Imitation has been put forward as a defining feature of memetic transmission. Since there is currently poor evidence for imitation in non-human animals, such definitions have been interpreted as restricting meme theory to the study of human behaviour patterns and birdsong. We believe this is a mistake. Human capacities for imitation, teaching and language may well account for the extraordinary diversity of human culture compared with animal proto-cultures, but imitation is simply one mechanism of transmitting acquired information between individuals. As long as information is transmitted with sufficient fidelity to be replicated in the brain of the receiver, any social learning process will do. Non-human animals may be poor imitators, but many are excellent social learners. We argue that the meme concept can, and should, be applied to animal cultural transmission.
Keywords: social learning, cultural transmission, acquired information, animals, imitation, replicator, meme.
In this article we present a counter position to Blackmore's, arguing that animals should be allowed the right to carry memes. We present our arguments, not as a criticism of Blackmore's position, but rather in an attempt to open up an informed debate over this interesting issue. We agree that evidence for non-human animals routinely imitating is weak, but argue that imitation need not, and should not, be the defining feature of a meme. Transmission fidelity, not the psychological process underlying transmission, is a determining feature of whether a meme can spread and replicate.
There is little doubt that animals regularly acquire learned information from one another. The term `social learning' refers to learning that is influenced by observation of, or interaction with, another animal or its products (Box 1984). Experimental investigations have revealed that imitation is just one of several processes that can result in social learning (Galef 1988). There exist numerous reports of novel behaviour patterns spreading through animal populations through social learning processes (for reviews see Lefebvre & Palameta 1988 or Galef 1988). Famous cases include termiting in chimpanzees (Goodall 1964), food washing in Japanese macaques (Kawai 1965), the opening of milk bottles in British birds (Fisher & Hinde 1949), dietary preferences in rats (Galef 1996), birdsong (Jenkins 1978), and fear of snakes in rhesus monkeys (Mineka & Cook 1988). In the majority of cases where the transmission process is investigated, behaviour patterns are not transmitted by imitation, but instead result from other, simpler processes such as local enhancement, where an animal's attention is drawn to an object by the actions of another, in a manner that results in learning (Galef 1988). Yet animals clearly have behavioural traditions based on acquired information transmission. Moreover, many of these animal traditions appear to change over time (e.g. Gibbs 1990) in a manner perhaps consistent with the predictions of memetic evolution.
So why do researchers place such a premium on imitation? It would seem that, for Blackmore, the key to memetic transmission is that something is learned about the form of a behaviour rather than about the environment (1999, p.49). For example, Blackmore argues that milk bottle opening in British birds is not a meme since "the tits already knew how to peck; they only learned what to peck" (Blackmore 1999, p.49, reversed original italics).
However, while in the past definitions of imitation have stressed the learning of a new behaviour through observation (e.g. Galef 1988), it is now widely recognised that imitation is not solely, and may rarely be, concerned with learning a new motor pattern: imitation is also concerned with learning the context and consequences of an established motor pattern (Heyes 1993; Heyes 1996, p.381). For illustration, consider the case of someone learning to play tennis through imitation. Here the apprentice is not learning to run, hold racket like objects, hit things or wave their arms around - all of the required motor patterns are already part of their repertoire. They are learning to move their body in a particular way (for instance, to move the racket towards the ball), at a particular time (when the ball is approaching), to generate a certain result (to hit the ball in the desired manner). The tennis playing meme is not exclusively concerned with motor patterns: it is a complex of information that also concerns the location appropriate for the behaviour, the objects with which one interacts, and the consequences of the behaviour. To take another example, this time the `making pumpkin soup' meme described by Blackmore (1999, p.61), we have a case where the recipient of the new recipe is not preparing food, cooking, or even making soup for the first time. The cook is simply carrying out an existing motor pattern (that of making soup) with an ingredient novel to the soup-making context but familiar in other contexts.
Exactly the same logic applies to the milk-bottle-top opening birds. They are not learning to peck any more than the tennis apprentice is learning to run around or hold rackets: that motor pattern is already part of their repertoire. They are learning to peck a particular object (the milk-bottle), found in a particular location (on a doorstep), to generate a particular consequence (the cream reward). Similarly, Japanese macaques are not learning to move their hands in water, they are learning that if they move particular objects (the sand-covered sweet potatoes) in water they can generate a desirable reward (that is, sand-free food). Imitation is not a criterion upon which the meme-carrying of animals and humans can be distinguished.
This raises a policy decision for memetics. Should reconstructed memes count as genuine memes? We argue that they should for two reasons. First, virtually all memes, including those of humans, involve an element of reconstruction (Sperber 1996). Humans when they imitate rarely do so perfectly, and they are typically forced to re-evaluate and adjust their behaviour in the light of sensory feedback (Piaget 1962; Yando et al. 1978; Custance et al. 1995). To argue that largely reconstructed memes are not memes would require an arbitrary and unenforceable rule to be employed regarding just how much reconstruction is allowed before acquired information qualifies as a meme. Reconstruction cannot be quantified in any meaningful sense. Second, reconstructed memes fit the Darwinian model as effectively as perfectly transmitted memes. They too can replicate and evolve, and to eliminate them on arbitrary grounds at this early stage in the science of memetics risks eliminating a large number, maybe even the majority, of interesting cases of social transmission that may benefit from memetic analysis. What counts is not how transmission occurs, but whether the product is similar in the transmitter and receiver - in other words, what counts (amongst other things) is fidelity (Dawkins 1989).
So why is an exception made of birdsong? We suggest two reasons, both unconnected to the psychological processes that underlie learning. First, Dawkins mentions birdsong in The Selfish Gene (1989, p.189). Dawkins's authority carries a great deal of weight in memetics, and if he says birds have memes that is enough for many. Second, birdsong has been subject to some of the most elegant empirical work on memes (e.g. Burnell 1998; Lynch 1996; Lynch et al. 1989). It turns out that birdsong lends itself to the meme concept beautifully, perhaps better than almost anything else, even human culture. We would welcome similar memetic analyses of other forms of animal social learning.
The meme concept need not be restricted to animals phylogenetically close to humans. In an experimental study, Curio et al. (1978) demonstrated that observer birds could learn to mob a non-raptorial bird, and even a plastic bottle, as a consequence of witnessing another bird mob at its presentation. This behaviour was transmitted along chain of six individuals, with each individual acting as a demonstrator for the next bird in the chain. Here a meme for the context in which mobbing is an appropriate response is transmitted. However, there were limitations to what could be transmitted. For example, birds did not learn to mob an empty cage, and learned to mob the plastic bottle stimulus less strongly than the bird stimulus. There may be similar restrictions on the successful transmission of human memes.
Third, an example from fish. Mating sites of the bluehead wrasse (a coral reef fish) have remained in daily use over 12 years (four generations) without changing location (Warner 1988). There was no obvious correlation of sites with resources, and many more potential mating sites available. Entire populations were experimentally replaced, and the transplanted populations chose new sites; but where females were transplanted to locations containing native females, the newcomers only mated at established sites, implying that tradition rather than resources is important in determining mating site location. After further experiments Warner proposed that these arbitrary traditions were probably transmitted by females following one another (Warner 1990), a very simple social learning process. Here we have an example of a long-lived, arbitrary tradition transmitted without imitation. Again, we can consider the mating site preference that each fish learns as a discrete, replicating unit of information, and examine the diffusion dynamics of these memes.
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