not all culture is symbolic, so the death of "semes" seems to be no great loss. However, IMO, it is fascinating historical tit-bit that Cavalli-Sforza and Marc Feldman join all the other students of cultural evolution that publicly toyed around with "-eme" words.
 In case anyone needs a recap, August Weismann was the guy who cut the tails off of rats. He concluded that a barrier prevented somatic cells from contribution to evolution - now known as the Weismann barrier. The Weismann barrier is a real phenomenon, but people vary on how permeable they think the Weismann barrier is.
There's a corresponding barrier in cultural evolution. Cakes contribute to recipes, but mainly through selection. Anywhere where reverse engineering is complex and difficult, something like the cultural equivalent of the Weismann barrier comes into play. It is hard to dispute the idea that there is less of a barrier in the cultural domain, though. The difference between the domains is partly because there is more reverse engineering in cultural evolution. Genetic engineers can reverse engineer designs originally coded in DNA as well, but for various reasons, they rarely bother. There's plenty of secret computer software in the world, but only a small number of secret genomes. So, people are forced to reverse engineer software from web sites, while few are forced to reverse engineer gene sequences from organic phenotypes.
It's important to recognize that Weismann's barrier is also pretty seriously compromised even if you ignore cultural transmission. As I explained in my article on Lamarckian inheritance in cultural evolution:
Some acquired characteristics are inherited while others are not. If Weismann had chosen a different trait - for example, stress - he might well have drawn the opposite conclusion.
Another problem for Weismann's barrier is genetic engineering. These days a scientist can transfer whatever information they like into the genome. Acquired characteristics, information from somatic cells, detailed sequence information, genes from other species, prime numbers - whatever they like, really. The permeability of Weismann's barrier is increasing as time passes and biotechnology improves.
 Bonobos exhibit a number of human-like traits when compared to common chimpanzees. They are famously more sociable than common chimpanzees. They exhibit a domestication syndrome and have a range of paedomorphic traits. In humans these traits seem linked to cultural inheritance. Paedomorphism gives brains more time to adapt to culture and reduced levels of aggression facilitate cultural transmission. These observations lead to the hypothesis that bonobos have more cultural transmission than common chimpanzees do.
One point which may conflict with this idea is that bonobo brains are significantly smaller than chimpanzee brains - while in humans, culture seems to have made brains larger. Small brains are consistent with the bonobo domestication syndrome, though.
Key data which evaluating this hypothesis depends on involve comparisons of common chimpanzee vs bonobo cultural learning abilities.
There's some comparative data here. It says (on p.27) that bonobos use tools to acquire food much less than common chimpanzees do. Bonobos do have sophisticated social learning skills, though, as Kanzi demonstrates.
If bonobos are more like humans than chimps because they have more culture, then that would be a fantastic discovery from the perspective of students of cultural evolution. It would make bonobos a snapshot of the development of culture in progress - with common chimpanzees providing a baseline.
 Joe Brewer once wrote:
It is a beautiful irony that the “meme” meme won out (as indicated by the amount of attention garnered with it as the label for cultural transmission) using the very evolutionary processes that constitute how cultural evolution actually works.
The story of the "meme" meme is indeed an interesting one. There have been some histories of memetics published. One of the more comprehensive ones is:
I think there's some further scope for analysis of the rise of the "meme" meme.
One of the features of the story is the repeated attacks on the meme concept by confused academics who don't properly understand it. Cultural evolution has featured an ongoing war between academics and popularizers which has, I believe, been a destructive war which has done damage to the field. Presumably, without this conflict, the term "meme" would be even more widely used and understood.
Another interesting feature of the rise of the "meme" meme is its use of memetic hitchhiking on the most viral content on the internet. A massive marketing department probably couldn't have come up with a better plan for promoting the "meme" meme - and yet the whole business with internet memes apparently happened with very little central planning.
Anyway, the rise of the "meme" meme - in competition with the various meme synonyms - is an interesting topic for students of cultural evolution. It's metamemetics! I look forward to the whole topic being given a more comprehensive treatment.
 The Selfish Gene was published 40 years ago. That makes the meme 40 years old.
Matt Ridley has penned an article celebrating 40 years of Selfish Genes. The article is a good one - but it fails to mention the M word.
Memes could turn out to be the most significant contribution made in The Selfish Gene. Dawkins popularized the nascent field of cultural evolution - making it comprehensible to the layman. Five years later four academics published books on the topic, dragging the topic into the academic mainstream, and the rest is history. While the history of cultural evolution long predates The Selfish Gene, retrospectively, 1976 seems like a significant turning point for the field.
Theories broadly based on Darwinian evolution apply at various levels in the brain.
Long-term memories are copied to preserve them. Learned information is repeatedly
rehearsed. Mistakes are repeatedly dwelled on. Promising ideas and plans are copied
in short term memory as variations on them are generated and selectively maintained.
Also, various low level processes behave in a Darwinian fashion: most significantly,
neurite tips are copied with selective retention, and nerve impulses themselves are
copied as they travel down branching axons and up branching dendrites.
I think that it makes reasonable sense to refer to these low-level brain processes
using the term 'Neural Darwinism'. There's a bit of a problem though - which is that
the term 'neural Darwinism' has been widely used to refer to the particular theories
put forth in the late Gerald Edelman's
1987 book titled neural Darwinism.
Gerald Edelman was a pioneer in applying Darwinism to the brain - and I don't want to
diminish his contribution too much. Developmental selection is a reasonable idea.
Experiential selection is a bit more of a dubiously-named idea. I read neural Darwinism
in the 1990s and found it dry, tedious and unconvincing. However, retrospectively,
Edelman's concept of neural reentry
looks important. It could be the key to
understanding how the brain does something functionally similar to
back propagation without having a good quality
bi-directional signal propagation mechanism.
Overall, though, I think things have moved on a bit since 1987. It is now clearer
that there are at least three types of low-level signal copying in the brain: the
conventional reproduction of cells - including neural stem cells, the splitting
of axon and dentrite growth tips, and the splitting of signals travelling how
axons and back-propagating up dendrites. Despite the progress, I think we
can still use the term "Neural Darwinism". I would hate "Neural Darwinism" to
become a reference to an out-dated and discredited theory: it deserves better
than that.
Signals propagation in axons and dentrites has been effectively simulated by
synthetic neural network enthusiasts. However few of them bother simulating
the other main copying processes in the brain: cell splitting during development
and neurite tip splitting. Maybe simulating these Darwinian brain processes
would help to build better synthetic neural networks.
For references see my Keeping Darwin in mind essay.
 Recapitulation theory refers to the idea that organism development recapitulates evolutionary history. As Ernst Haeckel's put it, "ontogeny recapitulates phylogeny". Most biologists would describe Haeckel's ideas as being widely discredited. The story of Haeckel doctoring his embryo illustrations to support recapitulation theory has been widely retold.
Recapitulation theory encapsulates a rather simple truth - that development often proceeds by strapping on extra developmental phases. When this happens, ontogeny does indeed recapitulate phylogeny.
The brain makes a simple example. Brains are divided into layers, and over evolutionary time, more layers have been added.The human neocortex, for example consists of six layers. During development the neural layers are formed progressively. Migrating neurons climb up a scaffolding made of radial glial cells and bypass previous layers of neurons in the cerebral cortex, creating a new layer on top of their predecessors. This mirrors development over evolutionary time - where ancestors had fewer layers of neurons.
A simple, pure example of cultural recapitulation theory can be found in knot theory. Knots are often formed by tying knows on top of other knots. The simpler knots come first historically as well as during knot construction. A reef knot is a simple example of one knot being tied on top of another knot in order to strengthen it.
The concept of "cultural recapitulation" is often used in another way. Some say that the way that individual learning mirrors historical discovery is a case of cultural recapitulation. For example, the order in which scientific concepts are taught to children might mirror the order in which they are discovered. This is certainly recapitulation and it does involve culture, but the analogy with Ernst Haeckel's idea is weaker than with the knot example I gave. The reason is that the development involved is the development of a young animal - a memetic host rather than a memetic product. The corresponding biological analogy would be if the way in which organisms were infected with pathogens reflect the sequence in which those pathogens evolved. If is easy to imagine reasons why that might be true - for example new pathogens might be less expert at spreading to new hosts. However, this seems a bit different to classical recapitulation theory - since that does not normally involve symbiosis.
Will cultural recapitulation theory suffer the same fate as its organic counterpart? It is, perhaps too early to tell. However, maybe cultural recapitulation theory will help to revitalize recapitulation theory in the organic domain. It is a bit of a shame that recapitulation theory is so widely labelled as a dud idea. Perhaps cultural examples will help to illuminate the truth at the core of the idea.
 In 2003, Geoffrey Hodgson proposed that we use the terms "habit" and "routine" as replacements for the term "meme". As with most other meme synonyms, this suggestion doesn't seem to have been very popular. Retrospectively, it appears to me that this proposal has critical technical limitations that put it out of the running.
Hodgson says that "habits" represent individual transmission while "routines" represent group-level transmission. The dictionary seems to think that individuals can have routines as well, muddying this proposed distinction. Hodgson defends the idea that these entities can act as units of cultural transmission. What he fails to defend is the idea that all cultural transmission is mediated by habits or routines. This claim seems straightforwardly incorrect. For example, the Bible is a bunch of memes, but it isn't a bunch of habits. Habits are associated with individuals, but no individual counts the bible as being among their habits. Nor is the bible a bunch of routines.
This makes Hodgson's proposal incomplete basis of a theory of cultural evolution. If adopting his terminology, we would need one theory for the evolution of habits and routines, and another theory for the evolution of other aspects of culture. Or we would need to redefine these terms and give them counter-intuitive technical meanings. Can we patch up Hodgson's proposal by finding another term (apart from 'habits' and 'routines') to represent other inherited aspects of culture? Maybe - but it looks like a dustbin category to me.
The other issue with Hodgson's proposal is that "habits" and "routines" are not necessarily socially transmitted. We already have terms for mental content that isn't necessarily socially transmitted: 'ideas' and 'concepts'. Part of the reason that term 'meme' found its niche is that it expressed a different idea from the terms 'idea' and 'concept'. If 'meme' had been another synonym for 'idea' and 'concept', it would have failed.
I think Hodgson's proposal is now dead. This post is a post-mortem that attempts to explain where it went wrong.
 One of the dodgy memes perpetuated by some cultural evolution enthusiasts is that there is more to inheritance than genes. Boyd and Richerson say that human evolution progressed "not by genes alone". Lee Alan Dugatkin calls imitation "evolution beyond the gene". Jablonka and Lamb say that "Genetic" is only one of four dimensions of variation, the others being the "Epigenetic", "Behavioral" and "Symbolic" dimensions. Steven Rose says there is "life beyond the gene". David Sloane Wilson says:
core evolutionary theory needs to expand beyond genetics to include other inheritance systems, such as environmentally induced changes in gene expression (epigenetics), mechanisms of social learning found in many species, and the human capacity for symbolic thought that results in an almost unlimited variety of cognitive constructions, each motivating a suite of behaviors subject to selection (Jablonka & Lamb 2006; Penn et al. 2008).
Expand evolutionary theory without expanding genetics? It makes no sense to me: science needs an expanded genetics too. Expanding the domain of evolutionary theory without expanding the domain of genetics would be a very lop-sided approach.
Memetics pioneered the expansion of genetics to cultural evolution way back in the the 1980s. Efforts to establish a new science of 'replicators' to compete with genetics have gone nowhere - and don't make much sense. The latest attempt to establish a science of non-genetic inheritance - epigenetics - is an absolute joke. What a "dustbin" category that is. It is good for one thing - being an example of how not to do science. Waddington's excellent notion of epigenetics is in the process of being hijacked by foolish and ignorant scientific punks. What we need is a generalized gene and a generalized genetics. When generalizing evolution, scientists should not neglect to generalize genes and genetics! These concepts are absolutely needed for any sensible grounding of evolutionary theory on information theory. Genes should be the units of heredity and genetics should be the study of heredity.
Some say that genetic algorithms are not really "genetic". It is nonsense: genetic algorithms really are genetic. Take genes and genetics seriously, dammit. Don't confine them to the special theory of evolution - that's not where they belong.
Molecular biologists may have appropriated the term "gene" - but it isn't theirs to define. As Steven Pinker puts it:
Molecular biologists have appropriated the term "gene" to refer to stretches of DNA that code for a protein. Unfortunately, this sense differs from the one used in population genetics, behavioral genetics, and evolutionary theory, namely any information carrier that is transmissible across generations and has sustained effects on the phenotype.
If we don't expand genetics now, it will only need doing later. Cultural evolution's scientific lag in academia is large, but forward thinking individuals should still be able to see that the need to generalize genetics is now clear and obvious.
IMO, Boyd, Richerson, Dugatkin, Jablonka and Wilson are not doing scientific progress any favours by dragging their feet on this issue. Get with the program, folks. The 'beyond the gene' meme might look progressive to you - but it looks backwards to me. The best way forwards is to generalize genes and genetics when generalizing evolution. This is scientific evolution, rather than scientific revolution - and evolution is usually less painful and more likely.
Don't tell me that genes and genetics have more inertia and are harder to move. That much is now obvious. The point is that they need to move, and will have to move eventually. So, who is helping? ...and who is not? Which side of this issue are you on?
|
|
Posts
