Gene and Meme

For The Encyclopedia of Animal Behavior

Susan Blackmore, May 2018


The concept of memes is derived from the principles of universal Darwinism; that whenever information is copied with variation and selection, that information is a replicator and inevitably evolves. The core definition of a meme is ‘that which is imitated’. Genes are the first replicator; memes the second replicator that emerged when human ancestors became capable of imitating sounds and actions. Criticisms of memetics include claims that they do not exist, that the analogy with genes is false, that the units cannot be specified, that there is no clear equivalent of the germ line in biology, and that the sources of variation are intelligently designed rather than random. Reasons for rejecting these criticisms are discussed. For memetics, memes are selfish replicators that evolve for their own benefit, while other theories of cultural evolution look to biological advantage, providing very different accounts of the origins of the large human brain and language. Imitation is observed in song birds, cetaceans and great apes but animal cultures may arise through forms of social learning other than true imitation and are not memetic. Overimitation in children as compared with other great apes may be important in providing memetic advantage. The possibility of a third replicator, technological memes or tremes, is briefly considered.


The word ‘meme’ was coined in Richard Dawkins’ 1976 book The Selfish Gene. Although, more than forty years on, most people associate memes with the notion of Internet memes, it is helpful to return to Dawkins’ original definition to understand the theory of memetics and its differences from other theories of cultural evolution.

In The Selfish Gene Dawkins (1976) popularised the gene’s eye view of biological evolution. Yet, rather than confining his argument to selfish genes, Dawkins wanted to emphasise what he called ‘universal Darwinism’. This is the general principle that whenever information is copied with variation and selection, then evolution is inevitable. Campbell (1960) describes the same process as “blind variation with selective retention”. Dawkins distinguished between the information that is copied (the replicator), and the carrier of that information (the vehicle). This is similar to Hull’s (1988) distinction between replicators and interactors. In the case of biology, genes are replicators and their phenotypes are their vehicles or interactors. Animals, in Dawkins’ view, are the ‘lumbering robots’ that carry their replicators around and protect them.

The critical point is that these three steps: variation, selection and heredity, can be understood as a simple three-step algorithm, the evolutionary algorithm. This is ‘Darwin’s dangerous idea’, a ‘universal acid’ that rips through all other explanations. “a scheme for creating Design out of Chaos without the aid of Mind.” (Dennett 1995, 50). It was the generality of this scheme that Dawkins wanted to emphasise; ‘Darwinism is too big a theory to be confined to the narrow context of the gene” (1976 p 191). He wanted “to claim almost limitless power for slightly inaccurate self-replicating entities, once they arise anywhere in the universe” (Dawkins 1989 p 322).

Genes may be replicators but they are only one example.  In Dawkins’ opinion, life anywhere in the universe would be driven by some kind of replicator, whether based on chemistry, electronic circuitry or anything else. This is why he asked, ‘do we have to go to distant worlds to find other kinds of replicator and other, consequent, kinds of evolution?’ and answered, ‘no’. Staring us in the face is a second replicator, ‘It is still in its infancy, still drifting clumsily about in its primeval soup … the soup of human culture’ He wanted a name to convey the sense of a unit of imitation, and one that would rhyme with ‘gene’. Taking, mimeme from the Greek meaning ‘imitated thing’ or ‘that which is imitated’, he chose the word ‘meme’. Examples are ‘tunes, ideas, catch-phrases, clothes fashions, ways of making pots or of building arches’ (Dawkins 1976 p 192). We might add to his list, stories, poems, works of art, money, financial institutions, scientific theories, and much more. All these are copied from person to person with variation and selection; they fit the definition of a meme as replicator. In this way memetics echoes much earlier ideas, and indeed versions of universal Darwinism have a long history (Plotkin 1993, Campbell 2011).

In what sense then are replicators, whether genes or memes, selfish? The Selfish Gene popularised the view that biological evolution proceeds not for the benefit of the species or even the individual organism, but for the benefit of the underlying replicator. Dennett (1995), presses the important question ‘Cui Bono?’ or ‘Who benefits?’ revealing the sense in which genes are ‘selfish’. They will get copied whenever and however they can, regardless of the consequences to anything else. This does not mean, as Dawkins pointed out over and over again, that the resulting organisms are selfish. Humans are just one of many species in which cooperation and altruism, as well as competition, have evolved from the underlying competition between genes.

Yet something else occurred during human evolution. Our ancestors, unlike other gene vehicles, became capable of imitating with high enough fidelity to give rise to a new replicator, the meme. Memes are selfish in the same sense as genes are. They use human beings and their technology to get copied whenever they can regardless of the consequences; we are the meme machines that copy, vary and select memes (Blackmore 1999). We were the one species that let loose a second replicator with interests of its own.

The main differences between memetics and other theories of cultural evolution depend on this point. For memetics, memes sustain a new evolutionary process operating on top of the old. They ‘should be regarded as living structures, not just metaphorically but technically.’ (Humphrey in Dawkins 1976 p 192). Christiansen and Chater (2008) disagree. They note that language is akin to an “organism” but they reject memetics and put “organism” in scare quotes, writing, ‘Following Darwin (1900), we argue that it is useful metaphorically to view languages as “organisms,”’ (p 2).

Yet in The Descent of Man (1871) Darwin does not use the word “metaphor”. He discusses parallels, homologies, and analogies, writing that, “The survival and preservation of certain favoured words in the struggle for existence is natural selection. (Darwin 1871 p. 91); the proofs that both different languages and distinct species ‘have been developed through a gradual process, are curiously the same’ (Darwin 1871 p 59). Both involve selection and modification with descent. In this way, Darwin’s view is closer to that taken by memetics. The large human brain and language provide fine examples of how the arguments over this point have been played out and I shall return to these after reviewing some of the classic objections to memetics.

Criticisms of memetics

Do memes even exist?

McGrath (2005) complains that there is “no direct evidence for the existence of ‘memes’ themselves” (p. 121), Aunger (2000) that “their existence has yet to be proven” (p. 7) and Wimsatt that they are “both misnamed and mischaracterized, and perhaps even that they do not exist” (1999 p. 280) or exist only if “conceptualised broadly and inclusively” (Wimsatt 2010 p 273). Yet, I would argue that even asking the existence question reveals misunderstanding of the original concept of memes (Blackmore 2010).

Do dollars exist? asks Dennett (2017). Obviously they seem to but this is only because dollar bills and coins exist; the ‘dollars’ themselves are essentially no different from bitcoins or other digital currencies. Do words exist? he asks. Of course they do, and words are the quintessential  memes. There is no question about the existence of bicycles, phones, furniture, skyscrapers, tattoos, holiday brochures, cricket, or the days of the week.  They are all information encoded in some kind of matter and energy, and all are sufficiently stable to either be copied or not.  Since the core definition of memes is “that which is imitated”, all these examples count. Even if it is hard to pin down, whatever is copied is, by definition, a meme. So “the pertinent question is not whether memes exist, … but whether they are a useful theoretical expedient” (Laland & Odling-Smee, 2000), whether thinking of words, stories, skills, habits and technologies as a new replicator is scientifically or philosophically useful.  I say yes, many others say no. This, unlike the existence question, needs answering.

Memetics has been called ‘pseudoscientific dogma’, ‘a dangerous idea that poses a threat to the serious study of consciousness and cultural evolution’ (Benitez-Bribiesca 2001). Its prospects of unifying the human sciences in the light of Darwin’s theory have been described as ‘rather dim’ (Bradie & Bouzat 2016) and the whole idea dismissed as ‘rife with conceptual problems and utterly lacking in empirical support’ (Poulshock 2002).

Ignoring the existence question and these outright rejections, leaves us with some serious questions to answer. How helpful is the analogy with genes? What should and should not count as a meme? Are memes really replicators? Do they live inside brains, outside brains, both or neither?  And does memetics gives rise to any useful theoretical or practical progress?

Analogies, units and other confusions

The idea that memes are analogous with genes has fuelled many objections and sown much confusion. Midgley calls the idea ‘an empty and misleading metaphor; a ‘useless and essentially superstitious notion’ (Midgley 1994) and rejects it on the grounds that unless we can show “what kind of entity memes are purported to be, the parallel between them and genes surely vanishes, and the claim to scientific status with it.” (Midgley 2003 p 97). Gould points out many significant differences between genes and memes, and suggests that comparisons between biological and cultural evolution ‘have done more harm than good’ (Gould 1991). And McGrath claims that the “case for the existence of the ‘meme’ rests on the questionable assumption of a direct analogy with the gene, which proves incapable of bearing the theoretical weight that is placed upon it” (2005 p 121).

All these miss the point of Dawkins’ reason for inventing the term, which was not to draw an analogy but to illustrate the power of universal Darwinism by treating cultural information as another example of a replicator. Understanding this shows that analogies between genes and memes are secondary to the basic idea, not primary. Some analogies will be fruitful because memes and genes are both replicators, but others will not because memes use entirely different copying mechanisms from genes, totally different sources of variation, and different means of storing information. As Richerson & Boyd point out, “the best evidence suggests that cultural variants are only loosely analogous to genes” (2005 p. 60). This should not be a reason for rejecting memetics.

Another point to remember is that genes and their associated processes have co-evolved for approximately four billion years, creating a high fidelity system with effective error-correction machinery, often separating phenotypes from a germ line, whereas memes have been here for at most a few million years and are still, in Dawkins’ terms, drifting about in their new soup of human culture (1976). We should therefore use gene-meme analogies cautiously, using them to build hypotheses, but not assuming precise similarity.

Another source of confusion concerns units.  Memes are often described as “units of culture” or “units of imitation”, although other descriptions include “elements of culture,” “contagious ideas,” or “cultural instructions”.  Some critics have made the problem of units into a major objection.  For example, Midgley (2000) complains that culture cannot be neatly divided up into units, and nothing is to be gained by “atomising thought,” since “thought is not granular” (p. 67).  Similarly, Jablonka & Lamb (2005) argue that, in cultural evolution, there are no “discrete unchanging units with unchanging boundaries that can be followed from one generation to the next” (p. 212), and Richerson & Boyd (2005) object to the idea that “culture must be divisible into tiny, independent gene-like bits that are faithfully replicated” or “tiny snippets of information” (p. 60).  However, in defining memes as “discrete, faithfully replicating, gene-like bits of information,” (p. 6) they are departing far from the original definition.  Some memes are discrete and some are not; some replicate with very high fidelity (e.g., printed text) and some do not (e.g., dance steps); some memes are like genes in some ways and not in others.  Going back to the original idea of memes as information that is copied, or “that which is imitated,” avoids these unrealistic claims.

The problem arises partly because it is easier to talk about entities than to talk about “that which is imitated,” in the abstract. Dennett tackles the problem head-on, calling memes ‘mindless informational things’ (2017 p 173) and ‘a new kind of evolutionary replicator – culturally transmitted informational entities’ (2017 p 175). He argues for defining the units as “the smallest elements that replicate themselves with reliability and fecundity” (1995 p. 344), or as a cultural item “with enough Design to be worth saving – or stealing or replicating.” (p. 143). In art, for example, an entire gallery is too large a unit for selection to work on; but a blob of pink paint is too small.  The single painting is the natural unit, as when we remember Picasso’s Guernica or buy postcards of Monet’s Water Lilies.  A single word is too short to copyright and an entire library too long, but we can and do copyright three-word advertising jingles and 300,000-word books. A favourite musical meme is the first four notes of Beethoven’s Fifth Symphony, a tiny unit which has spread across the globe, probably even more widely than the larger unit represented by CDs and downloads of the whole work. In all these cases, both large and small amounts of information can count as a meme because they are copied as a unit with sufficient reliability and fecundity for selection to operate. In this view it is not a problem whether memes have to be considered as units or not.

Where are memes?

A further confusion concerns whether memes are stored inside brains, in the physical culture they create, in both or neither. For example, if I tell you an excellent joke and you pass it on to someone else, and they pass it on again, is this meme the precise joke as told, the gist of the joke, the spoken words, or the representations of any of these inside people’s heads? This, often unproductive, debate is complicated by the question whether memes have the equivalent of vehicles, interactors or phenotypes – some kind of physical product of memetic information sometimes referred to as a ‘phemotype’ (Speel 1997). If so, should we say that the meme itself is the joke and its phemotype is the information stored in the person’s brain, or the written version or what?

People have argued each way without reaching any consensus, and just to complicate the issue, Dawkins himself changed his mind. In The Selfish Gene (1976), he made no distinction between memes and their vehicles.  Later, in The Extended Phenotype (1982) he revised this, making “the distinction between the meme itself, as replicator, on the one hand, and its ‘phenotypic effects’ or ‘meme products’ on the other.  A meme should be regarded as a unit of information residing in a brain” (1982 p. 109).  These two views are sometimes known as “Dawkins A” and “Dawkins B” (Gatherer, 1998).

Even before Dawkins coined the term ‘meme’, Cloak (1975) distinguished between i-culture (cultural instructions inside people’s heads) and m-culture (the products of those instructions); the ultimate function of both being to maintain the i-culture.  Subsequently others proposed that memes are neural patterns inside brains, with their products or vehicles being outside the brain (e.g. Delius 1989, Aunger 2002), or “an information pattern, held in an individual’s memory, which is capable of being copied to another individual’s memory” (Heylighen and Chielens 2009). By contrast, Benzon (1996) argues precisely the opposite:  that the replicators are outside brains and their vehicles inside.

This debate has been used by some to discredit or reject memetics (e.g. Jablonka & Lamb 2005, Wimsatt 1999) but others simply avoid it by reverting to the basic idea that memes are whatever is copied; so sounds, written text, physical objects and effects inside brains can all be considered as memes. This might make sense considering that memes are a relatively new replicator that has not settled down into a system with vehicles or interactors distinct from the memes themselves (e.g. Blackmore 1999, Dennett 1995, Speel (1997).

This problem of distinguishing vehicles from replicators has given rise to further contentious and unresolved arguments that memetic inheritance is ‘Lamarckian’ (involving the inheritance of acquired characteristics) and that cultural variation is directed, not random (Aunger, 2000, Gould 1991, Richerson & Boyd, 2005). For some critics, these seem to rule out memetics as a valid enterprise, while others are more concerned with whether culture can evolve if these two claims are true (Kronfeldner, 2007). Yet the concern over Lamarckism is misplaced because the whole idea of calling memetic evolution “Lamarckian” rests on drawing a memotype/phenotype distinction which may be false (Blackmore 1999). For example, Sperber and Claidière claim there is nothing ‘resembling a cultural germline’ (2008 p 285). This is true for any memes that are naked replicators, having no phenotypes, and for these any suggestion of Lamarckism is irrelevant. For others it may not be.

As with previous disputes it may help to step back and remember the difference between one very old replicator (genes) and one very new replicator (memes). Genes are part of an ancient system that has been evolving for about four billion years, probably from simpler precursor systems (Maynard Smith and Szathmary 1995), creating an extraordinarily complex and high-fidelity system that produces all manner of creatures based on the recombination and mutation of information stored in a single type of molecule. The protection of information in the germ line and the separation of the genotype from phenotype was a critical step because it is more effective to copy the instructions for making something than it is to copy the product itself (Blackmore 1999, 2001).  This is not only because multiple copies are made from the same instructions (increasing fecundity), but also because any accidents that befall the product – for example, during its construction or lifetime – are not passed on (increasing fidelity).  This system more effectively retains any “good tricks” that the evolutionary exploration of design space stumbles upon (Dennett 1995).

By contrast, the first memes can have appeared only when our ancestors began to imitate with sufficient fidelity to sustain cumulative culture. Although we have no clear idea of what those early memes were, we can guess that they might include ways of lighting and maintaining fires, carrying and preparing food, or covering and decorating the body. When anyone acquired any such new skill they would have done so by observing someone else doing it first, fitting Thorndike’s (1898) well-known definition of imitation as ‘learning to do an act from seeing it done’. This was Dawkins’ ‘primeval soup’ of culture.

A few million years later, many memes are still in this state with direct copying of the product, low fidelity copying, no distinct generations, and a wide variety of copying methods, but others are not, and we may be seeing clear signs of culture evolving towards a system of separated germ line and phemotype. For example, people may still hear someone sing or play an instrument and try to copy what they hear, but this low fidelity copying is increasingly rare. If we like the music we hear we go and buy the CD or download it from copies that are stored with close to perfect fidelity, elsewhere. When you buy a film on DVD, it was not copied from a previous version of that film (i.e. copy-the-product); instead, multiple copies of the film were produced in a factory from the same original, making the equivalent of a germ line (i.e. copy-the-instructions for making the product, Blackmore 2001). Damage to any particular copy does not matter because it does not affect subsequent copies. The same is true of printed books, cars, planes, and indeed anything made on a production line. Along with these changes, but only in the extremely brief period of the past century or so, has come the switch from analogue to digital storage which further improves fidelity.

As for that other source of arguments against memetics, that the main sources of variation in biology are random unlike the directed variation in so much of culture, we might guess that many aspects of culture will end up finding a way to produce random variation. Indeed this is already the case when genetic algorithms are used in evolutionary computation (Simon 2013) and in much of the software we routinely use. All this suggests that memetic evolution is discovering ever more effective means of copying with variation and selection just as biology has been doing for billions of years.

A New Replicator or Culture on a Leash?

The whole concept of memes depends on the idea that they are a new replicator, with their own replicator power, creating a new evolutionary process operating on top of the old. But is this valid?

One question concerns their method of reproduction. The evolutionary algorithm requires copying with variation and selection, and this implies an important difference between behaviours that are re-produced (i.e. were already in the second individual’s repertoire) and those that are truly imitated (i.e. new behaviours acquired by observing another individual carrying them out) (Thorndike 1898, Jablonka 1999). Sperber argues that a “fundamental objection to the meme model” is that most cultural items are “re-produced” in the sense of being produced many times, but are not “reproduced” in the sense of being copied from another (2000 p. 164). He compares two drawings, a more or less random scribble and a slightly wonky five-pointed star. When people try to copy them, the first is hard and results in a degenerating chain of copies; the second is easy and results in a series that tends towards the familiar star shape. As Dawkins points out, an independent observer might easily be able to put the first set of drawings into the order in which they were made, but not the second; it is “self-normalizing” (Dawkins 1999).

Sperber sees this as a serious objection to memetics because the drawing is not copied or imitated but is “re-produced”, meaning it cannot count as a replicator, whereas Dawkins sees it as an example of the fidelity of meme transmission increasing as people learn to recognise and draw familiar shapes. This same increase in fidelity happens when people learn to speak, dividing the space of possible sounds into a limited number of discrete words that can then be mixed in potentially infinite ways. Reading and writing further increase fidelity, as well as the longevity and fecundity of written memes. Similar, although perhaps less obvious, developments can be seen in chess moves, dance steps, sports, cooking and many other everyday learned skills.

Richerson & Boyd effectively reject memetics by claiming that “cultural variants are not replicators” (2005, p. 82), citing the peculiarities of biased transmission, behavioural attractors, and error prone imitation.  However, this may show only how complex memetic transmission must be (Blackmore 2010).  They also contend that copying must be perfect for a replicator to count as such, suggesting that memes must be “discrete, faithfully replicating, genelike bits of information” (2005 p 6) but this makes no sense. If copying were perfect, whether of genes or memes, there would be no variation on which selection could operate and therefore no evolution.  More interesting is to ask how high the fidelity has to be for a cumulative evolutionary process to get started.  If human imitation is good enough, which it certainly seems to be, we may be justified in treating memes as replicators.

This question is important because it affects the perceived relationship between genes and memes.  Theories of gene-culture co-evolution tend to follow Lumsden and Wilson’s famous claim that “genetic natural selection operates in such a way as to keep culture on a leash” (1981 p 13). In other words, culture is an adaptation of benefit to human genes and, ultimately, kept under control by them.  For memetics, memes did not evolve as an adaptation of benefit to genes; they were a new replicator that was accidentally let loose by the human capacity for imitation and may be either adaptive or maladaptive from the genes’ point of view.  In 1976, Dawkins complained of his colleagues that “In the last analysis they wish always to go back to ‘biological advantage’” (p. 193).  Richerson and Boyd provide a perfect example. Their ‘costly information hypothesis’ comes very close to memetics in some ways; they occasionally use the term “selfish meme” and describe us, and our culture, as like obligate mutualists. Yet they still maintain that “culture is an adaptation” and that “culture is on a leash, all right” even if the dog on the end is clever and strong.

This is especially clear when it comes to the highly contentious issue of the origins of language. Most theories assume that language must have evolved as an adaptation, i.e. using language was adaptive for early humans’ genes. Although this may seem obvious, memetics does not assume this but instead sees language, like the rest of culture, as a parasitic second-level replicator that appeared when our ancestors became capable of imitation. These new, imitated behaviours required energy, time and brain power both to acquire and retain. In this view, our relatively enormous brains were a burden created by memetic drive, by the demand to become ever better at imitation when the meme pool expanded. And human language was not a biological adaptation but a parasite turned symbiont (Blackmore 1999, 2007).

Both memetics and gene-culture co-evolution theories claim that brains and language have shaped each other (Blackmore 2008, Christiansen and Chater 2008) but again with the same fundamental difference. Most theories appeal to genetic advantage to explain this process and either reject or ignore memetics altogether (e.g. Botha and Everaert 2013, Christiansen et al 2009, Chater and Christiansen 2010). As though this nullifies the value of any memetic analysis, some specifically reject it, pointing to sources of variation and claiming that memes ‘seem patently to be products of sighted watchmakers; that is, they are products, in part at least, of many generations of intelligent designers, imitators, and critics.’ (Christiansen et al 2016 p 61. They ignore the fact that some memes are created intelligently and deliberately while many are not. Much imitation occurs quite unconsciously as when people in conversation mimic each other’s gestures or body position, pick up each other’s accents and mannerisms, or find themselves helplessly humming a tune they just overheard.

For those who accept memetics, there is a fascinating argument to be had over the extent to which memes are mindlessly copied and varied as opposed to intelligently designed and whether this has changed over the course of memetic evolution.

As with respect to the germline/phenotype distinction discussed above, I have argued that memetic evolution has, in some ways, become more Darwinian in the sense that early memes were naked replicators being copied with no such distinction, while modern memes such as printed books or cars produced in a factory do. By contrast, Dennett (2017) uses Godfrey-Smith’s (2009) concept of Darwinian spaces to argue that human culture is gradually becoming less Darwinian or ‘de-Darwinizing’, with early memes being more or less mindlessly copied while modern memes are ever more deliberately and intelligently designed. There is much to be explored here and possibly both changes are taking place at once.

Do other animals have memes?

If a meme is that which is imitated, as Dawkins originally defined it, this question is equivalent to asking whether other species can imitate and, if so, what they can imitate and whether this gives rise to copying with variation and selection. However, the situation is not that simple because not everyone agrees with defining memes this way and even Dawkins referred to ‘imitation in the broad sense’ which might allow other types of social learning to count as creating memes, as some other theorists have suggested (e.g. Durham 1991, Runciman 1998, Laland and Odling Smee 2000, Brodie 1996, Ball 1984, Gabora 1997). Reader and Laland argue that while human imitation, teaching and language may explain why human culture and language are so diverse compared with animal proto-cultures, many non-human animals are excellent social learners. As long as information is transmitted with sufficient fidelity to be replicated in another’s brain, this should count as memetic transmission; ‘the meme concept can, and should, be applied to animal cultural transmission’ (Reader and Laland 1999 p100).

A related question is whether to apply the word “culture” to behaviours that are spread by forms of social learning other than true imitation. If these are included as cultural, then some primates, rats and birds have culture but, by a strict definition of memes, they do not sustain memetic culture. This may be the same as asking whether all culture has to be cumulative culture or involve a ratchet effect by which culture builds up (Tomasello 1999). These questions are important because memetics rests upon the claim that memes are a new replicator. Any process that gives rise to memes must therefore be capable not only of reproducing old behaviours by observing others performing them but creating new behaviours that can be passed on with variation and then become subject to selection. The question then is whether other forms of social or individual learning can do this.

Contagion is a clear example of non-memetic transmission. For example, coughing, yawning and laughing can all be contagious, spreading quickly through groups of people. But laughing when someone else laughs, or joining in a spate of annoying coughing at a concert, involves no copying.  The second person’s laugh is triggered by hearing the first, and is their own laugh, not a copy of the first person’s laugh (that is, unless they are deliberately mimicking).  Contagion in other species includes picking up a warning cry from another, or fleeing when another does.  This is entirely triggered re-production, and is neither cultural nor memetic.

Two famous examples illustrate the way animal culture can arise through social learning processes other than imitation; tits opening milk bottles in England and macaques washing sweet potatoes in Japan.

Throughout much of the twentieth century in England, unhomogenised milk was delivered to people’s doors in glass bottles with shiny silver or gold-coloured foil tops, and in 1921 a new phenomenon was observed; the foil tops were found broken and jagged. The culprits turned out to be various small birds, mostly blue tits (Cyanistes caeruleus), and the new behaviour spread from village to village, having started at several separate locations (Fisher and Hinde 1949). This can reasonably be thought of as bird culture but was this spreading innovation a meme; would it count as a replicator? In other words, did it spread by a process capable of sustaining copying of behaviours with variation and selection? The answer may be no; although this looks like imitation, the new behaviour might have spread by stimulus enhancement rather than imitation (Sherry & Galef 1984). Blue tits often tear at bark or leaves to find food, and peck to eat it, and experimental studies have shown that blue tits, especially juvenile females, are proficient at social learning and this ability is correlated with innovative problem solving (Aplin et al 2013). It is therefore not surprising that some innovative birds may have tried the foil tops and discovered the rich cream underneath.  They would then have left jagged silver tops which would be conspicuous to other birds who would then also discover the cream and associate this treat with milk bottles (Sherry & Galef 1984).  On this account, the birds had not learned a new skill from seeing it done, but only a new stimulus at which to tear and peck. There was no indication of variations in the behaviour that could be selected, and no evidence of cumulative culture suggesting a true replicator.

Similar doubt can be cast over the case of the Japanese macaques on Koshima Island who, in 1948, were found to be taking the sweet potatoes that were provided for them, and washing the sand off them in the sea (Hirata et al 2008). This new cultural behaviour was long thought to have been spread by imitation but there are other possibilities. Macaques seem to enjoy water and there are many other instances of groups that regularly go into rivers or pools. Juveniles also tend to follow their mothers. So it is possible that some individuals accidentally discovered that sweet potatoes dropped into the sea were nicer to eat. Then, by following them, others discovered the trick for themselves. This would enable an animal culture but not one that could evolve by generating variations available for selection. In other words, sweet potato washing is not a meme.

There are, however, many instances of what appears to be true imitation in other species and a long history of speculation on the topic (Galef 2013, Nehaniv et al 2002). Song birds imitate sounds, and there have been several studies of bird song memes and their survival and propagation in the meme pool (Bonner 1980, Baker 1996, Lynch et al 1989, Payne et al 1988). The brain changes entailed in such imitative learning have been studied in zebra finches that listen to their father’s song from about a month after hatching and practice each song hundreds of thousands of times (Vallentin et al 2016).

Many cetaceans and some apes are known to be capable of imitation and have been studied both in the wild and the laboratory (Whiten (2001). Recent experimental studies have used methods such as the ‘do-as-other-does’ paradigm in which animals learn to copy familiar and novel behaviours by watching a demonstrator. Killer whales, for example, have been shown to acquire novel behaviours this way (Abramson et al 2013). Dolphins have been taught to imitate on demand, can imitate both other dolphins and humans, and are capable of deferred imitation, with juveniles more often displaying spontaneous imitation, and adults being better at elicited imitation (Kuczaj & Yeater 2006). But whether these abilities underlie dolphin culture, and whether the acquired behaviours should be considered as memes, is less certain.

Such studies show that some species are capable of true imitation. Do the copied behaviours then necessarily count as memes? And can they sustain evolving cultures? Fierce arguments have concerned the comparison between human imitation and that of the other great apes, with disputes over the question of whether ‘apes ape’ (Tomasello 1993). Some have argued that while children are true imitators, chimpanzees display only emulation. In other words they copy only the goal of an observed behaviour, such as gaining food, and use their own methods for completing the task (see Whiten et al 2009 for a review). Experimental studies comparing children with chimpanzees have provided intriguing results suggesting ‘overimitation’ in children but not in other apes. That is, children readily imitate specific actions even when these do not obviously lead to any reward, are irrelevant to the demonstrated outcome and even when there are more efficient methods of achieving the demonstrated outcome or their own actions result in failure  (Nielsen, 2009).

For example, in some experiments demonstrators used a mixture of relevant and irrelevant actions to open an ‘artificial fruit’ with food or a toy inside; when the box containing a reward was opaque both children and chimpanzees copied the demonstrated actions fairly accurately but when it was transparent and they could see that some actions were irrelevant to opening the box, the children continued to copy all the actions while the chimpanzees used only relevant actions. In another study, two-year-old children, chimpanzees and orangutans were shown how to open a plastic tube to get at food or a toy inside. Whereas the children mostly copied the demonstrated actions the other apes tended to try to smash or tear their way into the tube (Horner & Whiten, 2005, Nielsen 2009, Whiten 2017). Similar differences are observed comparing 3 to 5 year-old children with bonobos, in which the children readily copied causally irrelevant actions whereas none of 46 bonobos did so (Clay & Tennie 2017).

This difference could be highly significant from a meme’s eye point of view. Overimitation means copying almost any demonstrated memes; emulation (or goal emulation) means that only memes perceived by the animal as useful or relevant to its goals are copied. For the evolution of culture this makes an enormous difference. If memes are replicators, the tendency to copy all or any actions is a far more effective basis for their evolution, providing more actions, more variation in actions, and therefore greater scope for selection to operate. From the perspective of meme theory, overimitation – or non-selective imitation – is precisely what would be needed to let loose a second replicator which would then evolve in its own direction for its own benefit. This would force meme-gene co-evolution driven by the competition between the genetic advantage of selectively copying only useful memes and the memetic advantage of copying all.  In this view, human culture really is different from the cultures of other species.

The future of memetics

For decades after it was proposed in 1976, the concept of memes was widely ignored or derided and very little theoretical or experimental progress was made. Many reasons were suggested, ranging from the difficulty of making testable predictions to people’s fears that memetics takes away much of human self-determination and power. Nevertheless, empirical work that either tests or uses memetics is increasingly being reported, for example on the evolution of music and musical notation (Jan 2017), comparison of chemical terms in Chinese and Western science (Wright 2000), art and design (Langrish 2004, Steadman 2008), team problem solving (Rosen et al 2017), talent management (Swailes 2016), economics and cultural values (Waddock 2016), the evolution and globalisation of jokes (Shifman & Thelwall 2009, Shifman et al 2014) and, of course, the survival of replicating text in the Internet (Pocklington & Best 1997) and the spread, popularity and effects of Internet memes (Diaz & Mauricio 2013, Miltner 2014, Gal et al 2016).

I have suggested a further possibility that builds on the principles of universal Darwinism, that perhaps even more new replicators might piggy-back on existing ones. Memes arose when one of the products or phenotypes of genes became capable of a new kind of copying – imitating sounds and actions. I have suggested that a third replicator might, in an analogous process, emerge from products of the second. Indeed, such new replicators may already be emerging as digital information copied by computers, phones, servers and other devices (Blackmore 2010). I initially referred to these new replicators as ‘temes’ but when this caused much confusion I changed the name to ‘treme’. Tremes are digital information copied, varied and selected by silicon based technology, independent of human control. If this is correct then tremes would provide another example of a selfish replicator, blindly propagating for its own benefit rather than for the benefit of human beings, their genes, their technology, the other animals with which we share our world, or the environment in which all these live.




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