Reconsidering Cultural Selection Theory

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Brit. J. Phil. Sci. 59 (2008), 455479

Reconsidering Cultural SelectionTheory

G. K. D. Crozier

ABSTRACT

This paper examines conceptual issues that arise in applications of Darwinian naturalselection to cultural systems. I argue that many criticisms of cultural selectionist models

have been based on an over-detailed reading of the analogy between biological and

cultural units of selection. I identify five of the most powerful objections to cultural

selection theory and argue that none cuts to its heart. Some objections are based on

mistaken assumptions about the simplicity of the mechanisms of biological heredity.

Other objections are attributable, rather, to mistaken inferences from observations of

biological subject matter to what is essential in natural selection. I argue that such

features are idiosyncratic of biological systems, but not essential for natural selection.

My arguments throughout are illustrated by examples from biological and cultural

evolution, and counter-factual illustrations from the history of theoretical biology.

1 Introduction

2 Cultural Selection Theory

3 First Objection: Lamarckianism

4 Second Objection: GenotypePhenotype Distinction

5 Third Objection: Common Hereditary Architecture

6 Fourth Objection: Biological Analogue for Cultural Units

6.1 Regarding strict analogues

6.2 Regarding the trait analogue

6.3 Regarding the virus analogue

7 Fifth Objection: Environmental Interaction

8 Conclusion

1 Introduction

The mechanism of evolution by natural selection has been applied to fields

as diverse as immunology, computation, neural programming, and cultural

change. Though selectionist explanations of cultural change seem plausible and

show potential for providing valuable insight, this field has not developed into a

strong research area. It lacks several key features, including a widely accepted,

CThe Author(2008). Publishedby Oxford University Press on behalf ofBritish Society for the Philosophyof Science. All rights reserved.

doi:10.1093/bjps/axn018 For permissions, please email: [email protected]

Advance Access published on July 22, 2008

atTheUniversityofMontanaon

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456 G. K. D. Crozier

clear case study and an account of the role of environmental variables in the

evolution of cultural traits.

This paper examines the conceptual issues that arise in applications of Dar-

winian natural selection to cultural systems, defending cultural selectionist

models that are somewhat analogous to models of genetic selection. Many

criticisms of cultural selectionist models have been based on an over-detailed

reading of the analogy between biological and cultural units of selection. Specif-

ically, such objections are based on misconceptions about the relationships

between biological evolution, cultural evolution, and the theory of natural

selection.

2 Cultural Selection Theory

Cultural selection theory studies cultural changes by modelling them after

theories of biological evolution. Three prominent approaches to cultural se-

lection are social contagion theory, evolutionary epistemology, and memetics.

Social contagion theorys epidemiological approach construes social entities

as analogous to parasites that are transmitted virally through a population of

biological organisms. Evolutionary epistemologys focus lies in causally con-

necting evolutionary biology and rationality by generating explanations for

why traits for rational behaviour or thought patterns would have been selected

for in a species evolutionary history. Memetics models cultural change after

population genetics, taking cultural units to be analogous to genes.

This essay is concerned with systems of cultural evolution that are unam-

biguously non-strategic: that is, the mechanisms underlying the systems of

interest are not forward-thinking or rational in any sense, but rather result

from blind variation and selective retention of characteristics resulting from

environmental interaction. Some theorists contend that even quintessentially

strategic, forward-thinking, rational processes are essentially the product of

non-strategic, Darwinian selection. For example, some of Poppers ([1972])work in the philosophy of science and Campbells ([1974]) evolutionary episte-

mology can be interpreted as supporting this view. This essay, however, is not

concerned with such processes.

Most of the debate to be considered takes place within the context of memet-

ics, but it applies more broadly as well. For example, in distinction from sup-

porters of memetics, I am committed neither to finding a cultural analogue of

the gene nor to the existence of some cultural unit that is faithfully replicated

from generation to generation. What I am committed to is searching for cultural

adaptations and the mechanisms responsible for them on a case-by-case basis,

and to remaining open-minded about what specific forms those mechanisms

can take within natural selection theory. I argue that we can learn from the

history of evolutionary biology which elements are truly central to selection


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Reconsidering Cultural Selection Theory 457

theory and which only appear to be so due to the successes of contemporary

genetics and the idiosyncrasies of the biological medium. Nevertheless, since

much of the relevant literature uses the terminology of memes and memetics,

this paper will make use of that vocabulary.

Although analogies had been drawn between cultural and biological evolu-

tion for some time, it was in the 1970s that memetics arose as a recognizable

metaphor around which to structure investigations and build a theory of cul-

tural evolution by natural selection analogous to genetics. Dawkins ([1976])

coined the term meme to refer to cultural units that are differentially repli-

cated during a process of natural selection, analogous to biological genes. The

memetic framework excited a generation of researchers with the possibility of

replicating the success of molecular biology. Despite all this attention, how-

ever, little coherence has been generated within the field of memetics. Thesystems that are chosen for memetic analyses are as diverse as the interests

of their authors: fads, ideologies, religious symbols, rumours, art, social insti-

tutions, chess moves, tool use, and so on. Rarely will two authors tackle the

same subject matter. Further, many of the cultural systems discussed in the

literature are hypothetical cases introduced to support a theoretical or defini-

tional point, and are without any empirical content. The consequent debates

rarely reach community consensus, and disagreements pass out of attention

due to their resistance to resolution rather than because theorists have reached

agreement.

The 35-year inability of memeticists to find compelling evidence or de-

vise decisive investigations in order to narrow the candidate hypotheses in-

dicates to some that memetics has failed to describe any robust causal features

of the cultural landscape. For example, Lanier ([1999], p. 2) contends that

memetics:

[. . .] is so variable as to provide no fixed target [. . .] Are memes a rhetorical

technique, a metaphor, a theory, or some other device? Depending on whoyou talk to, they can be so wispy as to be almost nothing [. . .] They make no

predictions and cannot be falsified. They are no more than a perspective.

Similarly, Gardner ([2000]) argues that:

[. . .] memetics is no more than a cumbersome terminology for saying what

everybody knows and that can be more usefully said in the dull terminology

of information transfer [. . .] A meme is so broadly defined by its proponents

as to be a useless concept, creating more confusion than light.

Recently, Edmonds ([2005]) dissolved the Journal of Memetics, of which he

was the founder and editor. He contends that memetic studies have produced

nothing more than whatcould be found in cultural evolutionary models without


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genetic analogies, and that this has resulted in a loss of academic interest in the

field. Edmonds ([2005]) writes:

I claim that the underlying reason memetics has failed is that it has not

provided any extra explanatory or predictive power beyond that availablewithout the genememe analogy. Thus whilst the idea of memes has re-

tained its attractiveness for some in terms of a framework for thinking

about phenomena, it has not provided any added value in terms of provid-

ing new understanding of phenomena. The fact that some who wear the

theoretical spectacles (Kuhn [1969]) of memetics insist on redescribing a

host of phenomena in these terms despite the lack of substantive results

merely confirms other academics opinion of the approach. The ability to

think of some phenomena in a particular way (or describe it using a certain

framework) does not mean that the phenomena has those properties in any

significant sense.

Academics who seek to study memetics in serious ways have suffered in

the respect that they are often confused with those on the penumbra for

whom memetics is a fad. However, this mistake is grounded in an element

of truth. The study of memetics has been characterized by theoretical

discussion of extreme abstraction and over ambition. Thus for example,

before any evidence is available or detailed causal models constructed,

attempts have been made to explain some immensely complex phenomena

such as religion in general or consciousness. This sort of discussion shifts

any study of memetics from the realm of science to that of philosophy and,on the whole, this philosophy has adopted the subsumption tactic (Hull

[2001]), seeking to generalize explanation rather than been productive of

essentially new insights.

I will review some theoretical debates regarding the application of Darwins

theory of natural selection to cultural evolutionary studies. One theme that

will emerge in these critiques is that many objections are based on disanalogies

between the mechanisms of biological evolution and their proposed cultural

counterparts. I will argue that critiques based on disanalogies between the me-dia of biological and cultural evolutionary systems rely on two categories of

controversial claims. First, these arguments rely on claims concerning the ease

with which the biological subject matter can be carved at the joints when con-

trasted with the relatively arbitrary distinctions that apparently characterize

their cultural counterparts. I will argue that these claims lose their force on

closer inspection. Second, these arguments rely on premises concerning what

auxiliary assumptions are incompatible with the theory of evolution by nat-

ural selection. I will argue that these premises are vulnerable to charges of

anachronism.

I contend that both proponents and critics of cultural evolutionary studies

must abandon their shared hidden premise that biological evolution is the

archetypal example of evolution by natural selection rather than merely one


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of several domains in which this process takes place. Further, until we have

case studies to demonstrate the viability of cultural selection theory, it may be

premature to continue pursuing theoretical debates.

3 First Objection: Lamarckianism

One predominant critique of cultural evolutionary studies is that it is inherently

Lamarckian. Lamarckian evolution has long been disconfirmed in biology,

supplanted by a Darwinian theory of blind variation and selective retention.

Consequently, when an evolutionary theory is charged with being Lamarckian,

generally this is meant to imply that the theory is seriously flawed. I will argue

that the charges of Lamarckianism do not undermine cultural evolutionary

studies. First, cultural evolution is not inherently Lamarckian. Charges to thecontrary are based upon misconceptions of the hereditary pattern of cultural

units. Second, even if cultural evolution did proceed by Lamarckian means,

the application of Darwinian natural selection in these domains would still

be possible. Darwinian and Lamarckian mechanisms can coexist in the same

system.

A generation before Darwin, Lamarck advocated the theory of adaptation

by the inheritance of acquired characteristics: the theory that evolutionary

adaptations are the cumulative effects of the use or disuse of organs during the

lifetimes of organisms. Lamarckian inheritance occurs when characteristics ac-

quired by predecessors during their lifetimes are inherited by their successors.

In its literal sense, evolution is Lamarckian if the environment changes the

phenotype of an organism so that this organism is better adapted to the envi-

ronmental factor that produced this change, and if the change is transmitted

somehow to the heritable material so that it can be passed on to the offspring of

the organism through reproduction. In Darwinian evolutionary theory, on the

other hand, heritable variations are random with respect to the fitness effects

they will have when the organism interacts with its environment.Cultural evolution may appear, at first glance, to be inherently Lamarckian

since cultural characteristics are acquired by an organism during its life and

can subsequently be passed on to its offspring. Some caution is in order here:

it must be remembered that cultural evolution is concerned with the evolution

of units of culture, rather than with the evolution of biological organisms.

Although the biological organisms phenotype is modified by the introduction

of new cultural traits during its lifetime, cultural evolution is concerned, rather,

with tracking changes in cultural systems themselves. In cultural evolution, it is

the effects of selection pressures on cultural entities that are of interest, as well

as the effects of these pressures on cultural populations to generate adaptations

or frequency changes. Biological organisms fade from view under cultural

evolutionary description since they participate in only part of the lifecycle of


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the cultural entities: biological entities are the medium of the copying process

of the cultural entities.

Hull ([2000], p. 56) puts the point this way:

For example, you did not understand the Pythagorean Theorem. You take

a course in plane geometry, and now you do. You have acquired a new

meme. You in turn can pass this increased knowledge on to someone

else. Isnt this an instance of the inheritance of acquired characteristics?

Not in the least. In the science of memetics, memes are analogous to

genes, not phenotypic characteristics. Hence, if memetics is anything, it is

the inheritance of acquired memes. How passing on memes (or fleas for

that matter) can count as Lamarckian inheritance in any comprehensible

fashion continues to elude me. It is made to look plausible only by running

together the genetic with the memetic perspectives.

If we were to draw a genealogical tree in memetics, it would be from the

point of view of the cultural unit rather than the organism. Speel ([1997])

coined the terms memotype and phemotype to refer to the cultural counterparts

of the biological genotype and phenotype, and I will follow this terminology

throughout. Only if thephemotypesnot phenotypesmanifest characteristics

that are both acquired during their own lifecycles and heritable to further

generations of cultural progeny, is their evolution properly to be described as

Lamarckian. Since it is the evolution of cultural and not biological units thatis under examination, cultural evolution is not inherently Lamarckian.

However, there may be genuine cases of Lamarckian cultural evolution. Still

more importantly, the discovery of such cases would neither undermine the ve-

racity of cultural evolutionary theory, nor would it devalue Darwinian natural

selectionist approaches to the description of cultural evolutionary systems. It

is generally conceded that Lamarckian evolution does not take place in bio-

logical evolution, since there is no mechanism by which changes to a parents

phenotype can cause corresponding changes to its genetic material thereby

making those changes transmittable to its offspring. (Epigenetic mechanisms

are one possible exception.) However, although Lamarckianism is often mis-

understood to be in tension with Darwinian evolution by natural selection, the

two mechanisms are compatible. Darwin himself contended that some evolu-

tion was likely the result of Lamarckian adaptation of acquired characteristics.

Cultural evolution is not inherently Lamarckian, but if cases of Lamarckian

evolution are discovered, they will not undermine cultural selection theory.

4 Second Objection: GenotypePhenotype Distinction

The identification of an instance of biological evolution as Lamarckian or Dar-

winian hinges on the genotypephenotype distinction. This is the foundation

of some objections to the importation of selection theory to cultural systems.


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Some cases of cultural evolution support the identification of phemotype

and memotype, while others resist the application of this distinction. Consider,

for example, the reproduction of chain letter e-mails that are coded in html.

The underlying html code of the e-mails can be identified as the memotype; the

version visible to the user, who may or may not forward it, can be identified

as the phemotype. Other casesbird songs, for examplehave no obvious

memotype and appear to be all phemotype. In such cases, the same cultural

entity seems to be serving two roles that are generally distinct in biological

evolutionary processes: replicator and vehicle/interactor.

I will argue that this conflation of these two kinds of units of selection,

held to be importantly distinct in evolutionary biology, should not undermine

the application of natural selection to the cultural domain. This is because the

distinction between these roles is based on a consensus that arose from the unitsof selection debate in theoretical biologya consensus whose conceptual basis

is potentially flawed: even in traditional evolutionary biology this distinction

does not necessarily apply.

At the forefront of the units of selection debate, Williams and Dawkins

supported genes as the true units of selection, while Mayr and Gould argued

that individual organisms hold this position. The resolution to this debate

by Hull and Dawkins through the replicatorvehicle/interactor distinction

convinced many theorists that natural selection requires something in the role

of the replicator. It is this that I wish to question.

Williams ([1966]) bookkeeping argument (a term coined by Wimsatt

[1980], [1981] in a criticism of genetic reductionism) advocated genes as the

proper unit of selection. He supported this on the basis that evolution-

ary biologists, and in particular population geneticists, are interested pri-

marily in tracking frequencies across generations. Williams wrote ([1966],

pp. 224):

The essence of the genetical theory of natural selection is a statistical biasin the relative rates of survival of alternatives (genes, individuals, etc.).

The effectiveness of such bias in producing adaptation is contingent on

the maintenance of certain quantitative relationships among the operative

factors. One necessary condition is that the selected entity must have a high

degree of permanence and a low rate of endogenous change relative to the

degree of bias (differences in selection coefficients). Permanence implies

reproduction with a potential geometric increase.

Acceptance of this theory necessitates the immediate rejection of the im-

portance of certain kinds of selection. The natural selection of phenotypescannot in itself produce cumulative change, because phenotypes are ex-

tremely temporary manifestations. [. . .] Socrates consisted of the genes his

parents gave him, the experiences they and his environment later provided,

and a growth and development mediated by numerous meals. [. . .] If the


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hemlock had not killed him, something else soon would have. So however

natural selection may have been acting on Greek phenotypes in the fourth

century B.C., it did not of itself produce any cumulative effect. [. . .] The

same argument also holds for genotypes. With Socrates death, not only

did his phenotype disappear, but also his genotype. [. . .]

It is only the meiotically dissociated fragments of the genotype that are

transmitted in sexual reproduction, and these fragments are further frag-

mented by meiosis in the next generation. If there is an ultimate indivisible

fragment it is, by definition, the gene that is treated in the abstract discus-

sions of population genetics.

Since biological individuals have a population frequency approaching zero,

Williams argued that genes rather than phenotypes are the units on whichnatural selection operates.

Building on the work of Williams, Dawkins ([1976]) also placed genes at the

centre of natural selection, with the units of selection defined as those things that

produce copies of themselves. Boyd and Richerson ([2000], p. 155) summarize

this conception of the units of selection as consisting of the following three

qualities:

(1) Copying Fidelity. The copying must be sufficiently accurate that

even after a long chain of copies the replicator remains almost

unchanged;

(2) Fecundity. At least some varieties of the replicator must be capable of

generating more than one copy of themselves;

(3) Longevity. Replicators must survive long enough to affect their own

rate of replication.

Dawkins argued that genes are the only biological units with sufficient fecun-

dity, longevity, and stability to be subject to the multiple iterations of selection

required for complex adaptations to arise.Mayr ([1963]) and Eldredge and Gould ([1972]), to the contrary, argued that

individuals are the proper units of selection because it is their properties that

interact with the environment to determine how many offspring will be pro-

duced in the next generation. They contended that, since gene frequencies only

change directionally because of environmental interaction with phenotypes,

individuals (phenotypes) are the proper units of selection.

This debate was accepted by many to have been resolved when Hull ([1981])

and Dawkins ([1982]) independently argued that there are two kinds of

units of selection (Reeve and Keller [1999]). Each party in the debate over

genes versus individuals was correct in identifying one of these two kinds of

units of selection. Hull ([1981], p. 33) describes these two types of units as

follows:


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Replicatorsentities that pass on their structure directly in replication;

Interactorsentities that produce differential replication by means of

directly interacting as cohesive wholes with their environments.

Replicators are units that can be replicated in subsequent generations. Theyrepresent the effects of selection by virtue of the changes in frequency with

which they appear in subsequent generations. This is defined in terms of copying

fidelity, fecundity, and longevity according to the specifications of Williams

and Dawkins. Vehicles (or interactors in Dawkins [1982] terminology) are units

subject to the direct causal effects of external selection pressures, and they

accord with Mayr, Eldredge, and Goulds notion of the units of selection.

The numbers in which replicators are represented in subsequent generations

are dependent upon the interactions between the selective environments and

the vehicles/interactors that are created by replicators, and through which

replicators are transmitted into the next generation.

Boyd and Richerson ([2000]) argue that the resolution to the units of selection

debate offered by Hull and Dawkins is based on the misconception that it is

necessary for replicators to exemplify near-perfect copying fidelity. They argue

that the term replicator is a potentially misleading one since it is correlation

between generations that is required for evolution by natural selection and

not replication per se: replication is one kind of correlation that can lead to

evolution by natural selection, but not the only kind. Boyd and Richerson([2000], p. 158) write:

We do not understand in detail how culture is stored and transmitted, so we

do not know whether culturally transmitted ideas and beliefs are replicators

or not. If the application of Darwinian thinking to understanding cultural

change depended on the existence of replicators, we would be in trouble.

Fortunately, culture need not be closely analogous to genes. Ideas must

be gene-like to the extent that they are somehow capable of carrying the

cultural information necessary to give rise to the cumulative evolution of

complex cultural patterns that differentiate human groups. They exhibitthe essential Darwinian properties of fidelity, fecundity and longevity, but,

as the example of phonemes shows, this can be accomplished by a most

ungene like, replicatorless process of error-prone phenotypic imitation.

All that is really required is that culture constitutes a system maintaining

heritable variation.

To this end, Boyd and Richerson advocate abandoning the genocentricity of

a memetic approach, and in particular the view that cultural units must be

replicators that are capable of being reproduced with near-perfect fidelity. While

we do need the reproduction or emulation of a prototype, we do not need

perfectly faithful reproduction in the sense of Dawkins and Hulls replicator.

Griesemer ([unpublished]) is currently working on a conception of reproduction

that may suit the needs of cultural selection.


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5 Third Objection: Common Hereditary Architecture

Another challenge levied against cultural selection theory is that, since there

is no hereditary architecture that is common to cultural systems, it is impos-

sible to consistently identify the units of cultural selection. Wimsatt ([1999])argues that the absence of such universal memetic architecture makes it too

difficult to define the cultural units of selection and environmental interaction;

if this is correct, he argues, models of cultural selection analogous to models

of genetic selection cannot possibly be enlightening. Problems in generating

standard memetic models crop up in parentage, and flow from there to deriva-

tive problems in characterizing cultural analogues to gametes, genomes, and

species (Wimsatt [1999], p. 281). Wimsatt claims that the comparative success

of biological models of natural selection is facilitated by the presence of a ge-

netic, hereditary architecture that is common to all systems; the participation

of all biological systems in common hereditary patterns greatly facilitates the

identification of genes and gene-centred models.

Wimsatts argument identifies some of the problems central to any memetic

approach to selectionist interpretations of cultural evolution. I will argue, how-

ever, that this path to a model of cultural evolution still has merit.

Wimsatt argues that it is not possible to identify the cultural analogues of

the units of selection and environmental interaction. It is not obvious, however,

that biological evolution is intrinsically clearer than cultural evolution in thisway. First, the common hereditary architecture shared by biological systems

certainly helps researchers to make generalizations; however, the identification

of this architecture was facilitated by prior observations of variations in the

frequency of traits in biological populations over time and geographic regions.

The success of genetics is attributable, in part, to the long tradition of research

in this area; by comparison, the study of cultural evolution in this area has

just recently begun. Darwin developed evolutionary theory quite extensively

without much information about the mechanisms of variation and heredity, or

of Mendelian genetics.

Second, the genetic architecture does not guarantee a hereditary structure

sufficiently rigid that the units of selection are obvious. Contrary to Wimsatts

claim, there are many biological systems that defy rigid definitions of parentage.

Wimsatt ([1999], p. 281) claims that, unlike cultural cases, there are no cases in

biology where:

(1a) an individual has more than two parents [. . .];

(1b) the parental contributions from different parents are radically unequal

[. . .];

(1c) any of these parametersthe number of parents, the social role which

places them in a parenting relation, or their relative contributionscan fluc-

tuate irregularly from generation to generation.


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These claims are inaccurate since biological systems do violate these categories.

Three examples of biological systems that do not approximate the standard

single or double parental hereditary models are bees, fish, and bacteria.

A female bee inherits two-thirds of her genes from her mother and one-

third from her father, but a male bee receives all of his genes from his mother.

Thus, bee heredity does not obey point (1b) since the parental contributions

are radically unequal.

The South American Amazon Molly (Poecilia formosa) is a particularly fas-

cinating fish. Most generations of this species consist of asexually reproducing

female clones. Amazon Mollies produce fully viable clone eggs; however, sperm

are necessary to trigger the developmental processes. Thus, an Amazon Molly

will mate with males of either of the related species Atlantic Molly (Poecilia

mexicana) or Sailfin Molly (Poecilia latipinna). Usually this mating only servesto trigger the cloning process; however, occasionally (and the necessary condi-

tions are still not well understood) the mating does result in cross-fertilization

between the species. Surprisingly, the offspring of these pairings are not hybrids

of the two species, but develop into female, asexually reproducing Amazon

Mollies. Therefore, contrary to point (1c), the Amazon Molly irregularly varies

its parental hereditary contributions.

Should it appear that I have been excavating obscure examples that are no-

table for being biologically exceptional, I present a phenomenon that is as

common as they come. Bacteria undergo a process of genetic exchange called

conjugation during which one bacterium will insert a portion of its genetic

material into another bacterium. The recipient bacterium will pass on its mod-

ified genome to its daughter-clone progeny. In theory, a donor bacterium can

transfer its entire genome to a recipient, but conjugation is usually interrupted

by contingent, environmental interventions. Although bacterial reproduction

takes place by asexual cloning, the number of individuals from which an off-

spring has obtained its hereditary information is limited only by the number of

cells that have engaged its mother cell in conjugation. This contradicts point(1a) that there are never more than two parents. It also contradicts point (1b),

since the parental contributions are radically unequal. Finally, bacterial conju-

gation also undermines point (1c), since the number of parents and the degree

of their contributions vary from generation to generation and from cell to

cell. Despite these complications, biologists have successfully pursued genetic

explanations of bacterial evolution.

These examples indicate that the same complications responsible for

Wimsatts first obstacle to a memetic approach to cultural evolution also apply

to biological evolution. Indeed, Hamiltons ([1964a], [1964b]) description of

the haplo-diploid structure of bee inheritance has been celebrated as a triumph

of genetic selectionism (Dawkins [1976]). Since a genetic approach has been

successful despite flexible definitions of parentage in biological models, we may


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yet expect cultural models to follow their example. This further suggests that

cultural evolutionists can learn from the ways in which evolutionary biologists

have contended with analogous difficulties.

Wimsatts position implies that Darwin would not have been able to achieve

the successes he did had there not been some common hereditary architec-

ture, despite his ignorance of its structure. Wimsatt concludes that, since no

such gene-like architecture gives structure to cultural evolutionary systems, it

is unreasonable to expect memetics to produce comparable success. There is

insufficient reason, however, to rule out this approach a priori. Further, some

cultural systems do appear to be based upon very strict architectural con-

straints. What is needed is a case study to demonstrate the viability of memetic

approaches to cultural selection. I will return to this point later.

6 Fourth Objection: Biological Analogue for Cultural Units

Another source of controversy for cultural selection theory lies in the difficulty

of finding a biological analogue for cultural units. I will argue that a strict

biological analogue is not necessary for a cultural evolutionary model to be

successful. Furthermore, caution is advised when adopting any close analogy

between cultural and biological selection entities (such as traits or viruses) since

these can place inappropriate constraints on cultural selection theory.

6.1 Regarding strict analogues

Wimsatt ([1999]) contends that, to extend the genetic model to cultural change

successfully, cultural units must be characterizable as analogous to some par-

ticular biological unit, such as traits or viruses: a successful extension would

have to specify which entities in the new system are analogous to entities that

are central in the old model. But, he argues, a precise and consistent ana-

logue for memes with a biological evolutionary model has not been provided,the prospects for it are poor, and this undermines the feasibility of memetic

approaches to cultural evolution. Consequently, Wimsatt advocates a develop-

mental approach, which he contends is more open to a pluralist view of the

cultural units and whereby the categorization of cultural units is determined

by the characteristics of the particular system under investigation (Wimsatt

[1986]).

While I agree with Wimsatt that pluralism with respect to the biological ana-

logues of cultural units is warranted, I disagree that this undermines the value

of alternative approaches to cultural selection and necessitates a developmental

approach. For example, social contagion theory and memetics can be expected

to provide distinct descriptions of the same cultural system that are not only

fully compatible but also inter-reducible, much as biological viruses can be


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studied both epidemiologically and genealogically (Marsden [1998]): both ap-

proaches describe the same objects operating according to the same laws, yet

they assume distinct vantage points from which to describe these operations.

On the one hand, social contagion theorys epidemiological approach is hor-

izontal because it looks at a cross-section of the distribution and frequencies

of various cultural units throughout a biological population or other body.

This approach takes snapshots of this distribution at various points in time

and space and compares them to determine what factors might explain the

observed similarities and differences. On the other hand, memetics genealog-

ical or population genetic approach provides vertical descriptions of cultural

lineages or of changes in the frequencies of different versions (or alleles) of cul-

tural units. This approach tracks the continuous evolution of the unit through

successive generations.The horizontal and vertical descriptions of selection of the same evolution-

ary system are complementary because they reveal different facets of the same

process. There is good reason to remain open-minded about which, and in-

deed whether any, strict biological analogue is required for cultural selection

theoryeven one that is developmental.

6.2 Regarding the traitanalogue

Some analogues, however, are more vulnerable to misinterpretation than others.

For example, analogies between cultural units and the traits of organisms (very

common in evolutionary epistemology, but apparent in other approaches as

well) tend to shift focus from the differential fitnesses and consequent evolution

of cultural entities to the effects that cultural entities have on the differential

fitnesses and consequent evolution of biological organisms.

It should be remembered that the fitness of a cultural unit consists in its

ability to get itself reproduced and consequently to be represented in higher

proportions in the cultural milieu. This point is central to clarifying manymisconceptions about the application of natural selection to cultural evolution:

a cultural units reproductive success is not equivalent to the reproductive

successes of the organisms in whose cultural system the unit exists. Analogies

between cultural units and the traits of organisms blur this vital distinction.

For example, in the second edition of Philosophy of Biology, Sober ([2000])

contends that a model of cultural evolution cannot explain why a meme is so

psychologically attractive. I quite agree that an evolutionary model must restrict

itself to answering the questions what has evolved and how has it done so.

Why one trait is more fit than another in a given context cannot be addressed

until the evolutionary model is supplemented with information from another

source. An explanation of the fitness granted to a flying bird by its wings must

refereven if only implicitlyto the principles of aerodynamics (Christensen


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and Hooker [1999]). However, it is misleading for Sober to use the term psy-

chological attractiveness synonymously with cultural fitness because the factors

shaping the fitness of a cultural trait may be external to the psychological state

of the organism. While cultural traits might spend a phase of their reproductive

cycles in the psychologies of organisms, a good scientific explanation for why

a trait is fitter than its alternatives may require that more attention be paid to

non-psychological systems, such as those governed by the organisms habitat.

Sobers term psychological attractiveness is analogous to describing the fitness

of a physiological trait as its genetic attractivenessit is descriptively accurate

but explanatorily impoverished.

One of the interesting challenges of describing cultural evolutionary pro-

cesses is to explain the relative fitnesses of memes. This is further complicated

by the intricate interactions that can exist between the fitnesses of cultural unitsand the biological fitnesses of the organisms through which the cultural units

are propagated. Dawkins ([1976], p. 207) makes a point similar to Sobers when

discussing why the meme for godhas been so successful:

The survival value of the god meme in the meme pool results from its great

psychological appeal. It provides a superficially plausible answer to deep

and troubling questions about existence. It suggests that injustices in this

world may be rectified in the next. The everlasting arms hold out a cushion

against our own inadequacies which, like a doctors placebo, is none the

less effective for being imaginary. These are some of the reasons why the

idea of God is copied so readily by successive generations of individual

brains. God exists, if only in the form of a meme with high survival value,

or infective power, in the environment provided by human culture.

Some of my colleagues have suggested to me that this account of the

survival value of the god meme begs the question. In the last analysis they

wish always to go back to biological advantage. To them it is not good

enough to say that the idea of a god has great psychological appeal. They

want to know why it has great psychological appeal. Psychological appeal

means appeal to brains, and brains are shaped by natural selection of genes

in gene-pools. They want to find some way in which having a brain like

that improves gene survival.

Dawkins colleagues are motivated to pursue biological explanations because

the factors identified as responsible for psychological appeal require further

justification. This justification may be grounded in biological explanations, but

alternatives are available: epistemological, biochemical, cultural, and so on.

The greater the support for the causal categorization of the relevant memetic

environment, the more compelling will be the case that a genuine cultural

adaptation exists.

Elsewhere, Dawkins refers to blind faith as a successful cultural adapta-

tion: The meme for blind faith secures its own perpetuation by the simple


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unconscious expedient of discouraging rational inquiry (Dawkins [1976],

p. 213). If it is true that once we adopt the cultural trait of blind faith the

only way to eliminate it is rational inquiry, but that the trait itself undermines

rational inquiry, then the only way to avoid carrying this trait is to not adopt it

in the first place. This example appeals to a notion of rationality in its assign-

ment of fitness, and as such can avoid circular reasoning. However, the purely

cultural or psychological nature of the traits environment sheds doubt on the

traits ontological status. Can we say that blind faith is itself a trait, or is it

rather a feature of other traits like religious belief and political ideologies? Is it

instead a kind of psychological predisposition?

Our ability to identify adaptations depends on the clarity with which we can

define the relevant causal relationships within the evolutionary environment.

Cultural evolution should not be expected to provide an explanation for the psy-chological attractiveness of a trait any more than biological evolution should ex-

plain why a birds wings are aerodynamically sound: that is where the theory of

aerodynamics comes in. An evolutionary theory, however, should indicate what

features of the environment an adaptation has the function of accommodating,

it should be able to predict that other features with similar functions would

have similar fitness advantages, and it should be able to accommodate contrary

evidence without appealing to ad hoc reasoning. The conceptual ambiguity

surrounding the fitness relationship between cultural and biological selection

units is responsible for many misconceptions about cultural selection, includ-

ing the Lamarckian objection. To avoid blurring this vital distinction, caution

is advised when using the traits of organisms as analogies for cultural units.

6.3 Regarding the virus analogue

The viral analogy is less treacherous than that of traits of organisms, insofar

as the fitnesses of viruses are understood to be importantly distinct from, and

indeed often in conflict with, those of their host organisms. The effect of acultural unit on the organisms through which it is copied will fall into one of

the following categories:

(1) Negative: Competition, such as a meme for suicide, which is clearly

detrimental to the fitness of the individuals demonstrating the

behaviour;

(2) Neutral: Commensalism, such as the meme for a particular nursery

rhyme, which is arguably irrelevant to the fitness of the individuals

repeating the rhyme;

(3) Positive: Mutualism, such as, arguably, a meme for a fashion trend,

which enhances the social statuses and thereby the fitnesses of the

individuals who have it.


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This viral approach to memetics provides a way to track the relationship be-

tween the fitnesses of organisms and memes while maintaining a distinction

between these two quantities. This distinction becomes confused when memes

are described as traits of organisms. The viral analogy lends itself well to cultural

selection by providing the conceptual tools for both horizontal epidemiological

descriptions and vertical population genetic approaches. It also provides the

means to clarify the value of this distinction.

Employing a viral analogy, however, requires care of a different sort.

Gatherer ([1998]) argues that memetics has been hindered by the description of

memes as contagious beliefs. The problem lies in the tendency to locate cultural

selection units in the minds of individualsfor example, as beliefs, skills, or

talents (Dawkins [1982], [1983])which renders the primary subject matter

of memetics essentially unobservable. Gatherer recommends that, to developmemetics as a credible empirical enterprise, models adopting a viral analogy

should be discarded in favour of models wherein memes are defined in terms of

observable properties such as songs, words, Windsor knots, and other cultural

artefacts (Dawkins [1976]).

In a sense, rather than viewing memes as objects possessedby individuals or

existing as patterns of information in the brains of individuals, the individuals

through which the memes are transmitted should fade from view. Gatherer

([1998]) writes:

If we are to have a cultural evolutionary theory, then we need units of

replication/selection, and the meme concept would seem to be as good a

way as any of approaching this problem. The mistake lies in the frequent

assumption that individuals have memes. But because, as will be shown,

we cannot assign memes to individuals with sufficient reliability or regu-

larity, we cannot produce meme frequencies, defined as the proportion of

individuals in a population that possess or have a meme. Gene frequencies

are absolutely necessary to population genetics, and if there is to be a pop-

ulation memetics we correspondingly need to have unambiguous meme

frequencies. There are some circumstances in which we can derive a statis-

tic of this sort but, as will be demonstrated, this kind of meme frequency

statistic, if we are to have it at all, must be defined in a purely behaviourist

manner.

Notice that this point dovetails with Wimsatts critique that memetics is

based on the absence of a common hereditary architecture underlying cultural

evolutionary processes, and also with the previous discussion on the replicator

interactor distinction. Though this observation leads Wimsatt to advocate a

developmental approach, Gatherer is led, rather, to suggest ways to overcome

this obstacle to memetic models of cultural selection.

Gatherer ([1998]) argues that memes must be defined as cultural artefacts

rather than as information patterns in the minds of individuals:


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By contrast, individuals do have genes. They may or may not pass them on

to the next generation of individuals. However, whatever the reproductive

success or otherwise of an individual, that individual carries those genes

around in its body for the duration of its life. A population geneticist may

derive allele frequencies as the proportion of individuals who carry onegene sequence or another. Gene frequencies are of course abstractions,

pure quantitative values, but they are abstractions which relate directly to

a physical reality, i.e. the reality of gene sequences within the bodies of

individuals. Thus, providing the required technical methods are available,

one may derive an unambiguous estimation of gene frequencies which are

comparable between one population and another.

Gatherer argues that we cannot assign these meme artefacts to individuals with

sufficient precision and regularity to measure meme frequencies in a population,

and that consequently population memetic studies are impossible.

Hull ([2000]) objects to Gatherers position, arguing that it rests on nave,

behaviourist assumptions that have long been rejected in the philosophy of

science. Though Gatherers argument is motivated by a desire to eliminate

commitments to unobservable entities (here mental entities or the beliefs of

others), Hull contends that it ultimately sneaks in a commitment to the non-

existence of these entities. Such a strong conclusion, Hull objects, would rule

out the inclusion of any theoretical entities in science. Of course genes, quarks,

and many other theoretical entities are not directly observed and yet are accept-able aspects of successful scientific theories. Still, Gatherers quasi-behaviourist

argument has a valuable lesson: caution must be taken when identifying theo-

retical entities in cultural selection theory. While viral approaches to cultural

selection need not altogether eliminate the introduction of theoretical entities,

these introductions should be minimized.

Although care is in order when adopting a strict viral analogy for cultural

units, just as it is for the analogy of traits of organisms, the latters propensity for

masking selective forces in cultural evolution is more pernicious. Nevertheless,

a strict analogy between biological and cultural selection is not necessary.

7 Fifth Objection: Environmental Interaction

The final source of controversy surrounding cultural selection that I will discuss

concerns the difficulty in distinguishing various phases in cultural evolutionary

processes, and in particular the role of environmental interaction. Wimsatt

([1999], p. 288) writes:

In genetics, and in evolutionary biology, the structure of our theories sup-poses that we can separate out processes of heredity, development, and

selectionif not physically, then at least analytically, each from the other,

in our models of the evolutionary process. But these three dimensions

of the evolutionary process are inextricably fused and confounded in the


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process of cultural evolution. This happens because of the pivotal role

development and the life cycle assume in cultural transmission. I believe

that it will be shown with increasing clarity that we cannot do important

parts of evolutionary biology either without recognising a central role for

development there. (That is largely a work for the future, even though wehave made some excellent beginnings.) But for culture, we cannot even get

started.

Wimsatt contends that the indistinguishability of the stages in cultural re-

productive cycles necessitates a developmental systems approach to cultural

evolution.

This objection to memetic approaches to cultural selection theory hinges

on the opacity of the memeenvironment relationship, which touches on

the themes of environmental interaction, vertical versus horizontal transmis-

sion, and the memotypephemotype distinctionall three of which have been

central to the debates considered throughout this paper. As Hull ([2000],

pp. 567) writes:

The distinction between Lamarckian and non-Lamarckian inheritance

turns on the genotype-phenotype distinction. One reason why concep-

tual change tends to look deceptively Lamarckian is that this distinction

is not easy to make in memetic change. Memes play the role of genes in

replication, but what counts as environmental interaction [?] [. . .] If genesdetermine which transmissions are vertical in traditional gene-based se-

lection, then memes must determine which transmissions are vertical in

meme-based selection. The reason that this conclusion is unsettling is that

we do not have a very clear idea of memetic environmental interaction.

While memetic replication seems clear enough, memetic environmental

interaction does not.

Like other controversies explored in this paper, this objection assumes an anal-

ogy between genetics and memetics that is stricter than necessary for culturalapplications of natural selection to succeed. Wimsatts position presupposes

that memetic approaches to cultural selection must have an analogue of the

genotypephenotype relationship; however, this assumption is unwarranted.

Hull ([2000]) argues that researchers have adhered too rigidly to the analogy

with biological evolution, which in turn has caused cultural selection theory

to suffer. Genetics, developmental biology, and epidemiology are but three

of a plethora of biological arenas in which the effects of natural selection

manifest. The frameworks provided by the models of any one of these fields

should not necessarily be tightly held onto while exploring putative cases of

natural selection in other media. Instead, all selection processes should be

taken as special cases of natural selection, each on a par with the others in the

development of a generalized theory of evolution by natural selection.


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Although, as Hull points out, a strict analogy between traditional evolu-

tionary biology and cultural selection theory is not necessary, Hull ([2000],

pp. 456) also contends that the disanalogies between the two are overstated:

Given an incredibly simplistic notion of genes, memes are not in the least

like genes. The genetics that was incorporated into population genetics was

Mendelian genetics. Mendelian genetics, so the critics claim, is particulate

and concerns only pairs of alleles at a single locus (For a response, see

Wilkins [1998a], [1998b]). In contrast, memes are not all that particulate,

and more than two alternative memes can compete with each other. Of

course, one does not need to know very much Mendelian genetics to know

that Mendelian genes are not all that particulate and that numerous alter-

natives to Mendelian diploid inheritance exist (Crow [1979], [1999]). One

problem with interdisciplinary work is that any one worker is likely to know

much more about one area than any of the others. Geneticists know much

more about the complexities of genetics than of social groups. Conversely,

anthropologists and sociologists tend to be well versed in the details of

social groups. To them genetics looks pretty simple. Contrary to what we

were all taught in high school, genes are nothing like beads on a string. So

both memes and genes are likely to have comparably complex structures.

According to Hull, disanalogies between genetics and memetics are often

based on the misunderstanding that the subject matter of genetics is relatively

simple and that its concepts are clearly defined.Mendelian genes were initially largely defined operationally by positing the

number and kind of genes that could produce the observed patterns of inher-

itance. This operational definition was not employed with much consistency.

As Hull ([2000], p. 47) argues:

Of course, lots of genetic material could not be subdivided into dis-

tinct genes by Mendelian mechanisms, but it was still genetic material

and might one day be revealed by other mechanisms. Even limiting one-

self to Mendelian breeding experiments, additional genetic units werediscoveredmutons, codons, cistrons, and operons (Wilkins [1998a]).

With the advent of molecular biology, even more gene concepts were

introducedstructural genes, regulatory genes, introns, exons, nucleotides,

junk DNA, you name it. Mendelian geneticists complained that by terming

all these molecularly defined entities genes, molecular biologists were de-

stroying the clarity of the Mendelian gene. Of course, they did not mention

that they themselves had already destroyed much of its clarity.

Various sub-fields in evolutionary biology uniquely modified the gene concept

for the model systems they study.

It was Williams selectionist conception of a gene as any hereditary infor-

mation for which there is a favorable or unfavorable selection bias equal to

several or many times its rate of endogenous change (Williams [1966], p. 25)


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that Dawkins extended to his notion of replicators and ultimately to memes.

But, as Hull ([2000], p. 48) observes:

[. . .] why wait until this definition is extrapolated to memes to raise [. . .]

objections to it? Williams definition of evolutionary genes is just as difficultto apply as is its memetic correlate. In general, critics of memetics assume

standards so high for scientific knowledge that few, if any, areas of science

can possibly meet them.

Hull advocates ignoring in-principle objections to memetics, whether based

on disanalogies between cultural and biological instances of natural selection,

definitions of key concepts, or other higher-level theoretical debates. Rather,

memeticists should move from general discussions to focusing on empirical

investigations, as Hull ([2000], p. 49) argues:

Crude empirical investigations lead you to develop your theoretical per-

spective more clearly and extensively, and as it improves, you are in a better

position to run more sophisticated empirical investigations [. . .] memeti-

cists cannot begin to understand what the science of memetics is until they

generate some general beliefs about conceptual change and try to test them

[. . .] I want to urge memeticists to ignore the in-principle objections that

have been raised to memetics no matter how cogent they may turn out

to be and proceed to develop their theory in the context of attempts to

test it.

Hull defends memetics on the grounds that it is still a young discipline and

expects that the necessary developments will take place as the field matures.

Wimsatts objection is based on the difficulty of characterizing environmen-

tal interaction in cultural systems. In biological selection processes, traits are

identified by the fact that the proportion of the population in which they are

found in successive generations is dependent upon some common environ-

mental factor. Unless memetic traits are transmitted differentially to the nextcultural generation due to a process of environmental interaction, the evolution

of the system is merely a product of random drift rather than selection. What

cultural selectionists, then, must be primarily concerned with is generating no

mere description of cultural change, but an explanation of cultural traits as

adaptations to particular environmental circumstances.

8 Conclusion

One theme that dominates objections to cultural selection theory is the identi-

fication of disanalogies between cultural and biological systems. Some of these

objections present serious challenges to the fieldchallenges that must be ad-

dressed from within cultural selection theory. These challenges indicate genuine


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differences between biological and cultural systems, and they are based on aux-

iliary assumptions that are fairly well established in evolutionary biology: that

is, there is a consensus in theoretical and empirical biology that the identi-

fied biological disanalogue is essential to evolutionary biology and to natural

selection in general.

This paper has examined several objections in this category. One such ob-

jection is based on the absence of a mechanism guaranteeing the high-fidelity

replication of cultural units. Another is based on the absence of a common

hereditary architecture in cultural systems analogous to the structure of DNA.

A further such objection is based on the absence of clear phases in the cultural

units life cycle: that the memotypephemotype distinction does not character-

ize all cultural selection processes, and that the interaction between culture and

environment is not well defined.I have argued that these objections, significant though they may be, are not

fatal to the field but rather indicate areas that require further clarification.

With respect to objections where the consensus in biological and evolutionary

theory is potentially misled, it is essential that cultural selection produces case

studies that can clarify the relevant distinctions. For example, in response to the

objection that cultural selection fails because it lacks high-fidelity replication,

I have argued that consensus underlying the replicatorinteractor distinction

requires re-examination. I expect that cultural selectionand in particular, case

studies clarifying the role of environmental interactioncan produce examples

that will inform the requisite reformulation of the units of selection.

Other criticisms of cultural selection theory present less serious challenges

to the field. Like the challenges previously considered, these are also based

on disanalogies between biological and cultural systems; however, either the

articulated disanalogies are illusory, or they are not relevant to the applica-

tion of selection theory. The objection that cultural evolutionary systems are

inherently Lamarckian is one such example. I have argued that the failure of

these criticisms is attributable to two kinds of misconceptions: those aboutevolutionary biology and those about evolution by natural selection.

The absence of theoretical and definitional solidarity amongst proponents

of cultural evolution is likely to persist until more empirical investigations are

conducted. As Edmonds ([2002]) contends, such investigat

Reconsidering Cultural Selection Theory

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