Replication and Reproduction

First published Wed Dec 5, 2001; substantive revision Thu Oct 27, 2022

The problem of replication and reproduction arises out of the historyof genetics [see the entrygene for a historical review]. It is tied to the concept of the gene andits generalization in an evolutionary context [see the entryevolution]. Richard Dawkins introduced the notion of replicators—thingsthat self-replicate—as a universalization of evolutionaryunderstandings of genes. Dawkins argued that replicators are thesine qua non of evolution by natural selection [see the entrynatural selection], while other accounts only requirereproduction as one of itsdefining features. What exactly is a replicator? How are replicatorsdifferent from genes? Can evolution by natural selection occur withoutthe existence of replicators? Besides the biological domain, are thereany other domains in which replicators have been postulated? To answerthese questions, we will first provide some background forDawkins’ notion of the replicator and its ties with the conceptsof the gene and information. We will then introduce the distinctionbetweenreplicators andvehicles in the context ofbiological evolution, followed by the extension of this to otherdomains. Finally, we will discuss some of the challenges to the ideathat replicators are necessary conditions for evolution by naturalselection.

1. Background

The notion of replication arose as a concern in the period immediatelyafter the Second World War (seeFigure 1), with the rise of information theory, communication theory,cybernetics, and Erwin Schrödinger’s 1944 bookWhat isLife? Biologists immediately began to consider theinformational aspect of life, in each of these theoretical frameworks,in areas ranging from neurology to molecular biology, but especiallythe latter (Brookes 1956; Eigen & de Maeyer 1966; Gatlin 1972;Johnson 1970; Linschitz 1953; Margalef 1957 [1958]; Quastler 1953;Rashevsky 1955). It was only after the development of communicationstheory by Shannon (1948) and cybernetics by Wiener (1948) afterit that inheritance in biology took on a specificallyinformational tint (Griffiths & Stotz 2007). This is the case inparticular of the work of Watson and Crick on the double helixstructure of DNA (Watson & Crick 1953), as well as Crick’sCentral Dogma, which states that information cannot flow from proteinto DNA. The Central Dogma (Crick 1958) was initially cast in terms ofstructural information (Šustar 2007;Figure 2) and later recast by Crick as a more cybernetic or computational kind ofinformation (Crick 1970). [See Eugene V. Koonin 2012 & 2015 fordiscussion of the nature of the Central Dogma in current biology.]

a graph with a y axis ranging from 0% to 0.000160% and an x axis from 1800 to 2000; line stays below 0.000020% until 1960 at which point it rapidly rises to hit 0.000130% at 2000

Figure 1. Yearly frequency of count forthe word “replication” in Google text’s Englishcorpus from 1800–2000 with smoothing = 3. Data obtained fromGoogle Ngram Viewer (seeOther Internet Resources).

Structural information is effectively based on the transfer ofmolecular primary sequences (linear sequence) in the synthesis ofpolypeptides; that is, it is the literal sequence of basic groups(amino acids), before the protein folds into secondary and higherconformations. This is a physical and causal account of information:the new molecule has its sequence because it was templated in some wayby the “information”-imparting molecule (in this case, DNAto proteins). Griffiths and Stotz (see below) call this, accordingly,“Crick information”.

However, in the late 1950s, after the structure of DNA had become morewidely understood, chemist Homer Jacobson proposed acybernetic model of reproduction (Jacobson 1958), going sofar as to make a literal model out of train sets and electroniccircuits. Later, von Neumann proposed a theory of self-reproducingautomata (Neumann 1966), which has become a parallel but influentialresearch program to the issue in biology (Mitchell 2009). Jacob (1973)cemented the logical description of heredity, but the ultimateclassical source of the notion that genes are information comes fromWilliams’ classic 1966 bookAdaptation and NaturalSelection, in which he referred to genes as “cyberneticabstractions” (1966: 33).

The cybernetic notion of reproduction (and, later, replication) restedon the notion that genes werecontrollers of phenotypes;later, this came to be identified (e.g., by Gatlin) as synonymous withthe “communication” conception of information that Shannonproposed. In biology, genes came to be seen as “programs”for phenotypes [senders] that passed themselves on to progeny[receivers], around 1961 (Peluffo 2015).

Williams’ book was one inspiration for Dawkins’Selfish Gene (1976), and much of what Dawkins proposed aboutreplicators is an extension of Williams’ views, in conjunctionwith those of William D. Hamilton (1964) and various ethologicalideas. This view is now known as the gene’s eye view (fordiscussions see Okasha 2006, Chapter 5; Ågren 2021).

Williams introduced the notion of a gene as a substrate neutralentity:

I use the term ‘gene’ to mean that which separates andrecombines with appreciable frequency, (1966: 20)

and he further defined anevolutionary gene (that is, onethat is selectively important):

[A] gene could be defined as any hereditary information for whichthere is a favorable or unfavorable selection bias equal to several ormany times its rate of endogenous change. (1966: 25)

His answer to the question of what a gene is, is that a gene isevolutionary just insofar as it is something heritable that is subjectto rates of selection that exceed the mutation rate. This is adefinition, not a discovery. If it is favorably or unfavorablyselected and heritable, it is an evolutionary gene. According to thisdefinition, items that are not made of DNA can be classed asevolutionary genes so long as they match Williams’ definition(Lu & Bourrat 2018).

a rectangle divided into three horizontal regions, the first region has a purple background and is titled 'Replication' and has a diagram with the word 'DNA' and an arrow going from 'DNA' and circling back to it; the second region has a yellow background and is titled 'Transcription' and the diagram continues with an arrow from 'DNA' in the previous region to the word 'RNA' in this region; the third region has a green background and is titled 'Translation' and the diagram continues with an arrow from 'RNA' in the previous region going into this region and ending.

Figure 2. The Central Dogma of geneticsas envisaged by Crick around 1965 (Thieffry & Sarkar 1998).

2. Genes and Information

The language of “codes” and “information”flows easily enough with respect to replication [see the entrybiological information]. Transcription, translation, punctuation, redundancy, synonymy,editing, proofreading, errors, repairing of errors, messages, copies,and information all sound natural enough. Yet, the literature dealingwith information is both extensive and factious. Several differentformal analyses of information can be found and very little agreementabout which analysis is best for which subjects. On one point, thesescholars tend to agree—cybernetic information andcommunication-theoretic information will not do for replication inbiological contexts. Some argue for semantic information(Godfrey-Smith 2000a; Sarkar 2000; Sterelny 2000b). The trouble isthat no widely accepted version of semantic information exists. Winnie(2000) distinguishes between Classical and Algorithmic InformationTheory and opts for a revised version of the Algorithmic Theory. But,once again, the problem is that no such formal analysis currentlyexists.

In the face of all this disagreement and unfinished business,biologists such as Maynard Smith (2000) maintain either that informalanalyses of “information” are good enough or that somefuture formal version of information theory will justify the sorts ofinferences that they make. The sense of “information” usedin the Central Dogma of molecular biology is more like a fit oftemplate, or the primary structure of the protein sequence compared tothe sequence of the DNA base pairs. Attempts have been made in what isnow known as bioinformatics to use Classical Information Theory(Shannon’s theory of communication) to extract functional andphylogenetic information (Brooks & Wiley 1988; Gatlin 1972;Maclaurin 1998; Rodrick Wallace & Wallace 1998), but they appearto have been unsuccessful in the main.

While the most likely conclusion is that no version of informationtheory as currently formulated can handle “information” asit functions in biology (Griffiths 2001), attempts have continued tobe made to formulate just such a version (Bergstrom & Rosvall2011; Sternberg 2008). However, this undercuts the motivation forappealing to information theory to elucidate genes in the firstinstance. Griffiths & Stotz (2013: 153) have argued for“Crick” information, which they define as “thespecification of the order of amino acids in a polypeptidechain”. This specificity definition is more chemical thanabstract [see the section below onDevelopmental Systems Theory].

The “informational gene” conception, as James R. Griesemer(2005) terms it, is a highly abstract concept of a process and theentities that comprise it. This is intentional, as it permitsDarwinian dynamics to be applied to a range of phenomena, such associocultural, cybernetic, and medical processes. However, this raisesa metaphysical concern, namely that the view that a gene is nothingbut information is a version of Aristotelian hylomorphism [see theentryform versus matter]. That is, information is a type of form, which can be used to explainphysical processes. Classical hylomorphism was roundly demolished as ascientific hypothesis when Daltonian elements were named andinvestigated in the nineteenth century. By 1900, terms like“substance” (for matter that is propertyless apart frommass and extension in space) and “form” had taken on alargely philosophical sense that differed extensively fromAristotle’s own physical views. Griesemer notes:

We know that the informational genes are tied to matter and structure,but if evolutionary theory – to be general enough to covercultural and conceptual change – must be devoid of all referenceto concrete mechanism, it cannot follow from the theory, for example,that genes are inside organisms or are even parts of organisms, asDawkins’s language suggests. Strictly, only the correlationsbetween replicator and vehicle due to causal connections of acompletely unspecified sort can be implied by such a theory. Strivingto get matter and specific structure out of the theory in order tomake it apply to immaterial realms may thus leave it bankrupt as anaccount of causal connection for the material, biological cases.(2005: 79)

And yet, this kind of hylomorphism has remained, even in science.Biologists have argued that form determined many properties oforganisms in ways that could not be reduced to the actions of theirparts, and this kind of thinking remained and was co-opted by themolecular biologists and geneticists. Thus, we get the Central Dogmaof molecular biology:

The Central Dogma
This states that once ‘information’ has passed intoproteinit cannot get out again. In more detail, the transferof information from nucleic acid to nucleic acid, or from nucleic acidto protein may be possible, but transfer from protein to protein, orfrom protein to nucleic acid is impossible. Information means here theprecise determination of sequence, either of bases in thenucleic acid or of amino acid residues in the protein. (Crick1958)

Read physically, this means only that the structure of the DNAmolecule is not reproduced from the structure of proteins, a perfectlyreasonable account of the molecular processes. However, because Crickused the word “information”, some scientists, includingDawkins, took this to mean that genes are informational entities,which “code” for organismic traits from the molecularlevel up to the entire organism and even beyond. This is striking in apurple passage ofThe Blind Watchmaker, where Dawkinsdescribes genes as follows:

It is raining DNA outside. [downy seeds from willow trees] …The cotton wool is mostly made of cellulose, and it dwarfs the tinycapsule that contains the DNA, the genetic information. The DNAcontent must be a small proportion of the total, so why did I say thatit was raining DNA rather than cellulose? The answer is that it is theDNA that matters… whose coded characters spell out specificinstructions for building willow trees… It is raininginstructions out there, it’s raining programs; it’sraining tree-growing, fluff spreading, algorithms. That is not ametaphor, it is the plain truth. It couldn’t be any plainer ifit were raining floppy disks. (1986: 111)

So, we now turn to Dawkins’ view of genes asreplicators and the way in which he takes information to becybernetic and physical at the same time.

3. Dawkins’ View

Richard Dawkins introduced the distinction between replicators andvehicles inThe Selfish Gene (1976). For his purposes,Dawkins found the contrast between genes and organisms too restrictiveand specific. Everyone agrees that genes are replicators, but genesmay not be the only replicators. Dawkins also argued that perhaps moreinclusive entities than single genes might also function asreplicators. At the very least, the possibility should not be ignored.So, Dawkins adopted “replicator” as a more inclusive andgeneral term than “gene”. InThe ExtendedPhenotype, he defined a replicator as “anything in theuniverse of which copies are made” (Dawkins 1982b: 83). He alsointroduced the term “vehicle” for those entities producedby replicators that help these replicators increase in numbers byinteracting effectively with their environments. This distinction canbe expressed in terms of either entities or processes. According toDawkins, replicators function in replication, while vehicles functionin environmental interaction.

3.1 Genes as Replicators

A longstanding dispute in evolutionary biology concerns the levels atwhich selection can occur (Bourrat 2015b, 2021; Brandon 1996;Kerr & Godfrey-Smith 2002; Keller 1999; Lloyd 1988; Okasha 2006,2016; Sober & Wilson 1998; Williams 1966) [see the entry onunits and levels of selection]. Some authors see this dispute as concerning the levels at whichreplication can take place. Other authors take the levels of selectiondispute to concern environmental interaction and insist thatenvironmental interaction can take place at a variety of levels, fromsingle genes, cells, and organisms to colonies, demes, and possiblyentire species. Organisms certainly interact with their environmentsin ways that bias the transmission of their genes, but then so doentities that are both less inclusive than entire organisms (e.g.,sperm cells) and more inclusive (e.g., beehives).

Dawkins argued that replication inbiological evolutionoccurs exclusively at the level of the genetic material. The term“replication” refers first and foremost to copying, andgenes are the self-replicating molecules of biology. Some critics(e.g., Lewontin 1991: 48) interpreted this to mean that a strand ofDNA placed on a glass slide might start replicating all on its own. Ofcourse, no biologist has ever held such a view. Genes do replicatethemselves, but only with the aid of highly complicated molecularmachinery. Too often, however, the importance of this machinery hasgone unnoticed. To be sure, when we make copies on a photocopier, weare interested in the texts, figures, or just scrawls that appear onthese sheets of paper. We are not interested in how the photocopierworks, even if it doesall the work.

In Dawkins’ early writings, replicators and vehicles playeddifferent but complementary and equally important roles in selection;however, as he honed his view of the evolutionary process, vehiclesbecame less and less fundamental. Initially, Dawkins was content todethrone the organism from its pride of place in biology. It is animportant focus of environmental interaction, but other entities, bothbelow and above the organismic level, can also function as vehicles.In later writings, however, Dawkins went even further to argue thatphenotypic traits are what really matter for evolution by naturalselection to occur and that they can be treated independently of theirbeing organized into vehicles. More than that, features such as spiderwebs should be viewed as part of a spider’s phenotype. Hence,Dawkins titled his second bookThe Extended Phenotype(Dawkins 1982b).

Dawkins never lost his fascination with vehicular adaptations, afascination that his critics denigrate as Panglossian adaptationism.He filled his books with adaptationist scenarios, some more firmlysupported by data than others; however, from the perspective ofthe structure of evolutionary theory, he held that replicators aremuch more important than vehicles. For example, Dawkins argued at somelength that adaptations are always for the good of replicators, notvehicles (Lloyd 1992) [see the entry onunits and levels of selection]. Vehiclesexhibit these adaptations, but ultimately alladaptations must beexplicable in terms of changes in genefrequencies. Thus, it comes as no surprise when Dawkins (1994: 617)proclaims that he “coined the term ‘vehicle’ not topraise it but to bury it”. As prevalent as organisms might be,as definite as the causal roles that they play in selection are,reference to them can and must be omitted from any perspicuouscharacterization of selection in the evolutionary process. AlthoughDawkins is far from a geneticdeterminist, he is certainly ageneticreductionist. Whether reductionism itself is good orbad is a moot question (Sarkar 1998).

According to Dawkins, replicators have three basicproperties—longevity,fecundity andcopy-fidelity. Longevity means longevity of the genetype in the form of copies through descent (Dawkins 1982b:84; Hull 1980), although the stability of genetokens isincluded in the definition inThe Selfish Gene (1976: 18). Nogene as a physical body lasts all that long. In mitosis, a gene loseshalf of its substance at each replication. What endures, he says, isnot the entity itself but the information incorporated in itsstructure. It is this information that is copied with such highfidelity. Mutations do occur, but at very low frequencies. Even so, insome organisms, mutation rates must be too high because mechanismshave evolved that search out and repair such errors. Therefore, thegenotype is an informational notion; information is equivalent toAristotelian form. The type is theform of the tokens, inDawkins’ view.

One source of variation in genes of sexual organisms supplementingmutation is crossover. Pairs of homologous chromosomes line up next toeach other at meiosis, crossover, and recombine. For stretches of DNAin which different alleles exist, the result can be a change ininformation. Quite obviously, the shorter the stretch of DNA involved,the less likely that crossover will occur and the message change.Dawkins appeals to such dismantling of entities to argue againstorganisms functioning as replicators. In sexual organisms, organismsthemselves are torn apart and repeatedly assembled each generation(Caporael 2003). If long stretches of DNA lack the necessary identityby descent to function as replicators, then sexual organisms certainlylack it. However, some other explanation has to be provided forasexual organisms because they pass on their overall structure largelyunchanged from generation to generation. For example, R. A. Fisher,inThe Genetical Theory of Natural Selection (Fisher1930) considered the entire genetic complement of asexual organisms tobe a single gene, and this view has been repeated from time to timesince. According to Dawkins, genes, and only genes, can function asreplicators in biological evolution. How large these genetic units aredepends on such things as the prevalence of sex, the frequency ofcrossover or lateral gene transfers, and the intensity ofselection.

If there were sex but no crossing-over, each chromosome would be areplicator, and we should speak of adaptations as being for the goodof the chromosome. If there is no sex we can, by the same token, treatthe entire genome of an asexual organism as a replicator. But theorganism itself is not a replicator. (Dawkins 1982a: 95)

Dawkins offered two reasons for organisms not being able to functionas replicators. The first is the one he uses to delineate evolutionarygenes. As is the case for long stretches of DNA, organisms are tooeasily and frequently broken up to be considered units of replication.A second reason is that they cannot pass on changes in theirstructure, although some phenotypic change may result in generationalchange. In fact,some epigenetic mechanisms have been shownto be passed on across generations (Jablonka & Raz 2009; see theentry oninheritance systems). The amount of DNA that counts as a replicator certainly varies; but,according to Dawkins, nothing more inclusive than the genetic materialfunctions as replicators in biological evolution.

InThe Selfish Gene, although Dawkins wanted his definitionof gene to be close to that of Williams’ (1966)“evolutionary gene”, it is not quite:

A gene is defined as any portion of chromosomal material whichpotentially lasts for enough generations to serve as a unit of naturalselection. (Dawkins 1976: 30)

Where Williams’ definition is substrate neutral, Dawkins’is explicitly chromosomal and DNA-based. Where William’s“gene” refers toany entity, which is the reasonwhy it is an informational notion, Dawkins’ gene, by contrast,is tied to DNA, a view later heavily criticized by Stent (1977),an influential molecular biologist at the time.

On Dawkins’ account, the limits of genes need not be absolutelysharp. Nor must all genes be of the same length. The greater theselection pressure, the smaller the gene. At the most fundamentallevel, selection takes place between alternative alleles residing atthe same locus.

As far as a gene is concerned, its alleles are its deadly rivals, butother genes are just a part of its environment, comparable totemperature, food, and predators, or companions. (Dawkins 1976:40)

Alleles cannot cooperate with each other, only compete. That iswhere the “selfish” in “selfish gene” comesin. According to Dawkins (1976: 95), the selfish gene is not just onephysical bit of DNA. It is “all replicas of a particular bit ofDNA, distributed throughout the world” (for a recent defense ofa similar approach, see Haig 2012). Hence, genes do not form classesof spatiotemporally unrelated individuals but trees. They must bereplicas. But, being a replica is not enough. The linear repetition ofthe “same gene” in the form of several hundred copies isquite common. These replicas, however, do not reside at the samelocus. Although identical in structure, these genes do notcompete with each other in the way that alleles at the same locus can.In the simplest and most basic sense, alleles compete with alternativealleles at the same locus. Any other sorts of competition andcooperation are merely extrapolations from this fundamental sense ofallelic competition. Although genes may cooperate with each other invery complicated ways in embryological development, in replicationthey can be treated as “separate and distinct”. Indevelopment, the effects of genes blend. In replication, replicatorsdo not blend.

3.2 Hull’s Interactors

Dawkins introduced the general notions of replicator and vehicle sothat selection need not be limited exclusively to gene-basedbiological evolution. However, as the preceding discussion indicates,his later revisions to his general theoretical outlook were influencedstrongly by the traditional perspective of genes and organisms. Genescontain the information necessary to produce organisms and theiradaptations. Genes “ride around” in and“guide” organisms. As Dawkins describes them, vehicles arerelatively discrete entities that “house” replicators andthat can be regarded as machines programmed to preserve and propagatethe replicators that ride inside them. Although these terms may beappropriate for the relations between genes and organisms, theyinterfere with a more general analysis of replication and selection.What really matters in selection is that entities at various levels oforganization interact with their environments in such a way that therelevant replicators increase in relative frequency. The actual causalchain that connects replicators and vehicles need not be limited todevelopment.

For example, Dawkins argues at some length that genes and only genescan function as replicators in biological evolution. He adds that“all adaptations are for the preservation of DNA; DNA itselfjust is” (Dawkins 1982a: 45). But, DNA itself exhibitsadaptations. Anyone who has spent much time examining the molecularstructure of DNA soon realizes that it is adapted to replicate. Inaddition, the proliferation of junk DNA, transposons, and meioticdrive are three examples in which the only phenotypes that matter arephenotypic characteristics of genes (Brandon 1996; Sterelny 1996:388). Dawkins’ characterizations of replicators, vehicles, andthe relations between the two are too closely tied to genes,organisms, and development. DNA can certainly replicate itself, but itcan also function as a “vehicle”, even though it cannotcode for, ride around in, or direct itself. In sum, a more generalcharacterization of selection is needed, a characterization that doesnot assume that the only causal connection between replicators andvehicles is development from embryo to maturity. Such a generalizationwas offered by Hull (1980, 1981, 1988), who proposed that the relevantnotion is “interactor” rather than “vehicle”,as interactors are causal and active while vehicles are passiveentities.

4. Other Examples of Replicators

Although Dawkins finds it desirable to replace genes with replicatorsin his general characterization of the evolutionary process, he saysvery little about this more general notion inThe SelfishGene. Only one chapter (11) is dedicated to “memetic”evolution. Instead, as we saw, he discusses the special case of genesas replicators. The primacy of the genetic perspective in thecharacterization of replicators is claimed by some (Griesemer 2000b)to be one of the chief weaknesses not only in Dawkins’discussions but also in the work of his successors [see the sectionbelow onReproducers]. Hull and his colleagues (Hull, Langman, & Glenn 2001) claim to beproviding a general notion of replication, adequate to handle alldifferent sorts of replication, when too often we are simply readingpeculiarities of genetic replication into our general analysis ofreplication.

4.1 The Immune System

Does replication play the same role in other sorts of selection thatit plays in gene-based biological evolution? The reaction of theimmune system to antigens is closest to the standard biologicalexample of replication and selection (Hull et al. 2001). Certainly,immune systems arose through the same process as other functionalsystems. The basic structure of the human immune system is“built into our genes”, and the only genes that count inevolution are the ones that we received from our ancestors and cansubsequently pass on to our progeny. In Dawkins’ (1982b: 83)terminology, the genes that code for our immune system are active,germ-line replicators. They are active because they influence theirprobability to be copied; they are germ-line because they canpotentially have an infinity of descendants.

However, the genes that function in the reaction of the immune systemto antigens have two peculiarities. First, they incorporate mechanismsdesigned to produce very high frequencies of mutation; second, none ofthe genes involved in the functioning of the immune system aregerm-line. For instance, the genes that give rise to B-cells aredesigned to mutate extensively until one of these cells identifies aninvader as foreign. It then proliferates extensively as it attacks theinvader. As an organism matures, it accumulates more and more of theB-cells that have been successful in its past. More than this, asthe process of proliferation continues, the strength of the affinitiesto binding sites increases. Initially, the primary antibodies almostalways exhibit a weak affinity for their targets; however, as thereaction to the antigen continues, these affinities becomestronger.

Within the confines of a single organism, the reaction of the immunesystem to antigens has all the characteristics of selection processes;however, when the organism dies, all of these adaptations are lost. Insome species, females pass on not only the genes for the basicstructure of their immune systems but also some of the machinery thatpast invasions of antigens have produced in them. However, these cellsare rapidly removed from the offspring as it develops its own immunesystem. The reaction of the immune system to antigens departs fromgene-based selection in biological evolution in two ways. First, fromthe organismal perspective, the genes that function in protecting anorganism from invaders are not germ-line. They are somatic. Second,the relevant mutation rates are much, much higher in the immune systemthan in ordinary gene-based selection. Instead of mechanisms existingto discover mutations and repair them, mechanisms exist that encouragemutations—massively so. If the functioning of the immune systemis to count as an adaptive process, some changes must be made.One possibility is to clarify what counts as a “germ-line”replicator and reject the notion that extremely low mutationrates are inherent in all selection processes [see the section belowonChallenges to the Replicator].

4.2 Sociocultural Evolution

The temptation to regard cultures and societies as evolving entitiesin a way analogous to organisms has a long tradition. InTheSelfish Gene (Chapter 11), Dawkins proposes a version of it thatmakes use of the concept of the replicator. He chose the word“meme” for a sociocultural replicator due to itssimilarities with the word gene:

We need a name for the new replicator, a noun that conveys the idea ofa unit of cultural transmission, or a unit of imitation.‘Mimeme’ comes from a suitable Greek root, but I want amonosyllable that sounds a bit like ‘gene’ (Dawkins 1976:192).

The basic idea underlying Dawkins’ proposal is that sinceevolution by natural selection in the biological domain requiresreplication, it must also be the case in the sociocultural domain, andthat imitation, at least in many cases, is the way through whichsociocultural entities, the memes, replicate.

Important differences exist between sociocultural evolution andbiological evolution (Claidière & André 2012) [seethe entry oncultural evolution]. For example, Dawkins (1976) admits that he is on shaky ground when itcomes to the high-fidelity copying required of replicators. Memes seemto get changed much more frequently than genes (see Sterelny (2006)for a review). In general, however, it is fair to say that thestandards used to evaluate sociocultural transmission are not on parwith those used to evaluate biological transmission. An overlyidealized view of Mendelian genetics is contrasted with a much morerealistic view of cultural change. So the criticism goes, one problemwith memes is that they do not have discrete boundaries, do not allcome in the same size, and, in their functioning, are stronglyinfluenced by their environments. Genes, so the critics claim, havesharp boundaries, are all of the same size, and are immune toenvironmental influences. If sociocultural evolution is to beevaluated fairly, the same level of criticism must be applied to allputative examples of selection, from gene-based selection inbiological evolution and the reaction of the immune system to antigensto the development of the central nervous system and social learning(Hull et al. 2001). The collection of articles in Aunger (2000)present the arguments for and against a memetic approach toculture, and a more recent discussion is available in Lewens(2015).

Dawkins (1976) also places considerable emphasis on human brains asthe “vehicles” for memetic evolution. He defines“meme” as an entity capable of being transmitted from onebrain to another. Computers are also plausible vehicles for memes, aswell as artifacts more generally, which can “store” andthen release information when a copy of the artifact is made byemulation (mimesis): that is, without the presence of ademonstration. However, Dawkins’ discussion of memesis marred by the pervasiveness of the gene-organism perspective.For example, he defines “replicator” in terms oftransmission of information—memes leaping from brain to brain orfrom brains to computers and back again. But, memes do not leap frombrain to brain or from computer to computer. Their content istransmitted in a variety of ways, including books, audiotapes,conversations, and the like. As much as the physical basis changes,the message remains sufficiently unchanged. All instances of thismessage are equally memes, not just the ones residing in human brainsand computers.

Several accounts of memetic replication have been proposed in theliterature. Some consider that there is no replication in culturalevolution, but that memes are “attractor points” inculture (Atran 1998). Others consider that the interactor is the memeitself (Blackmore 1999) or that the meme is the selected culturalheritable information, just as Williams’ “evolutionarygene” is the selected genetic heritable information, and thememetic interactor is the repertoire of behaviors it elicits (Wilkins1998). One view that has been offered several times is that memes areactive neural structures (Aunger 1998, 2002; Delius 1991).

All the objections to the gene-meme analogy to one side, Dawkins(1976: 211) finds the chief difference between genetic and memeticchange is that biological evolution is at bottom a war between allelesresiding at the same locus: “Memes seem to have nothingequivalent to chromosomes and nothing equivalent to alleles”.First, the usual depiction of alleles residing at the same locus onhomologous chromosomes so central to Mendelian genetics is anoversimplification; but, and more importantly, for at least half ofthe history of life on earth, replication and selection took place inthe absence of chromosomes, meiosis, and the like. If gene-basedbiological evolution took place for so long in the absence of theMendelian apparatus and still does so in many extant organisms, thendemanding that memetic evolution proceeds by this very special andpossibly aberrant sort of inheritance seems too strong. The cost ofmeiosis remains a serious problem in ordinary biological evolution.Demanding that cultural evolution incorporates this same highlyproblematic element in its own makeup seems strange in the extreme. Ifwe are to develop a general analysis of selection, we mustdistinguish between essential and contingent features of thisprocess.

Numerous evolutionary biologists question how fundamental to selectionthe perspective of alleles at a locus actually is. Almost everyoneagrees that evolution involves changes in gene frequencies. However,few go on to add that evolution isnothing but changes ingene frequencies. When one looks at evolutionary biology, onediscovers that it involves much more than changes in gene frequencies.To be successful, memetic evolution must be fleshed out.

In 2000, Aunger summarized the disputes between proponents andskeptics of the memetic approach as revolving around threelevels:

whether culture is properly seen as composed of independentlytransmitted information units;
whether these so-called memes have the necessary qualifications toserve as replicators; and
whether a Darwinian or selectionist approach such as memetics isthe most feasible or desirable form for a science of culture to take.(Aunger 2000: 11)

However, over 20 years after the diagnosis, it seems that thememetic approach has, by and large, been abandoned, with some of itsinsights incorporated into other evolutionary approaches to culture.Perhaps the most successful is dual-inheritance theory (Richerson& Boyd 2008), also known as gene-culture coevolution. A verygeneral characterization of sociocultural evolution has been attemptedby El Mouden, André, Morin, and Nettle (2014). This approach,which relies on the most abstract formalism used by evolutionarytheorists—namely, the Price equation (for an introduction to thePrice equation see Okasha 2006: Chapter 1)—highlights someof the differences between biological and sociocultural evolution, andwhy a pure memetic approach seems difficult to implement as a generalframework.

4.3 The Extended Replicator

Some of Dawkins’ critics think that genes play too important arole in his notion of replication. Replication, so they argue, canoccur at other levels of organization as well. Just as Dawkinsextended the notion of the phenotype, these authors propose to makethe notion of replicators more general as well—to extend thereplicator so to speak. Dawkins introduced hisExtendedPhenotype conception for two reasons. First, he wanted to extendthe notion of a “phenotypic trait”. The sort of nest thata bird builds or the song that it sings can count as phenotypic traitsjust as much as the shape of its bill. Second, as mentioned earlier,Dawkins wanted to break the hold that organisms have over how weconceptualize the living world. Traits do tend to come bundled intoreasonably discrete entities; but, for making inferences about theevolutionary process, traits can be treated as separate and distinct.However, as he proceeded, Dawkins eventually decided that extendingthe phenotype was not enough. He had to bury it. All sorts offascinating mechanisms to one side, what really matters in selectiontakes place at the level of replicators.

Although such critics of Dawkins as Sterelny, Smith, and Dickison(1996) declined to join with Dawkins in his demotion of environmentalinteraction, they did agree with him that replicators are special.They play a special role in development. However, they do not limitreplicators to genes, even in biological evolution. Sterelny et al.propose to extend the replicator to include nonstandard entities:“the set of developmental resources that are adapted from thetransmission of similarity across the generations” (Sterelny2001: 338). For example, the sort of burrow that a particular organismdigs is influenced by its genetic makeup; however, if these burrowsare used over and over again, characteristics of these burrows canthemselves be viewed as replicators. The effects of these burrows getpassed on from generation to generation, but not via the DNA.

As more and more non-DNA replicators, including epigeneticinheritance, are acknowledged, Dawkins’ view began to grade intothe conception of the Extended Replicator, which Sterelny et al.(1996) specify as any non-genetic as well as genetic replicators where“genetic” refers to DNA. These two views are sogeneral that any case that can be described in one can be redescribedin the other. Differences lie in ease of description. According toSterelny et al. (1996), the burrows that some organisms dig canfunction as replicators—extended replicators—while Dawkinsportrays them as instances of extended phenotypes. The contrast isbetween selfish burrows and selfish genes for burrowing.

In the version of replication formulated by Sterelny et al. (1996),both copying and biofunction are crucial. Copying is quite obviously acausal phenomenon, but not any old causal connection will do.Similarity of copies is necessary but not sufficient. Thesecopies must be copies of copies. One copy must produce another copyand that copy produce still another copy and so on. For one entity tobe a copy of another, it must be the output of a process whosebiofunction is to conserve function. The function of copying is toproduce from one token another token, which is similar in the relevantrespects. Genes fit this definition, but so do lots of examples ofnon-genetic transmission, such as habitat stability resulting fromnest site imprinting, the song that a bird learns, and variousmicroorganism symbionts, not to mention socioculturaltransmission.

5. Challenges to the Replicator

5.1 Developmental Systems Theory

In the wake of Oyama’sThe Ontogeny of Information(1985, 2000) and other work, notably that of Daniel Lehrman (1953) andPatrick Bateson (2001), a view of biological evolution has arisen thatemphasizes development (see also Griffiths & Gray 1994a). In thenineteenth century, development was an extremely active researchprogram. The next great discoveries in biology were predicted to be inthe area of development; however, this was not to be. FirstMendelian genetics and then a version of evolutionary theory centeredon population genetics took over biology, and they did so whileavoiding development. Everyone knew that development was central toboth evolution and reproduction, but no one could see how to integratethe masses of developmental data available into the emerging synthesisof evolutionary biology and genetics. As a result, development wasleft out of the Modern Synthesis (Beurton, Falk, & Rheinberger2000; Gilbert 1991). Considering how central development actually isin metazoan and metaphyte biology, the advances made while ignoring itare staggering.

Even so, developmental biology continued on its course until, at longlast, we seem to understand development well enough to beginintegrating it into the rest of biology. On the most conservativeview, current versions of evolutionary theory can remain largelyunchanged as development is grafted onto them. On a second view, bothperspectives are likely to require some modification to achieve thisintegration. Our understanding of development may have to be modified,but so too for evolutionary theory (Pigliucci & Müller 2010).Finally, at the other extreme, development will all but replaceevolutionary theory. In their more exuberant moments, some advocatesof Developmental Systems Theory (DST) seem to be claiming just that.Just as some molecular biologists think that molecular biology israpidly replacing all the rest of biology, some advocates of DST arguethat developmental theories will simply replace current versions ofevolutionary theory.

On some versions of the DST view, genes have no uniquely privilegedrole in repeated cycles of development (Griffiths 2001; Griffiths& Gray 1994b). In fact, no element of the developmental matrixplays any privileged causal role—not genes, not organisms, notthe environment, not anything. Everything counts as a resource;however, in particular situations, certain resources will playmore important roles than other resources. In rejecting any privilegedrole for genes, advocates of DST are especially skeptical of oneparticular role supposedly played by genes—the transmission ofinformation. According to some, information is central todevelopmentalism, but genes are not the only mechanisms forinformation transfer. According to others, information plays no rolein the emerging developmentalist perspective. The developmental systemas a whole is the unit of selection [see the entry onunits and levels of selection]. If genes have no special role and whole developmental cycles arereplicators, as some developmental systems theorists argue, thedistinction between replicator and interactor becomes blurred.

In the continuing debate over developmental versus traditionaltheories of evolutionary biology, Sterelny et al. (1996) hold a fairlyconservative position. They agree with the developmentalists thatgenes play no privileged role in the development of phenotypes fromgenotypes. Genes do play a role in this process, simply not aprivileged role. Genes can serve as a causal bridge fromphenotype to phenotype, but other entities can do so as well. Genesare not the only replicators in biological evolution. The repeatedcycles in inheritance include many different sorts of constancies andrepetitions—genes, cellular machinery, phenotypic traitsincluding behaviors, and social structures. Information remainscentral to selection processes, but genes are not the only carriers ofsuch information. Genes predict phenotypic characters only in the samesense that environmental factors predict them. Sterelny (2000a) hasargued that developmental considerations require no fundamentalreevaluation of evolutionary conceptions along the lines of DST.

5.2 Evolution by Natural Selection without Replication

Godfrey-Smith (2009; see also Godfrey-Smith 2000b; Griesemer 2005)contrasts the replicator/interactor approach to natural selection witha more classical approach. This latter approach is usually given as a“recipe” for evolution by natural selection, to useGodfrey-Smith’s terminology (see also Brandon 1990: 3.1). Eversince Darwin, a number of authors have proposed that for evolution bynatural selection to occur, three conditions or ingredients arenecessary. A population should exhibit (1) variation that (2)leads to differences in fitness between the entities forming thepopulation [see the entry onfitness], (3) which are heritable (reviewed in Godfrey-Smith 2007). The mostfamous version of these recipes is that provided by Lewontin(1970).

There is a difference in emphasis between the two approaches in thatthe replicator/interactor approach focuses on theentitiesforming the population, while the classical approach provides apopulation description of evolutionary change. Another differencebetween the two approaches is that under the classical approach,replication is not required for evolution by natural selection tooccur. In fact, as long as heritability is positive, and the two otherconditions are satisfied, evolution by natural selection can occur(but will not necessarily do so, cf. Godfrey-Smith 2009: 25). Anabstract definition of heritability (\(h^2\))—although by nomeans fully general (see Bourrat 2015a, 2022)—is that itrepresents the slope of the regression of average offspring characteron parent or mid-parent (in case of sexual species) character [see theentry onheritability]. Given that a positive heritability does not entail replication ofcharacter between parent and offspring, but is rather a specialcase in which heritability would be exactly 1 in the absence ofenvironmental variation, the replicator/interactor perspective issubsumed by the classical approach. Godfrey-Smith concurs:

From this point of view, we can see the replicator analysis as pickingout a special case of what is covered (or supposed to be covered) bythe classical view. (2009: 36; see also Godfrey-Smith 2000b)

To see why a lack of replication does not prevent evolution by naturalselection from occurring, suppose a population of entities exists withdifferent sizes, as presented inFigure 3(a), in which there is selection for larger entities. We assume that theentities reproduce asexually in discrete generations and that there isno environmental variation. At the parental generation, there are twotypes of individuals, with respective sizes of \(\frac{3}{4}\) and 1.Due to the selection for size, entities of size \(\frac{3}{4}\)produce 3 offspring while entities of size 1 produce 4 offspring.Suppose now that there is a positive heritability, as defined above,so that, on average, offspring resemble more their parent than theyresemble other entities of the parental generation, as displayed onFigure 3(a), but only slightly so (\(h^2 = 0.2\)). After one generation, althoughthere is no high-fidelity replication, the average size of thepopulation has changed due to the effect of natural selection.This simple example demonstrates quite straightforwardly thatreplication is not required for evolution by natural selection tooccur.

We said earlier that replication would imply a heritability of exactly1 in the absence of environmental variation. Although this is correct,the reverse is not true. In fact, take again a similar population asinFigure 3(a), but this time assume a different pattern of inheritance for the twotypes of entities, as displayed inFigure 3(b). With this pattern of inheritance, although there is no replication,entitieson average have the exact same charactervalue as their parent. In the case of the entities of size\(\frac{3}{4}\),some offspring have the exact same characteras their parent, but that is not required either, as is the case withthe entities of size 1, of which none of the offspring has the samecharacter as their parent.

One possible response against regarding the classical approach toevolution by natural selection as unconditionally superior to thereplicator/interactor approach is to argue that althoughevolution by natural selection can occur without replication, whenthis happens, the evolutionary change resulting is not apurecase of evolution by natural selection. Rather, it is evolution bynatural selection mixed with some other evolutionary processes. Arationale for this claim is that the fact that offspring do notresemble their parent introduces some variation into the population ofwhich the origin is not natural selection. In abstract terms, theintroduction of variation can be regarded as a form of mutation(Bourrat 2019). Under this interpretation, the classicalapproach, on the one hand, and the replicator/interactor approach, onthe other hand, simply emphasize different aspects of evolutionaryexplanations. The classical approach gives us evolutionaryexplanationsin the context of other evolutionary processesoccurring, while the replicator/interactor approach gives usevolutionary explanationsof what the effect of natural selectionwould be in the absence of any other evolutionary process acting inthe population—that is, in its pure form. Understood thatway, the two approaches do not necessarily need to be opposed.

Another problem with the idea of replication is that it is descriptiondependent. Described finely, two objects can never be identical. Thesame applies to parents and offspring. This, in itself, does notrepresent a challenge to the replicator/interactor view. However,and more problematically, a coarser grain of description mightlead parents and offspring to be regarded as copies of one anotherwhen a finer-grained description would show that they are not. Such acoarse-grained way of viewing evolutionary processes has been endorsedby some memeticists (see Boudry 2018; Dennett 2017 for explicitdefences). However, without a principled way to choose whichgrain(s) of description ought to be used for the study of a particularevolutionary setting, the claim that replication is fundamental toevolution by natural selection in the context of other evolutionaryprocesses occurring becomes arbitrary. This problem is a general onein evolutionary theory (see Bourrat, 2020), but is even moreproblematic in the context of cultural evolution where theline between genotype and phenotype is difficult to drawand multiple channels of transmission exist (Charbonneau& Bourrat 2021; Bourrat & Charbonneau2022).

5.3 Origins of Replicators

Another challenge to the replicator/interactor approach to evolutionby natural selection is to see replicators themselves as aproduct of natural selection rather than as a necessarystarting point. One premise of this argument is that some form ofnatural selection must have occurred on entities that initially werenot replicating, perhaps not even reproducing. This is recognized byDawkins, who notes in passing in the second chapter ofThe SelfishGene that “Darwin’s ‘survival of thefittest’ is really a special case of a more general law ofsurvival of the stable” (Dawkins 1976: 13) and that lifeoriginated from selection of molecules for stability.

a diagram with two parts, the first consists of a black circle labeled '0.75' with arrows pointing to three black circles below labeled respectively '0.8', '0.8', and '0.95', below this is a legend stating 'Average: 0.85'. The second part consists of a black circle labeled '1' with arrows pointing to four black circles below labeled respectively '0.8', '0.8', '1', and '1', below is a legend stating 'Average: 0.9'. below all is an equation

(a) Population in which there is no replication and a low heritability(\(h^2\)), yet evolution by natural selection occurs.

(b) Population in which there is no replication, yet heritability is1. Evolution by natural selection occurs.

Figure 3. Illustration of the idea thatevolution by natural selection can occur without replication.

The idea that selection can occur without there being replicators wasfirst introduced as “chemical selection” in line with theviews of Oparin, who proposed that coacervates (colloidal droplets)could encapsulate protobiological chemicals and self-replicate withoutgenes through mechanical fission, a view also independently proposedby Haldane (Oparin 1936 [1938]; Haldane 1929; see Kolb 2016).This was later extended by the work of Sidney Fox, who showed thatproteinoid spheres could mechanically reproduce stably (Hsu & Fox1976).

If modern biological systems evolved from molecules that did not havethe ability to reproduce, let alone replicate, one might ask: what isthe simplest case of evolution by natural selection? It is a case inwhich different types of entities have different persistences but donot produce any offspring. In this model, the types with the propertyof persisting longer represent an increasing frequency of the totalpopulation over time. One problem, though in a model of persistencealone, is that each time one entity goes out of existence, it is notreplaced by another entity; thus, over time, the population size tendstowards zero. Although this does not mean that evolution by naturalselection cannot occur in such populations, this places importantconstraints on the evolutionary dynamics of this population (Van Valen1989). Further, it is possible to regard the ability for a populationto maintain its size or increase its size as a primordial form ofadaptation. This idea casts some doubt on the view that paradigmaticcases of evolution by natural selection require apopulation of entities that reproduce, a view championed byGodfrey-Smith (2009). Several authors, following in Van Valen’sfootsteps, have recently argued that important cases of evolutionby natural selection must involve reproduction, and that differentialpersistence with or without differential growth can be sufficient(Bouchard 2008, 2011; Bourrat, 2014; Charbonneau 2014;Doolittle 2014; Doolittle and Inkpen 2018; for a review, seePapale 2021). Nonetheless, assuming a metapopulation, it is clearthat populations with the ability to maintain or increase their sizewould out-compete those that are unable to do so. Thus, the abilityfor a population to sustain its population size might be regarded as aprimordial form of adaptation without which other adaptations would bealmost impossible to evolve.

A next “stage” in the evolution from primordialentities unable to reproduce to modern biological systems might thushave been for entities to produce new offspring entities (or multiply)without the ability to pass on their properties reliably. In such asituation, the size of a population could be maintained. The mostextreme form of that would be to imagine a population of two types inwhich each type has the same probability to produce an offspring ofthe other types as its own type. With entities of the two types havingdifferent persistence, a modest form of evolution by natural selectionis still possible. Versions of this model have been proposed byWilkins, Stanyon, and Musgrave (2012), Earnshaw-Whyte (2012), andBourrat (2015a). Since there is, on average, no more resemblancebetween an offspring and its parent than any other entity of theparental generation, and evolution by natural selection is stillobserved in such a scenario, one might think that heritability is,strictly speaking, not a necessary condition for evolution by naturalselection.

Wilkins et al. (2012) argue that, over time, in a reactor with limitedincoming chemical species, hypercycles (as defined by Eigen et al.1991) of reactions will tend to include reaction types that are morestable and capture more source molecules and energy than variantreaction types, leading to the evolution by natural selection ofreplicator-like molecules as well as energetically efficient“metabolic” molecules. Recent work in biochemistry hasadded strength to that suggestion (Altstein 2015). Hence, given thatno replicator is required for selection (in this case, chemical),replicator molecules can be seen as theoutcome of selectiveevolution, not a necessary prior condition for it. In effect, theevolution of replicators is the first major transition of evolution,despite the view of Szathmáry and Maynard Smith thatreplicators needed to come first (Maynard Smith & Szathmáry1995; Szathmáry & Maynard Smith 1997). Bourrat(2014) further developed this idea into a series of agent-basedmodels. He showed that replication is an attractor in populationslacking heritability once small random mutations increasing ordecreasing the fidelity of character inheritance are introduced.

5.4 Reproducers

The works on the origins of replicators tie in with the notion ofthereproducer proposed by Griesemer (2000a,b, 2002, 2005;Wimsatt & Griesemer 2007), although it was published first, basedon a circulated manuscript by Griesemer, by Szathmáry andMaynard Smith (1997). A reproducer is an entity that develops and hasa material overlap between the “parent” and“progeny”. This causal relationship between parents andprogeny is called “progeneration”, the “increase innumerically distinct objects of a given kind”. Reproduction is acomposite process of development and progeneration, leading to objectsthat are themselves capable of development and progeneration. Thus,the reproducer concept extends the notion of a developmentalsystem.

Here, inheritance is asystem property, not a property ofparts, unless they are also reproducing systems, and isdevelopmentally acquired. Organisms do not arise already capable ofreproduction in the main, but must undergo orderly transitions beforethey may reproduce.

The relation between reproducers, and replicators and interactors,varies according to author. Griesemer treats replication as theterminal form of reproduction, once coding mechanisms have beenevolved, which is itself an evolved form of multiplication (Griesemer2000a: 76). Unlike Dawkins’ replicator concept, which Griesemerbelieves is based on a similarity relationship of“copying”, a reproducer requiresmaterial overlapof systems. However, replication arises when material overlap evolvesprogeneration, and progeneration evolves a codical inheritance system,asFigure 4 indicates.

title is 'Modes of Multiplication'. This is a tree diagram. At the top is a node labeled 'Multiplication' below and to the left is a node labeled 'Copying' with the line connecting labeled 'nonmaterial overlap (property transmission)'. Below and to the right is 'Progeneration' with the line connecting labeled 'material overlap (parts transfer)'. Below 'Progeneration' is 'Reproduction' with the line connecting labeled 'material overlap of mechanisms of development'. Below 'Reproduction' is 'Inheritance' with the line connecting labeled 'material overlap of evolved mechanisms of development'. Below 'Inheritance' is 'Replication' with the line connected labeled 'material overlap of evolving coding mechanisms of development'.

Figure 4. Wimsatt and Griesemer’sreproducer concept (Wimsatt & Griesemer 2007, 268).

Others, particularly Jablonka and Lamb (2005, 2007), think thatreproducers are distinct from replication and that someepigenetic inheritance systems are reproducers without beingreplicators. For them, a reproducer is the “target ofselection” [see the entry oninheritance systems].

Godfrey-Smith (2009: 83; see also Okasha 2006: 14) takes issue withGriesemer’s requirement of “material overlap” asbeing too narrow a definiendum of reproducers. He thinks thatreproducers need not contribute actual material to their progeny(where they template for sequences, as in the case of viruses).Griesemer believes Godfrey-Smith has over-interpreted the requirement;he is primarily concerned with the idea that parentscauseprogeny (Griesemer 2014). Charbonneau (2014) furthers the move made byGodfrey-Smith, arguing that the existence of parent-offspringlineages is unnecessary for evolution by natural selection tooccur.

An aspect of reproducerssensu lato that has fascinated someis that it does not imply a particular level of reproduction orprogeneration; indeed, Szathmáry and Maynard Smith considerthat the major transitions of evolution each involve an evolved modeof progeneration that is more inclusive or novel than what it evolvedfrom (Maynard Smith & Szathmáry 1995; Szathmáry& Maynard Smith 1997).

Godfrey-Smith also proposed a taxonomy of reproducers, dividing theminto simple, collective, and scaffolded (Godfrey-Smith2009: 87ff). Simple reproducers are

entities that can reproduce … using their own machinery, inconjunction with external sources of energy and raw materials.

He gives an example of a bacterial cell. Collective reproducers

… are reproducing entities with parts that themselves have thecapacity to reproduce, where the parts do so largely through their ownresources rather than the coordinated activity as a whole.

And, scaffolded reproducers

are entities which get reproduced as part of the reproduction of somelarger unit (a simple reproducer), or that are reproduced by someother entity.

Examples here includes viruses and chromosomes. Godfrey-Smith appliesthe scaffolded reproducer category to sociocultural evolution.

The notion of reproducers has also been applied to speciation andcladogenesis (A. Hamilton & Haber 2006). While it may apply topopulations, given that taxa do not share the kind of causal unityfound in populations and organisms, it is unlikely to apply to speciesor supraspecific taxa, and macroevolution in general.

Recently, Veigl, Suarez, and Stencel (2022) have extendedGriesemer’s reproducer. In lieu of “reproducer”, theypropose the notion of the “reconstitutor”, which is astructure that is “re-produced” atevery generation irrespectively of the causal origin of thereconstruction. While, as presented above, the idea thatmaterial overlap is a necessary condition for an entity tobe a reproducer and, consequently, to be an entity involvedin a process of evolution by natural selection seems overlyrestrictive, the notion proposed by Veigl et al. appears overlyliberal for at least two reasons. First, without lookingfor the causal connections between entities involved inevolution by natural selection across generations, that is, whatis transmitted, predicting an evolutionary trajectory at the level ofthese entities would be difficult. Second, the idea of thereconstitutor erases an important distinction made in evolutionarytheory between two forms of evolution: namely,transformational and variational evolution (see Lewontin 1983).These two forms of evolution are exemplified by change dueto mutation and changes due to selection, respectively. Thedistinction between mutation and selection, along with drift, isat the heart of population genetics and general formulations ofevolutionary theory such as the Price equation (e.g., Frank 2012, whoemphasize the distinction between “selection” and “transmission” astwo important and distinct evolutionary processes). To adoptthe reconstitutor as a fundamental evolutionary entity wouldrequire extremely strong arguments against the view that mutation andselection described at one level are fundamentally differentevolutionary processes or, at the very least, clear advantagesfor the notion of the reconstitutor over the reproducer.

6. Conclusion

The idea that evolution by natural selection requires the existence ofreplicators has been a very useful heuristic in evolutionarybiology, but also a way of motivating the application of anevolutionary thinking beyond the boundaries of evolutionary biology.We started by showing how the notion of gene can be abstracted awayfrom its physical basis and characterized in purely informationalterms. We then showed how Dawkins and Hull pursued this abstractionand attempted to free the gene from its biological substrate. Thismotivated responses both from proponents of the replicator concept,who furthered the abstraction, and also critics. As successful andinspiring as such an abstraction has been, it should not be reified:evolution by natural selection and adaptation can occur even in theabsence of replicators. High fidelity certainly facilitates adaptiveevolution, but this might be because replicators are themselves theproduct of evolution rather than necessary for it.

Bibliography

Ågren, Arvid J., 2021,The Gene’s-Eye View ofEvolution, Oxford, New York: Oxford University Press.
Altstein, Anatoly D., 2015, “The Progene Hypothesis: TheNucleoprotein World and How Life Began”,BiologyDirect, 10(1): 67. doi:10.1186/s13062-015-0096-z
Atran, Scott, 1998, “Folk Biology and the Anthropology ofScience: Cognitive Universals and the Cultural Particulars”,Behavioral and Brain Sciences, 21(4): 547–609.doi:10.1017/S0140525X98001277
Aunger, Robert, 1998, “The‘Core Meme’Meme”,Behavioral and Brain Sciences, 21(4):569–570. doi:10.1017/S0140525X98221273
–––, 2000,Darwinizing Culture: The Statusof Memetics as a Science, Oxford: Oxford University Press.doi:10.1093/acprof:oso/9780192632449.001.0001
–––, 2002,The Electric Meme: A New Theoryof How We Think, New York: Free Press.
Bateson, Patrick, 2001, “Behavioral Development andDarwinian Evolution”, in Oyama et al. 2001: 149–166.
Bergstrom, Carl and Martin Rosvall, 2011, “The TransmissionSense of Information”,Biology and Philosophy, 26(2):159–176. doi:10.1007/s10539-009-9180-z
Beurton, Peter, Raphael Falk, and Hans-Jörg Rheinberger(eds), 2000,The Concept of the Gene in Development and Evolution:Historical and Epistemological Perspectives, Cambridge: CambridgeUniversity Press. doi:10.1017/CBO9780511527296
Blackmore, Susan J., 1999,The Meme Machine, Oxford; NewYork: Oxford University Press.
Bouchard, Frédéric, 2008, “Causal Processes,Fitness, and the Differential Persistence of Lineages”,Philosophy of Science, 75(5): 560–570.doi:10.1086/594507
–––, 2011, “Darwinism Without Populations:A More Inclusive Understanding of the ‘Survival of theFittest’”,Studies in History and Philosophy ofScience Part C: Studies in History and Philosophy of Biological andBiomedical Sciences, 42(1): 106–114.doi:10.1016/j.shpsc.2010.11.002
Boudry, Maarten, 2018, “Replicate after Reading: On theExtraction and Evocation of Cultural Information”,Biology& Philosophy, 33(3): 27. doi:10.1007/s10539-018-9637-z
Bourrat, Pierrick, 2014, “From Survivors to Replicators:Evolution by Natural Selection Revisited”,Biology andPhilosophy, 29(4): 517–538.doi:10.1007/s10539-013-9383-1
–––, 2015a, “How to Read‘Heritability’ in the Recipe Approach to NaturalSelection”,The British Journal for the Philosophy ofScience, 66(4): 883–903. doi:10.1093/bjps/axu015
–––, 2015b, “Levels of Selection AreArtefacts of Different Fitness Temporal Measures”,Ratio, 28(1): 40–50. doi:10.1111/rati.12053
–––, 2019, “In What Sense Can There BeEvolution by Natural Selection without Perfect Inheritance?”,International Studies in the Philosophy of Science, 32(1):39–77. doi:10.1080/02698595.2019.1623485
–––, 2020, “Natural Selection and theReference Grain Problem”,Studies in History andPhilosophy of Science (Part A), 80: 1–8.doi:10.1016/j.shpsa.2019.03.003
–––, 2021,Facts, Conventions, and theLevels of Selection, Cambridge: Cambridge University Press.doi:10.1017/9781108885812
Bourrat, Pierrick, and Mathieu Charbonneau,2022, “Grains of Description in Biological and CulturalTransmission”,Journal of Cognition and Culture,22(3–4): 185–202. doi:10.1163/15685373-12340131
Brandon, Robert N., 1990,Adaptation and Environment,Princeton, NJ: Princeton University Press.
–––, 1996,Concepts and Methods inEvolutionary Biology, Cambridge: Cambridge University Press.
Brookes, Bertram C., 1956, “An Introduction to theMathematical Theory of Information”,The MathematicalGazette, 40(333): 170–180. doi:10.2307/3608805
Brooks, Daniel R. and Edward O. Wiley, 1988,Evolution asEntropy: Toward a Unified Theory of Biology, second edition,Chicago: University of Chicago Press.
Caporael, Linnda, 2003, “Repeated Assembly”, in StevenJ. Scher & Frederick Rauscher (eds.),EvolutionaryPsychology, New York: Springer US, pp. 71–89.doi:10.1007/978-1-4615-0267-8_4
Charbonneau, Mathieu, 2014, “Populations WithoutReproduction”,Philosophy of Science, 81(5):727–740. doi:10.1086/677203
Charbonneau, Mathieu, and Pierrick Bourrat,2021, “Fidelity and the Grain Problem in CulturalEvolution”,Synthese 199(3–4):5815–36. doi:10.1007/s11229-021-03047-1
Claidière, Nicolas and Jean-Baptiste André, 2012,“The Transmission of Genes and Culture: A QuestionableAnalogy”,Evolutionary Biology, 39(1): 12–24.doi:10.1007/s11692-011-9141-8
Crick, Francis H.C., 1958, “On Protein Synthesis”, inFrank K. Sanders (ed.),Symposia of the Society for ExperimentalBiology, Number XII: The Biological Replication ofMacromolecules, Cambridge; New York: Cambridge University Press,pp. 138–163. [Crick 1958 available online]
–––, 1970, “Central Dogma of MolecularBiology”,Nature, 227(5258): 561–3.doi:10.1038/227561a0
Dawkins, Richard, 1976,The Selfish Gene, New York:Oxford University Press.
–––, 1982a, “Replicators andVehicles”, inCurrent Problems in Sociobiology, King’sCollege Sociobiology Group (eds), Cambridge: Cambridge UniversityPress, pp. 45–64.
–––, 1982b,The Extended Phenotype: The Geneas the Unit of Selection, Oxford; San Francisco: Freeman.
–––, 1986,The Blind Watchmaker,Harlow: Longman Scientific and Technical.
–––, 1994, “Burying the Vehicle”,Behavioral and Brain Sciences, 17(4): 616–617.doi:10.1017/S0140525X00036207
Delius, Juan, 1991, “The Nature of Culture”, in MarionStamp Dawkins, Tim R. Halliday, & Richard Dawkins (eds.),TheTinbergen Legacy, London: Chapman and Hall.
Dennett, Daniel C., 2017,From Bacteria to Bach and Back: TheEvolution of Minds, New York: W. W. Norton & Company.
Doolittle, W. Ford, 2014, “Natural Selection ThroughSurvival Alone, and the Possibility of Gaia”,Biology andPhilosophy, 29: 415–423. doi:10.1007/s10539-013-9384-0
Earnshaw-Whyte, Eugene, 2012, “Increasingly Radical ClaimsAbout Heredity and Fitness”,Philosophy of Science,79(3): 396–412. doi:10.1086/666060
Eigen, Manfred and Leo de Maeyer, 1966, “Chemical Means ofInformation Storage and Readout in Biological Systems”,Naturwissenschaften, 53(3): 50–57.doi:10.1007/BF00594747
Eigen, Manfred, Christof K. Biebricher, Michael Gebinoga, andWilliam C. Gardiner, 1991, “The Hypercycle. Coupling of RNA andProtein Biosynthesis in the Infection Cycle of an RNABacteriophage”,Biochemistry, 30(46):11005–11018. doi:10.1021/bi00110a001
El Mouden, Claire, Jean-Baptiste André, Olivier Morin, andDaniel Nettle, 2014, “Cultural Transmission and the Evolution ofHuman Behaviour: A General Approach Based on the PriceEquation”,Journal of Evolutionary Biology, 27(2):231–241. doi:10.1111/jeb.12296
Fisher, Ronald Aylmer, 1930,The Genetical Theory of NaturalSelection, Oxford: Clarendon Press.
Frank, S. A., 2012, “Natural Selection. III. Selectionversus Transmission and the Levels of Selection”,Journal ofEvolutionary Biology, 25: 227–43.
Gatlin, Lila L., 1972,Information Theory and the LivingSystem, New York: Columbia University Press.
Gilbert, Scott F., 1991,A Conceptual History of ModernEmbryology, New York: Plenum Press.
Godfrey-Smith, Peter, 2000a, “Information, Arbitrariness,and Selection: Comments on Maynard Smith”,Philosophy ofScience, 67(2): 202–207. doi:10.1086/392770
–––, 2000b, “The Replicator inRetrospect”,Biology and Philosophy, 15(3):403–423. doi:10.1023/A:1006704301415
–––, 2007, “Conditions for Evolution byNatural Selection”,The Journal of Philosophy, 104(10):489–516. doi:10.5840/jphil2007104103
–––, 2009,Darwinian Populations and NaturalSelection, Oxford: Oxford University Press.doi:10.1093/acprof:osobl/9780199552047.001.0001
Godfrey-Smith, Peter and Benjamin Kerr, 2002, “Group Fitnessand Multi-Level Selection: Replies to Commentaries”,Biologyand Philosophy, 17(4): 539–549.doi:10.1023/A:1020583723772
Griesemer, James R., 2000a, “Development, Culture, and theUnits of Inheritance”,Philosophy of Science,67(Supplement. Proceedings of the 1998 Biennial Meetings of thePhilosophy of Science Association. Part II: Symposia Papers):S348–S368. doi:10.1086/392831
–––, 2000b, “The Units of EvolutionaryTransition”,Selection, 1(1–3): 67–80.
–––, 2002, “What Is ‘Epi’About Epigenetics?”Annals of the New York Academy ofSciences, 981(1): 97–110.doi:10.1111/j.1749-6632.2002.tb04914.x
–––, 2005, “The Informational Gene and theSubstantial Body: On the Generalization of Evolutionary Theory byAbstraction”, in Martin R. Jones & Nancy Cartwright (eds.),Idealization XII: Correcting the Model. Idealization andAbstraction in the Sciences, Amsterdam: Rodopi Publishers, pp.59–115.
–––, 2014, “Reproduction and ScaffoldedDevelopmental Processes: An Integrated EvolutionaryPerspective”, in Alessandro Minelli & Thomas Pradeu (eds.),Towards a Theory of Development, Oxford: Oxford UniversityPress, pp. 183–202.doi:10.1093/acprof:oso/9780199671427.003.0012
Griffiths, Paul E., 2001, “Genetic Information: A Metaphorin Search of a Theory”,Philosophy of Science, 68(3):394–412. doi:10.1086/392891
Griffiths, Paul E. and Russell D. Gray, 1994a,“Developmental Systems and Evolutionary Explanation”,Journal of Philosophy, 91(6): 277–304.doi:10.2307/2940982
–––, 1994b, “Replicators and Vehicles? orDevelopmental Systems”,Behavioral and Brain Sciences,17(4): 623–624. doi:10.1017/S0140525X00036281
Griffiths, Paul E. and Karola Stotz, 2007, “Gene”, inMichael Ruse & David Hull (eds.),Cambridge Companion toPhilosophy of Biology, Cambridge: Cambridge University Press, pp.85–102. doi:10.1017/CCOL9780521851282.005
–––, 2013,Genetics and Philosophy: AnIntroduction, Cambridge: Cambridge University Press.doi:10.1017/CBO9780511744082
Haig, David, 2012, “The Strategic Gene”,Biologyand Philosophy, 27(4): 461–479.doi:10.1007/s10539-012-9315-5
Haldane, John Burdon Sanderson, 1929, “Origin ofLife”,The Rationalist Annual, 148: 3–10.
Hamilton, Andrew and Matthew H. Haber, 2006, “Clades AreReproducers”,Biological Theory, 1(4): 381–391.doi:10.1162/biot.2006.1.4.381
Hamilton, William D., 1964, “The Genetical Evolution ofSocial Behaviour. II”,Journal of Theoretical Biology,7(1): 17–52. doi:10.1016/0022-5193(64)90039-6
Hsu, Laura L. and Sidney W. Fox, 1976, “Interactions BetweenDiverse Proteinoids and Microspheres in Simulation of PrimordialEvolution”,Biosystems, 8(2): 89–101.doi:10.1016/0303-2647(76)90012-5
Hull, David L., 1980, “Individuality and Selection”,Annual Review of Ecology and Systematics, 11: 311–332.doi:10.1146/annurev.es.11.110180.001523
–––, 1981, “Units of Evolution: AMetaphysical Essay”, in U.L. Jensen & Rom Harré(eds.),The Philosophy of Evolution, Brighton UK: HarvesterPress, pp. 23–44.
–––, 1988, “Interactors VersusVehicles”, in Henry C. Plotkin (ed.),The Role of Behaviorin Evolution, Cambridge, MA: MIT Press, pp. 19–50.
Hull, David L., Rodney E. Langman, and Sigrid S. Glenn, 2001,“A General Account of Selection: Biology, Immunology, andBehavior”,Behavioral and Brain Sciences, 24(3):511–28; discussion 528–73.doi:10.1017/S0140525X0156416X
Jablonka, Eva and Marion J. Lamb, 2005,Evolution in FourDimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation inthe History of Life, Cambridge, MA: MIT Press.
–––, 2007, “Précis ofEvolutionin Four Dimensions”,Behavioral and BrainSciences, 30(4): 353–365.doi:10.1017/S0140525X07002221
Jablonka, Eva and Gal Raz, 2009, “TransgenerationalEpigenetic Inheritance: Prevalence, Mechanisms, and Implications forthe Study of Heredity and Evolution”,Quarterly Review ofBiology, 84(2): 131–76. doi:10.1086/598822
Jacob, François, 1973,The Logic of Life: A History ofHeredity, New York: Pantheon Books.
Jacobson, Homer, 1958, “On Models of Reproduction”,American Scientist, 46(3): 255–284.
Johnson, Horton A., 1970, “Information Theory in BiologyAfter 18 Years”,Science, 168(3939): 1545–1550.doi:10.1126/science.168.3939.1545
Keller, Laurent, 1999,Levels of Selection in Evolution,Princeton, NJ: Princeton University Press.
Kerr, Benjamin and Peter Godfrey-Smith, 2002, “Individualistand Multi-Level Perspectives on Selection in StructuredPopulations”,Biology and Philosophy, 17(4):477–517, with commentaries, and replies by the authors:Godfrey-Smith and Kerr 2002. doi:10.1023/A:1020504900646
Kolb, Vera M., 2016, “Origins of Life: Chemical andPhilosophical Approaches”,Evolutionary Biology, 43(4):506–515. doi:10.1007/s11692-015-9361-4
Koonin, Eugene V., 2012, “Does the Central Dogma StillStand?”Biology Direct, 7: 27–27.doi:10.1186/1745-6150-7-27
–––, 2015, “Why the Central Dogma? On theNature of the Great Biological Exclusion Principle”,BiologyDirect, 10: 52. doi:10.1186/s13062-015-0084-3
Lehrman, Daniel S., 1953, “A Critique of KonradLorenz’s Theory of Instinctive Behavior”,TheQuarterly Review of Biology, 28(4): 337–363.doi:10.1086/399858
Lewens, Tim, 2015,Cultural Evolution: ConceptualChallenges, Oxford, UK: Oxford University Press.doi:10.1093/acprof:oso/9780199674183.001.0001
Lewontin, Richard C., 1970, “The Units of Selection”,Annual Review of Ecology and Systematics, 1: 1–18.doi:10.1146/annurev.es.01.110170.000245
–––, 1983, “The Organism as the Subjectand Object of Evolution”,Scientia, 77(18): 65.
–––, 1991,Biology as Ideology: The Doctrineof DNA, Toronto: Anansi Press.
Linschitz, H., 1953,Information Theory in Biology,Urbana, IL: University of Illinois Press.
Lloyd, Elisabeth Anne, 1988,The Structure and Confirmation ofEvolutionary Theory, (Contributions in Philosophy, no. 37), NewYork: Greenwood Press.
–––, 1992, “Unit of Selection”, inKeywords In Evolutionary Biology, Evelyn Fox Keller andElisabeth A. Lloyd (eds), Cambridge MA: Harvard University Press, pp.334–342.
Lu, Qiaoying and Pierrick Bourrat, 2018, “The EvolutionaryGene and the Extended Evolutionary Synthesis”,The BritishJournal for the Philosophy of Science, 69(5): 775–800.doi:10.1093/bjps/axx019
Maclaurin, James, 1998, “Reinventing Molecular Weismannism:Information in Evolution”,Biology and Philosophy,13(1): 37–59. doi:10.1023/A:1006573021270
Margalef, Ramón, 1957 [1958], “La teoría de lainformación en ecología”,Memorias de la RealAcademia de Ciencias y Artes de Barcelona, 23: 373–449.Translated as “Information Theory in Ecology”, by WendellHall inYearbook of the Society for General Systems Research,3: 36–71. [Margalef 1957 [1958] available online (English)]
Maynard Smith, John, 2000, “The Concept of Information inBiology”,Philosophy of Science, 67(2): 177–194.doi:10.1086/392768
Maynard Smith, John and Eörs Szathmáry, 1995,TheMajor Transitions in Evolution, Oxford, New York and Heidelberg:WH Freeman/Spektrum.
Mitchell, Sandra D., 2009,Unsimple Truths: Science,Complexity, and Policy, Chicago: University of ChicagoPress.
Neumann, John Von, 1966,Theory of Self-ReproducingAutomata, Arthur W. Burks (ed.), Urbana, IL: University ofIllinois Press.
Okasha, Samir, 2006,Evolution and the Levels ofSelection, Oxford: Clarendon.doi:10.1093/acprof:oso/9780199267972.001.0001
–––, 2016, “The Relation Between Kin andMultilevel Selection: An Approach Using Causal Graphs”,TheBritish Journal for the Philosophy of Science, 67(2):435–470. doi:10.1093/bjps/axu047
Oparin, Aleksandr Ivanovich, 1936 [1938],Vozniknovenie zhiznina zemle, Moscow: Izd. Akad. Nauk SSSR. Translated asTheOrigin of Life, Sergius Morgulis (trans.), New York:Macmillan.
Oyama, Susan, 1985,The Ontogeny of Information: DevelopmentalSystems and Evolution, Cambridge; New York: Cambridge UniversityPress.
–––, 2000,The Ontogeny of Information:Developmental Systems and Evolution, second edition, Durham, NC:Duke University Press.
Oyama, Susan, Paul E. Griffiths, & Russell D. Gray (eds.),2001,Cycles of Contingency: Developmental Systems andEvolution, Cambridge, MA and London: MIT Press: A Bradford Book.
Papale, François, 2021, “Evolution by Means ofNatural Selection without Reproduction: Revamping Lewontin’sAccount”,Synthese, 198(11): 10429–55.doi:10.1007/s11229-020-02729-6
Peluffo, Alexandre E., 2015, “The ‘GeneticProgram’: Behind the Genesis of an Influential Metaphor”,Genetics, 200(3): 685–696.doi:10.1534/genetics.115.178418
Pigliucci, Massimo and Gerd B. Müller, 2010,Evolution:The Extended Synthesis, Cambridge MA: MIT Press.
Quastler, Henry, 1953,Information Theory in Biology,Urbana IL: University of Illinois Press.
Rashevsky, N., 1955, “Life, Information Theory, andTopology”,The Bulletin of Mathematical Biophysics,17(3): 229–235. doi:10.1007/BF02477860
Richerson, Peter J. and Robert Boyd, 2008,Not by Genes Alone:How Culture Transformed Human Evolution, Chicago: University ofChicago Press.
Sarkar, Sahotra, 1998,Genetics and Reductionism,Cambridge; New York: Cambridge University Press.doi:10.1017/CBO9781139173216
–––, 2000, “Information in Genetics andDevelopmental Biology: Comments on Maynard Smith”,Philosophy of Science, 67(2): 208–213.doi:10.1086/392771
Schrödinger, Erwin, 1944,What Is Life? The PhysicalAspect of the Living Cell, Cambridge: Cambridge UniversityPress.
Shannon, Claude E., 1948, “A Mathematical Theory ofCommunication”,The Bell System Technical Journal,27(3): 379–423, 27(4): 623–656.doi:10.1002/j.1538-7305.1948.tb01338.x anddoi:10.1002/j.1538-7305.1948.tb00917.x
Sober, Elliott and David Sloan Wilson, 1998,Unto Others: TheEvolution and Psychology of Unselfish Behavior, Cambridge, MA:Harvard University Press.
Stent, Gunther S., 1977, “You Can Take the Ethics Out ofAltruism but You Can’t Take the Altruism Out of Ethics”,Hastings Center Report, 7(6): 33–36.doi:10.2307/3560881
Sterelny, Kim, 1996, “The Return of the Group”,Philosophy of Science, 63(4): 562–584.doi:10.1086/289977
–––, 2000a, “Development, Evolution, andAdaptation”,Philosophy of Science, 67(Supplement.Proceedings of the 1998 Biennial Meetings of the Philosophy of ScienceAssociation): S369–S387. doi:10.1086/392832
–––, 2000b, “The ‘GeneticProgram’ Program: A Commentary on Maynard Smith on Informationin Biology”,Philosophy of Science, 67(2):195–201. doi:10.1086/392769
–––, 2001, “Niche Construction,Developmental Systems, and the Extended Replicator”, in SusanOyama et al. 2001: 333–350.
–––, 2006, “Memes Revisited”,British Journal for the Philosophy of Science, 57(1):145–165. doi:10.1093/bjps/axi157
Sterelny, Kim, Kelly C. Smith, and Michael Dickison, 1996,“The Extended Replicator”,Biology andPhilosophy, 11(3): 377–403. doi:10.1007/BF00128788
Sternberg, Richard, 2008, “DNA Codes and Information: FormalStructures and Relational Causes”,Acta Biotheoretica,56(3): 205–232. doi:10.1007/s10441-008-9049-6
Szathmáry, Eörs and John Maynard Smith, 1997,“From Replicators to Reproducers: The First Major TransitionsLeading to Life”,Journal of Theoretical Biology,187(4): 555–571. doi:10.1006/jtbi.1996.0389
Šustar, Predrag, 2007, “Crick’s Notion ofGenetic Information and the ’Central Dogma’ of MolecularBiology”,British Journal for the Philosophy ofScience, 58(1): 13–24. doi:10.1093/bjps/axl018
Thieffry, Denis and Sahotra Sarkar, 1998, “Forty Years Underthe Central Dogma”,Trends in Biochemical Sciences,23(8): 312–316. doi:10.1016/S0968-0004(98)01244-4
Veigl, Sophie J., Javier Suárez, and Adrian Stencel, 2022,“Rethinking Hereditary Relations: The Reconstitutor as theEvolutionary Unit of Heredity”,Synthese, 200(5): 367.doi:10.1007/s11229-022-03810-y
Wallace, Rodrick and Robert G. Wallace, 1998, “InformationTheory, Scaling Laws and the Thermodynamics of Evolution”,Journal of Theoretical Biology, 192: 545–559.
Watson, James D. and Francis H.C. Crick, 1953, “MolecularStructure of Nucleic Acids: A Structure for Deoxyribose NucleicAcid”,Nature, 171(4356): 737–738.doi:10.1038/171737a0
Wiener, Norbert, 1948,Cybernetics, or, Control andCommunication in the Animal and the Machine, Cambridge MA:Technology Press.
Wilkins, John S., 1998, “What’s in a Meme? Reflectionsfrom the Perspective of the History and Philosophy of EvolutionaryBiology”,Journal of Memetics – Evolutionary Models ofInformation Transmission, 2(1): 2–33. [Wilkins 1998 available online]
Wilkins, John S., Clem Stanyon, and Ian Musgrave, 2012,“Selection Without Replicators: The Origin of Genes, and theReplicator/Interactor Distinction in Etiobiology”,Biologyand Philosophy, 27(2): 215–239.doi:10.1007/s10539-011-9298-7
Williams, George C., 1966,Adaptation and Natural Selection: ACritique of Some Current Evolutionary Thought, Princeton NJ:Princeton University Press.
Wimsatt, William C. and James R. Griesemer, 2007,“Reproducing Entrenchments to Scaffold Culture: The Central Roleof Development in Cultural Evolution”,Integrating Evolutionand Development: From Theory to Practice, Roger Sansom and RobertN. Brandon (eds), Cambridge, MA: MIT Press, pp. 227–324.
Winnie, John A., 2000, “Information and Structure inMolecular Biology: Comments on Maynard Smith”,Philosophy ofScience, 67(3): 517–526. doi:10.1086/392793

Academic Tools

How to cite this entry.
Preview the PDF version of this entry at theFriends of the SEP Society.
Look up topics and thinkers related to this entry at the Internet Philosophy Ontology Project (InPhO).
Enhanced bibliography for this entryatPhilPapers, with links to its database.

Other Internet Resources

Information on von Neumann’s self-reproducing automata.
Introduction to the molecular aspects of DNA replication
Google Ngram Viewer for replication (seeFigure 1 above, note there are some differences from the figure due to changes in thedatabase Ngram uses)

Acknowledgments

The original version of this article, entitled“Replication”, was written by the late David Hull. InJanuary 2005, John Wilkins became a co-author, with the responsibilityfor revising and updating the entry, so as to keep it current. Thus,revisions on or after January 2005 were introduced byWilkins and, from October 2018, by Wilkins and Pierrick Bourrat.Although the current version includes some of Hull’s originalessay, it does not represent his final views.

The authors would like to thank Peter Godfrey-Smith and JamesGriesemer for their suggestions and comments, and the reviewers whomade many useful comments and suggestions.

Open access to the SEP is made possible by a world-wide funding initiative.
The Encyclopedia Now Needs Your Support
Please Read How You Can Help Keep the Encyclopedia Free

Browse

About

Support SEP

Mirror Sites

View this site from another server:

USA (Main Site)Philosophy, Stanford University

Info about mirror sites

Library of Congress Catalog Data: ISSN 1095-5054

Movatterモバイル変換