Selected articles on how a memetic understanding of altruism can lead to cultural transformation


 By: John R. Evers

I. Introduction: Genes and Altruism

Charles Darwin described the sterility of certain castes of social insects, and more generally, the reproductive self-sacrifice such organisms represented, as “one special difficulty, which at first appeared to me insuperable, and actually fatal to my whole theory.”[1] In the 1960’s, W.D. Hamilton “inaugurated” the theory of “kin selection,” which offered a brilliantly simple explanation for such altruistic behavior.[2] As Holldobler and Wilson explain, “Hamilton recognized the importance of a measure he called inclusive fitness, which incorporates both the individual’s personal reproduction (classical fitness) and its influence on the reproduction of collateral relatives.”[3]

The essentials of kin selection and inclusive fitness are summarized according to a simple equation, called “Hamilton’s Rule,” which is expressed: C/B < b. “This says that the cost C (which is the loss in expected personal reproductive success through the self-sacrificing behavior) divided by the benefit B (the increase in the relatives’ expected reproductive success) must be less than b, the probability that the relatives have the same allele,”[4]if the altruist gene is to survive natural selection.

Inclusive fitness begins by focusing solely at the level of the gene, but then widens that focus to encompass the entire group or population comprising any given gene’s extant copy-set. For example, assume organism X is a carrier of gene-A and that gene-A causes X to behave in a certain way which kills X prior to the production of any offspring. If X’s suicidal behavior directly enhances the survival and reproductive potential of X’s siblings, gene-A still has a chance at survival and even proliferation – not in X or X’s non-existent offspring, but in X’s siblings and their offspring since at least some of X’s siblings will carry copies of gene-A. One copy of gene-A is just as good as the next; any single gene-copy is just as transitory as the organism which carries it.[5] Indeed, inclusive fitness demonstrates that the true fundamental unit of natural selection is a gene’s entire copy-set.

For this reason, in the expression C/B < b, everything hinges on b – the probability that the altruist’s siblings carry copies of the same altruist gene. As Dawkins observed, “[k]in selection accounts for within-family altruism; the closer the relationship, the stronger the selection.”[6] Thus, the “family” is merely a short-hand approximation of a representative portion of any given gene’s copy-set (according to the degree of relation within the family), and changing degrees of relatedness will directly effect levels of intra-family altruism.[7]

The obvious limitation of Hamilton’s Rule is that it only justifies altruism expressed within the family. Alternative theories, such as reciprocal altruism, can account for the natural selection of generally cooperative behavior regardless of familial relationships, but theories such as reciprocal altruism do not necessarily account for altruism.[8] As Wright points out, “[reciprocal altruism] doesn’t involve sacrifice for anyone who doesn’t ultimately reciprocate.”[9] Hence, reciprocal altruism does not account for “sacrifice” at all. For the purposes of this analysis, an organism that is to be considered an “altruist” must suffer a net cost (in terms of reproductive potential) as a result of its behavior, as it was primarily this feature that established the monumental problem which challenged Darwin in 1859 and which was solved in part by Hamilton’s Rule.

The intra-family limitation of Hamilton’s Rule is a result of a certain feature of genetic reproduction; namely, that genes can only produce copies by making new carrier organisms in which to house these copies. In other words, gene copies are only distributed vertically (down family trees) within a population. Because genes distribute their copies in this fashion, the only easily identifiable groups of organisms that will have some a priorireason for possessing high b values in Hamilton’s Rule is the genetic family. Thus, while Hamilton’s Rule may have solved the paradox of completely self-sacrificial behavior within the confines of the genetic family, it simultaneously precludes the possibility of altruism directed beyond the genetic family since it is unlikely that a random, non-family member will share a copy of the relevant altruistic gene. When C/B is less than b, self-sacrificing behavior can be seen as successful altruism; but, when C/B is greater than b, the same behavior begins to look more like genetic suicide. In fact, altruism of any degree should face the threat of competitive elimination where the b-value is zero.

Moreover, any benefit enjoyed by an organism not carrying the altruist gene (a “non-carrier”) will necessarily dilute the net benefit that would otherwise accrue to the altruist gene’s copy-set. Therefore, if an altruist gene is to succeed, some mechanism must evolve which can eliminate (or, at least, offset) this potential dilution. By functioning to decrease any potential benefit to competitors and increase potential benefit to the altruist gene’s copy-set, this mechanism will manifest itself generally as an increase in competition between carriers of the altruist gene and their non-carrier competitors. Thus, the inverse operation of Hamilton’s Rule dictates levels of inter-family competition generally proportional to any degree of intra-family altruism.

However, if some non-genetic unit of natural selection can allow for a sufficiently high b-value among non-siblings (i.e., between any randomly selected members of a population), then the threat posed by the inverse operation of Hamilton’s Rule is removed. Richard Dawkins introduced the generally accepted name for this alternative unit of natural selection: it is the “meme.”[10]

II. Memes and Altruism

Any meme can be defined generally as a rule of behavior, encoded by functional neuronal groups or pathways. Behavior is action, whether mental or physical. Ideas such as tying shoe-laces or opening a door represent rules of physical action, i.e., rules of patterned neural-muscular interaction. Concepts such as apple, seven, or causality, represent rules of mental action, or rules of cognition, i.e., rules of patterned neural-neural interaction. Hence, physical movement is governed by memes which represent rules of physical action and thought is governed by memes which represent rules of mental action.[11]

If memes are units of natural selection (allowing for a memetic application of Hamilton’s Rule); and, if memes copy horizontally (allowing the “memetic family” to extend beyond the scope of the genetic family); and, if memes can be directly responsible for altruistic behavior, then memetically driven and inclusively fit altruism can extend to the whole of any given (freely communicating) human population according to the mathematical purity of Hamilton’s Rule.

A. Memes as units of selection

Most, if note all, models of selection share three basic elements: (1) variability of base units; (2) competition between base units for reproductive resources, or selective pressure; and, (3) selective reproduction (or, selective longevity).[12] Memetic variation is demonstrated by the approximately one trillion (1,000,000,000,000) neurons which make up the average human brain, with the cerebral cortex accounting for at least 50 billion of these nerve cells.[13] Moreover, because anyone who wants to share any idea with another person must create a physical substrate for the idea (e.g., language, whether spoken or written), the full scope of memetic variability is most accurately described by combining the subjective knowledge of each living human carrier with the full body of extant objective knowledge (e.g., knowledge encoded in books, or computers).

Considering a meme as a discrete neural group or pathway, it follows that memetic competition is fueled by the need for (finite) neurochemical resources and (limited) neural space.[14] That neural patterns or neural groups compete for space and neurochemical resources is not a novel proposition.[15] Thus, memetic competition for the limited space and biochemical resources available in the brain can be restated simply as competition between neurons or functional groups of neurons for stimulation.

Rules of behavior which are followed and reinforced to increase the probability of future stimulation are remembered. Memes which fail to acquire the necessary resources for reproduction, whether manifested in physiological or non-organic material substrates, will face competitive elimination, i.e., they will be forgotten. Every human activity, from basic cognition to daily “decision-making,” can be seen as a process of natural selection during which functional neuronal groups compete to occupy a behavioral (whether mental or physical) niche within a carrier organism’s nervous system. Hence, certain memes are selected over others based on the neurochemical characteristics of the underlying neuronal group(s) or pathway(s), as well as their resulting classical and inclusive fitness.

B. Horizontal Memetic Reproduction and Altruistic Memes

The second and third demonstrations, that memes are capable of horizontal reproduction and that memes are capable of encoding altruistic behavior (assuming such behavior exists at all), are easily made as they are both self-evident propositions.

Genetic reproduction occurs through an elaborate process of cellular fusion and the consequential growth and maturation of newly formed carrier organisms. Memetic reproduction occurs through an equally elaborate processes of imitation and/or linguistic communication, whereby memes are copied from one nervous system into another. That memes are capable of horizontal reproduction (i.e., that memes are not limited to copy distribution within genetic families) is a self-evident proposition, provided the recipient of the proposition is not a genetic sibling of the one advancing the proposition. Indeed, any attempt to dispute the proposition that human beings generally are capable of sharing ideas is to advance or attempt to share the idea that ideas cannot be shared. Therefore, it is literally beyond argument that memes are capable of horizontal reproduction.

Finally, while genes encode the assembly of polypeptides and thereby transcribe physiological structure, memes represent patterns of neural stimulation and thereby transcribe behavior. Therefore, if anything is to be directly responsible for altruistic behavior it will be a meme, whether the meme is genetically encoded (i.e., innately acquired), or learned (i.e., environmentally acquired).[16]

III. Memetic Application of Hamilton’s Rule

As currently applied, Hamilton’s Rule is based solely on the static nature of genetic identity: because an organism’s genetic makeup is fixed for the duration of the organism’s lifespan, any two organisms will either share a copy of a gene or they will not, and this objective fact is measured according to probability and familial relation. However, since memes are capable of horizontal reproduction (whether through imitation or linguistic communication), it is possible that any meme, including an altruist meme, could express itself in favor of (and, therefore, presumably in the presence of) a non-carrier competitor and thereby copy itself into the memetic (i.e., neuronal) structure of the competitor. In short, a competitor directly experiencing the benefits of another’s self-sacrifice might subsequently imitate such altruistic behavior for no other reason than having experienced such behavior first-hand. Consequently, one should expect various factors (whether genetically encoded, memetically encoded, or both) to evolve throughout a population which would enhance or detract from the probability of such imitation (e.g., empathy, admiration, so-called “open-” or “close-mindedness,” etc.).[17] Any given meme or set of memes could likewise evolve various mechanisms to increase the probability of such imitation (e.g., cognitive or psychological appeal, etc.).[18] Hence, it is conceivable that some factor or set of factors could be delineated that would determine the relative rate of conversion (from non-carrier-competitor to carrier-clone) for any given meme within a certain population.

This rate of conversion shall be expressed as a new variable, to be added to the right side of Hamilton’s Rule since it will operate to increase the likelihood that the beneficiary organism(s) will carry a copy of the same altruistic meme (following the altruistic encounter). Because the rate of conversion will have no application where a beneficiary already carries a copy of the altruist meme, it will only modify the probability that a beneficiary does not carry a copy of the altruist meme, which is described by (1-b). Therefore, the memetically adapted Rule is: C/B = b + c(1-b), where the rate of conversion [c] measures the likelihood that expression of a meme will “infect” a non-carrier and thereby convert the non-carrier into a carrier.

Operation of this adapted version of Hamilton’s Rule can be stated as follows. Assume that there exists a 1/4 (or 25%) degree of memetic relation within an intra-communicating population. Hence, each member of this population shares roughly 1/4 (or 25%) of the same memes. Assume also that a member of this population carries an altruist meme that has a conversion rate of 3/4 (or 75%). Hence, for every four non-carriers, three will be converted into carriers upon contact with the meme. According to these figures, there is a 13/16 (or, 81.25%) chance that expression of this meme in favor of any randomly selected member of the population will operate to confer a direct benefit on a representative member of the altruist meme’s copy-set. Whether a net benefit is likely to be conferred by expression of that meme will depend on comparing this figure with the Cost-Benefit ratio defined by the left side of Hamilton’s Rule.

IV. Conclusion

According to the memetic application of Hamilton’s Rule, the key factor to success for altruistic behavior is not a high probability of clone status (with regard to the altruistic gene) based on genetic familial relation, but a high probability of clone status (with regard to the altruistic meme), based on a population’s horizontal reproductive fertility. Indeed, a highly infectious, altruistic meme (e.g., 80% conversion rate) existing in a small fraction (e.g., 10%) of the population would still enjoy a very high probability (72%) that expression of any copy, in favor of any random member of the population, would serve to benefit that meme’s (growing) copy-set (all other things being equal). Assuming acceptable criteria and empirical data can be established to give real meaning to the “rate of conversion” for any given meme, the memetic application of Hamilton’s Rule offers a comprehensive justification for general (intra-societal, or intra-cultural) altruism. Meanwhile, it can be said with newfound certainty that purely altruistic behavior is possible within any memetically fertile population.


[1]Holldobler and Wilson, The Ants, (Harvard University Press, 1990), at p.181.




[5]A sequence of nucleotides which dictates the construction of a specific polypeptide can be classified as a “gene,” and can be analyzed as being fit or unfit according to natural selection; but, it cannot be called a “survivor.” Likewise, a discrete quantum of functionally synchronized neurons can be classified as a “meme,” and can be analyzed as being fit or unfit according to natural selection; but, it cannot be called a “survivor.” Any particular sequence of nucleotides or quantum of neurons will degrade prior to or contemporaneous with the death of the carrier organism. Indeed, nucleotides and neurons are as much a part of the organism as is any protein, enzyme, cell, organ, or physiological system. The only thing that is carried by an organism (in the sense that it can outlast the organism) is the underlying pattern of nucleotides or neurons, which exists purely as a speculative construct of subjective consciousness. The concept of the potentially immortal gene/meme can be made to represent a concrete reality only by redefining the functional “gene” or “meme” as the total extant copy-set of any given particular genetic/memetic manifestation, i.e., as a population of gene/meme copies. Any individual member of such a population may be transitory, but the population itself can survive indefinitely.

[6]Richard Dawkins, The Selfish Gene (Oxford University Press, 1989), at p. 94.

[7]See e.g., Robert Wright, The Moral Animal (Vintage Books, 1994), at 164.

[8] See Dawkins, The Selfish Gene, supra note 6, pp. 202-233 for explanations of various strategies for cooperation and mutual benefit. See also, Wright, The Moral Animal, supra note 7, at pp. 189-209. According to Wright, reciprocal altruism is the only available explanation for inter-family altruism and, therefore, “wins by default.” Id. at 202.

[9] The Moral Animal, supra note 7, at p.207.

[10]The Selfish Gene, supra note 6, at p. 192.

[11]A possible third category of action is linguistic action, which represents a means of converting physical action into mental action, i.e., language is movement that encodes thought.

[12]See e.g., Edelman, Neural Darwinism (Basic Books, 1987), at p. 9.

[13] Kandel & Schwartz, eds., Principles of Neural Science (N.Y.: Elsevier Science Publishing Co., 2nd Ed. 1985), at 223.

[14]It also follows that memes compete for access to the limited and finite materials required for manifestation in non-organic memetic carriers, e.g., book space, computer storage space, etc.

[15]For example, Edelman describes the “Mnemon hypothesis,” proposed by J.Z. Young in 1965, which describes a means for selective competition amongst neuronal groups competing for stimulation. (Neural Darwinism, supra note ___, at 14-15). Edelman’s own theory of neuronal group selection is based on the concept of neuronal competition for stimulation. (Id. at pp. 45-46).

[16]For the purposes of this analysis, innate behavior describes behavior which is the immediate result of memetic expression, but which is ultimately the result of genetic expression, i.e., the responsible memes are formed as a result of genetic expression. Learned behavior is the product of environmentally acquired memes which are, in turn, the product of epigenetic, somatic selection. An innately acquired meme that is incapable of horizontal reproduction is the functional equivalent of a gene.

[17]See generally Aaron Lynch, Thought Contagion (Basic Books, 1996), at pp.2-16.


What makes a meme successful? Selection criteria for cultural evolution

CLEA, Free University of Brussels, Pleinlaan 2, B-1050 Brussels, Belgium

To be published in: Proc. 15th Int. Congress on Cybernetics (Association Internat. de Cybernétique, Namur).

ABSTRACT. Meme replication is described as a 4-stage process, consisting of assimilation, retention, expression and transmission. The effect of different objective, subjective, intersubjective and meme-centered selection criteria on these different stages is discussed.


Cultural evolution, including the evolution of knowledge, can be modelled through the same basic principles of variation and selection that underlie biological evolution (Boyd & Richerson, 1985; Cavalli-Sforza & Feldman, 1981). This implies a shift from genes as (replicating) units of biological information to a new type of (replicating) units of cultural information: memes (Dawkins, 1976). A meme can be defined as an information pattern, held in an individual’s memory, which is capable of being copied to another individual’s memory. This includes anything that can be learned or remembered: ideas, knowledge, habits, beliefs, skills, images, etc. Memetics can then be defined as the theoretical and empirical science that studies the replication, spread and evolution of memes (Moritz, 1990).

To be replicated, a meme must pass successfully through four subsequent stages: 1) assimilation by an individual, who thereby becomes a host of the meme; 2) retention in that individual’s memory; 3) expression by the individual in language, behavior or another form that can be perceived by others; 4) transmission of the thus created message or meme vehicle to one or more other individuals. This last stage is followed again by stage 1, thus closing the replication loop. At each stage there is selection, meaning that some memes will be eliminated. The present paper will look in more detail at the mechanisms governing these four stages, and present a list of selection criteria that allow us to estimate the fitness of a meme relative to its competitors.

The four stages of meme replication


A successful meme must be able to “infect” a new host, that is, enter into its memory. Let us assume that a meme is presented to a potential new host. “Presented” means either that the individual encounters a meme vehicle, or that he or she independently discovers it, by observation of outside phenomena or by thought, i.e. recombination of existing cognitive elements. To be assimilated, the presented meme must be respectively noticed,understood and accepted by the host. Noticing requires that the meme vehicle be sufficiently salient to attract the host’s attention. Understanding means that the host recognizes the meme as something that can be represented in his or her cognitive system. The mind is not a blank slate on which any idea can be impressed. To be understood, a new idea or phenomenon must connect to cognitive structures that are already available to the individual. Finally, a host that has understood a new idea must also be willing to believe it or to take it serious. For example, although you are likely to understand the proposition that your car was built by little green men from Mars, you are unlikely to accept that proposition without very strong evidence. Therefore, you will in general not memorize it, and the meme will not manage to infect you.


The second stage of memetic replication is the retention of the meme in memory. By definition, memes must remain some time in memory, otherwise they cannot be called memes. The longer the meme stays, the more opportunities it will have to spread further by infecting other hosts. This is Dawkins’s (1976) longevity characteristic for replicators.

Just like assimilation, retention is characterized by strong selection, which few memes will survive. Indeed, most of the things we hear, see or understand during the day are not stored in memory for longer than a few hours. Although you may have very clearly assimilated the news that the progressive liberal party won the Swaziland elections with 54% of the votes, you are unlikely to remember anything of this a week later–unless you live in Swaziland, perhaps. Retention will depend on how important the idea is to you, and how often it is repeated, either by recurrent perception or by internal rehearsal. All learning paradigms agree that experiences are encoded more strongly into memory by frequent reinforcement.


To be communicated to other individuals, a meme must emerge from its storage as memory pattern and enter into a physical shape that can be perceived by others. This process may be called “expression”. The most obvious means of expression is speech. Other common means for meme expression are text, pictures, and behavior. Expression does not require the conscious decision of the host to communicate the meme. A meme can be expressed simply by the way somebody walks or manipulates an object, or by what he or she wears.

Some retained memes will never be expressed, for example because the host does not consider the meme interesting enough for others to know, uses it unconsciously without it showing up in his or her behavior, does not know how to express it, or wants to keep it secret. On the other hand, the host may be convinced that the meme is so important that it must be expressed again and again to everybody he or she meets.


To reach another individual, an expression needs a physical carrier or medium which is sufficiently stable to transmit the expression without too much loss or deformation. Speech, for example, uses sound to transmit an expression, while text will be transmitted through ink on paper or electrical impulses in a wire. The expression will take the form of a physical signal, modulating the carrier into a specific shape from which the original meme can be re-derived. This physical shape may be called the meme vehicle. For example, meme vehicles can be books, photographs, artefacts or CD-ROMs.

Selection at the transmission stage happens through either elimination of certain memes, when the vehicle is destroyed or gets corrupted before it is perceived by another individual, or through differential multiplication, when the vehicle is reproduced into many copies. For example, a manuscript may be put into the shredder or it may be turned into a book which is printed in thousands of copies. A radio communication may get lost because of noise, or it may be broadcasted to millions of listeners. Especially since the emergence of mass media, the transmission stage is the one where the contrast between successful and unsuccessful memes is largest, and where selection may have the largest impact.

Meme fitness

The overall survival rate of a meme m can be expressed as the meme fitness F(m), which measures the average number of memes at moment t divided by the average number of memes at the previous time step or “generation” t – 1. This fitness can be expressed in a simplified model as the product of the fitnesses or survival rates for each of the four stages, respectively assimilation A, retention R, expression E and transmission T:

F(m) = A(m) . R(m) . E(m) . T(m)

A denotes the proportion of memes vehicles encountered (or memes independently discovered) by the host that are assimilated. R represents the proportion of these assimilated memes that are retained in memory. Therefore, A <= 1, R <= 1. E is the number of times a retained meme is expressed by the host. T is the number of copies of an expression that is transmitted to a potential new host. Unlike A and R, E and T do not have an upper bound, although E is likely to be more restricted than T. Note that F is zero as soon as one of its components (A, R, E, T) is zero. This expresses the fact that a meme must successfully pass through all four stages in order to replicate. Also note that for a meme to spread (F > 1), you must have E > 1 or T > 1.

General Selection Criteria for Memes

Which memes will most successfully pass all these stages can be modelled by a series of selection criteria. These criteria are discussed in more detail in earlier papers (Heylighen, 1993, 1997). I will here basically situate them with respect to the four replication stages. The criteria can be grouped into different families, distinguished by the system responsible for the selection. At present, we have no method to derive the value of the fitness components from the degree to which a meme fulfils the different criteria. This does not mean that no predictions can be made, though. All other things being equal, a meme that scores better on one of these criteria is predicted to become more numerous in the population than a meme that scores worse.

This is a falsifiable hypothesis, which can be tested through experiments or observations. For that, it suffices to operationalize the tested criterion. This has already been done for criteria such as invariance (Van Overwalle & Heylighen, 1995), formality (Heylighen & Dewaele, 1998) or conformity (cf. Boyd & Richerson, 1985), and seems relatively easy to do for the others as well by using standard social science methodologies, e.g. for developing test for personality traits.

Objective Criteria

Objective criteria denote selection by phenomena or objects independent of the hosts and memes involved in the process. The distinctiveness criterion functions mainly during the assimilation stage. It states that phenomena that are distinct, detailed or contrasted are more likely to be noticed and understood, and therefore assimilated. The invariance and controllability criteria, on the other hand, apply mainly to the retention stage. According to the invariance criterion, phenomena that recur, independently of the way in which they are perceived, are more likely to be maintained in memory. Controllability notes that phenomena which react differentially to the subject’s actions are also more likely to leave a permanent memory trace.

Subjective Criteria

Subjective criteria represent selection by the subject who assimilates the meme. The main criteria at the assimilation stage are novelty (facilitates assimilation by attracting the subject’s attention) and simplicity (requires less processing for the meme to be understood). The criterion of coherence (connection, consistency and support between new perception and existing memory trace) facilitates the understanding and acceptance parts of the assimilation stage, since it represents the ease with which the new meme can “fit in” with the memory that is already there. It also facilitates the retention stage since memories that cohere are more easy to retrieve and use and are therefore less likely to be forgotten. The criterion of utility, like controllability, functions mainly at the retention stage, since useful memes are more likely to be effectively used and thus reinforced, although it will also help assimilation, by making it more worthwhile for the host to do the effort to assimilate.

Intersubjective Criteria

Intersubjective criteria represent selection through the interactions between different subjects. Group utility is an emergent criterion, that is implicit in all four stages: a memes that is useful to the group of all its hosts is more likely to survive because it helps the group itself to survive and grow, and thus to absorb other individuals. Authority functions mainly at the assimilation stage: memes from authoritative sources, i.e. hosts or vehicles that are held in high regard or considered to represent expertise in the domain, will be more easily noticed and accepted. Formality (i.e. precise, unambiguous expression) too helps assimilation, at least of the original memetic content of the expression. It will contribute basically to what Dawkins (1976) calls copying-fidelity. (On the other hand, informal expression, because it tends to be simpler, may facilitate assimilation, but of an idea different from the one initially expressed). Conformity, the reinforcement of the same meme by different hosts belonging to the same group, will boost acceptance and retention (cf. Boyd & Richerson, 1985). Expressivity, the ease with which the meme can be expressed in an intersubjective medium, will obviously contribute to the expression stage. Publicity, finally, the effort put by the host(s) into the broad distribution of the message, will maximize transmission.

Meme-centered Criteria

Finally, the meme-centered criteria represent selection on the level of the meme itself. They depend only on the internal structure of the meme, not on its “fit” to external selectors, such as subjects, objects, or groups. These criteria will typically select for “selfish” (cf. Heylighen, 1992) or “parasitic” (cf. Cullen, 1998) memes, whose only goal is to spread themselves, “infecting” a maximum of hosts without regard for their hosts’ well-being. This does not imply that the same meme cannot satisfy both selfish and non-selfish criteria. Religions often have this mixture of parasitic and beneficial traits (cf. Cullen, 1998)

Self-justification, the degree to which the components of a meme mutually support each other, will facilitate understanding and acceptance. Self-reinforcement, the degree to which the meme stimulates its host to rehearse itself, e.g. by repetition, meditation, prayer, etc., will strengthen retention. Intolerance, the degree to which a meme excludes rival memes from being assimilated or retained, will also help the meme to retain a stable position in memory. Proselytism, the degree to which the meme urges its host to maximally spread the meme to other hosts, will increase the rates of expression and transmission.

stages\selectors Objective Subjective Inter-subjective Meme-centered
Assimilation distinctiveness novelty simplicity


authority formality self-justification
Retention invariance controllability coherence utility conformity self-reinforcement intolerance
Expression expressivity proselytism
Transmission publicity proselytism

Table 1: a summary of the main selection criteria for memes, classified according to the stage during which they are most active, and the system responsible for the selection.


This simple four stage model helps us to analyse the mechanics of meme replication, and the different requirements a meme must satisfy to spread successfully. It moreover helps us to situate and to systematize a more intuitively developed list of objective, subjective, intersubjective and meme-centered selection criteria. Although the four stage model suggests a formula for calculating memetic fitness, the theory is as yet insufficiently developed to unambiguously determine the parameters of the equation. However, the list of selection criteria does produce a range of qualitative predictions, which can be empirically tested.


Francis Heylighen has been supported during these investigations by the Fund for Scientific Research – Flanders (FWO) as a Research Associate.


Boyd R. & Richerson P.J. (1985): Culture and the Evolutionary Process, (Chicago University Press, Chicago).

Cavalli-Sforza L.L. & Feldman M.W. (1981): Cultural Transmission and Evolution: a quantitative approach, (Princeton University Press, Princeton).

Cullen B. (1998): “Parasite Ecology and the Evolution of Religion”, in: Heylighen F. & Aerts D. (ed.) (1998): The Evolution of Complexity (Kluwer Academic, Dordrecht)

Dawkins R. (1976): The Selfish Gene, (Oxford University Press, New York).

Heylighen F. (1992) : “Selfish Memes and the Evolution of Cooperation”, Journal of Ideas , Vol. 2, #4, pp 77-84.

Heylighen F. (1993): “Selection Criteria for the Evolution of Knowledge”, in: Proc. 13th Int. Congress on Cybernetics (Association Internat. de Cybernétique, Namur), p. 524-528.

Heylighen F. (1997): “Objective, subjective and intersubjective selectors of knowledge”, Evolution and Cognition 3:1, p. 63-67.

Heylighen F. & Dewaele J-M. (1998): “Formality of Language: definition, measurement and behavioral determinants”, submitted to Behavioral and Brain Sciences

Moritz E. (1990): “Memetic Science: I – General Introduction”, Journal of Ideas 1, p. 1-23

Van Overwalle, F., & Heylighen, F. (1995): “Relating covariation information to causal dimensions through principles of contrast and invariance”, European Journal of Social Psychology,25, p. 435-455.

Cultural Evolution and Memetics

Article prepared for the Encyclopedia of Complexity and System Science

Francis Heylighen & Klaas Chielens

Evolution, Complexity and Cognition group Vrije Universiteit Brussel



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