Can Darwinism be “Generalized” and of what use would this be

J Evol Econ (2011) 21:545–562DOI 10.1007/s00191-011-0235-3

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Can Darwinism be “Generalized” and of what usewould this be?

Georgy S. Levit · Uwe Hossfeld · Ulrich Witt

Published online: 1 July 2011© Springer-Verlag 2011

Abstract It has been suggested that, by generalizing Darwinian principles, acommon foundation can be derived for all scientific disciplines dealing withevolutionary processes, especially for evolutionary economics. We show, how-ever, that in the development of evolutionary biology, the abstract principlesof so-called “Generalized Darwinism” have not been crucial for distinguishingDarwinian from non-Darwinian approaches and, hence, cannot be consideredgenuinely Darwinian. Moreover, we wonder what can be gained by invokingthe abstract principles of Generalized Darwinism given that they do not sufficeto substantiate an explanation of actual evolutionary processes. To that end,specific hypotheses are required. They neither follow from the suggestedabstract principles, nor are they more easily found on that basis. Accordingly,we find little evidence in the literature for the claim that generalized Darwinianprinciples enhance the explanatory power of an evolutionary approach toeconomics.

G. S. LevitUniversity of King’s College, Halifax, Canadae-mail: [email protected]

G. S. Levit · U. HossfeldAG Biodidactics, Friedrich Schiller University, Jena, Germany

U. Hossfelde-mail: [email protected]

U. Witt (B)Max Planck Institute of Economics, Jena, Germanye-mail: [email protected]

546 G.S. Levit et al.

Keywords Darwinism · Evolution · Evolutionary economics ·“Generalized Darwinism” · Variation · Selection · Retention

JEL Classification B25 · B40 · B52

1 Introduction

In reflecting on evolutionary phenomena in their respective domains, manydisciplines borrow conceptual tools from Darwinian evolutionary biology.Evolutionary economics is no exception. It is popular, for example, to con-struct analogies to natural selection, particularly in the neo-Schumpeterianliterature (see, e.g., Nelson and Winter 1982; Metcalfe 1998; Windrum 2007;Andersen 2009, chap 12). More recently, it has been argued, however, that it isnot only possible but also desirable to go beyond metaphors and analogies.It is claimed that general principles of evolution can be derived throughisolating abstraction from the Darwinian theory that explain alike evolutionin the biological domain and non-biological domains, e.g. that of economics(Hodgson and Knudsen 2006a, 2010). The claim that “Darwinism involves ageneral theory of all open, complex systems” (Hodgson 2002) ties in with anongoing debate in the philosophy of science originally triggered by Dawkin’s(1983) vision of “Universal Darwinism”. In that debate, variation, selection,and inheritance (for the latter also the terms “replication” or “retention” aresometimes used) have been postulated by some writers as domain-independentgeneral principles (see Campbell 1965; Dawkins 1983; Hallpike 1985, 1986;Wilkins 1998; Wimsatt 1999; Hull 2001; Crozier 2008). These principles alsoform the core of “Generalized Darwinism”, an approach propagated in aseries of works by Hodgson and Knudsen (2004, 2006a, b, 2010) with specialreference to evolutionary economics.

With their recent manifesto “In defense of Generalized Darwinism” in thisjournal (Aldrich et al. 2008), several prominent scholars have endorsed suchan encompassing approach to evolutionary economics. Its tacit presumptionis a fundamental homology between evolution in nature and evolution in theeconomy—“social evolution is Darwinian”, as Hodgson and Knudsen (2006a)have put it. In practice, the homology assumption provides a basis for adoptingstill other domain-specific, biological concepts in an abstract disguise. Forexample, the biological notions of genotype and phenotype are transformedinto an abstract “replicator–interactor” dichotomy which is then suggestedas a general analytic tool in evolutionary theories (Hull 1988; Hodgson andKnudsen 2004, 2006b).

The claims for Generalized Darwinism have stirred some opposition in evo-lutionary economics. Buenstorf (2006), for example, argues that the conceptof selection based on a replicator–interactor distinction contributes little tounderstanding industrial evolution: “The homology between biological inher-itance and the replication of routines is achieved at the price of abstractionfrom economically relevant details.” On the basis of his “ontological continuity

Can Darwinism be “Generalized” and of what use would this be? 547

hypothesis”, Witt (2003 chap 1; 2004, 2008) claims that the Darwinian theoryis relevant for understanding economic evolution, albeit not in the form ofabstract one-fits-all principles but rather as a meta-theory about how man-made evolution could emerge from, and is molded in, evolution in nature.The notable difference here is that, as a result of the intervention of humanintelligence, intentionality, creativity, and knowledge accumulation, the rulesof man-made evolution are likely to differ from those of biological evolution.Supporting this view, Cordes (2006) points to the fact that biological evolu-tion has created a human “behavioral repertoire” which forms the bedrockof cultural evolution and gives rise to specific adaptation dynamics for theunderstanding of which the abstract principles of variation, selection, andinheritance contribute little, if anything. Vromen (2008) notes that “Hodgsonand Knudsen do not counter this critique by dismissing the extra substancethat Witt’s continuity hypothesis adds. . . , but by diminishing the ontologicalsubstance of their own Generalized Darwinism. . . . The price they have to payfor this is that it leaves their Darwinian principles with virtually no content.”

With this background, the present paper sets out to do two things. First, itwill go beyond the criticism launched against Generalized Darwinism so far byshowing that its very core—the identification of Darwinism with the principlesof variation, selection, and inheritance—is highly problematic (an argumentthat can also be leveled against Dawkin’s Universal Darwinism). Conceptu-alizing evolution by means of these abstract principles not only comes at theprice of abstracting from economically relevant details, as Buenstorf (2006)notes, but also at the price of abstracting from crucial biological “details”. Inbiology, it depends on the specific features of the processes by which geneticinformation is varied and inherited whether different theories about varia-tion, selection, and inheritance qualify as Darwinian, non-Darwinian, or anti-Darwinian (see Levit et al. 2008a). These features require specific hypothesesand addenda (left out by the suggested abstraction) which ultimately decideabout the fit with the logic of Darwinism and the empirical evidence that hasbeen gathered for it. The three principles that remain as allegedly unifyingframework after the isolating abstraction has been carried out are but an emptyhull that, as Vromen has suggested, lacks explanatory substance.

Second, and in recognizing the history of evolutionary theorizing in biology,the paper will examine more closely the very idea inspiring Generalized Dar-winism. This is the claim that the abstract principles identified by GeneralizedDarwinism are helpful in, or even necessary for, building up an encompassingevolutionary theory in economics. To invoke abstract principles “top-down”to construct a theory of evolution, e.g., for the economic domain, means, ofcourse, that hypotheses on the “details” of the particular domain must befound that account for the specific conditions of, say, economic processes.However, such specific hypotheses can neither be logically derived from theabstract principles, nor do the abstract principles seem particularly useful oreven necessary for arriving at specific hypotheses by which explanations foreconomic evolution can be provided. Doubts may therefore be raised as towhether Generalized Darwinism offers any advantage in explaining economic


evolution. It seems worth noting in this context that evolutionary biology,the very discipline from which Generalized Darwinism borrows its principles,has never taken resort to the research strategy which the proponents ofGeneralized Darwinism suggest for evolutionary economics.

Accordingly, the plan of this paper is as follows. Sections 2 to 4 give anoverview of three different phases in the development of Darwinism. Eachof these phases is indicative of significant changes in the central tenets thatthe Darwinians were forced to accept in view of the problems emergingfrom the specificities discovered in the various interconnected research ac-tivities, reaching from the paleontological record to the molecular processesgoverning life and reproduction. In this process, the principles of variation,selection, and inheritance did play a role. But they were neither the onlygeneral principles considered relevant, nor did they serve as the criteriondistinguishing Darwinian from non-Darwinian interpretations. In the light ofthese insights, Section 5 turns to a discussion of the prospects and limitationsof the top-down approach suggested by the proponents of Generalized Dar-winism for the development of evolutionary economics. Section 6 presents theconclusions.

2 Darwin and “Classical Darwinism”

Darwinism is a dynamic and complex theoretical system consisting of severalnecessary, interconnected postulates and numerous theoretical addenda andspecific hypotheses. A difficulty that arises in defining Darwinism results fromthe fact that the theory of natural selection achieved its logical consistency andconceptual maturity only decades after Darwin’s death. For that reason, anappeal to Darwin’s own writings is not necessarily the best way to determinewhether a particular concept in question is, or is not, “Darwinian” in character.To account for this complication, it is necessary to distinguish different, histor-ically contingent, phases of “Darwinism” (and of anti-Darwinian concepts) inbiology. The majority of historians of biology agree that there are three majorphases in the growth of Darwinian thought (Reif 2000; Reif et al. 2000).

The first phase is Classical Darwinism, represented by Darwin’s The Originof Species by Means of Natural Selection (1859), subsequent works, and theirimmediate reception among biologists. Pushing the idea of organic evolutionand common descent, Darwin’s contemporaries grappled with how strictlyto interpret the relative importance of the principle of natural selection,the concept of inheritance combining both hard and soft (‘Lamarckian’)mechanisms, and the postulate of gradualism. In fact, Darwin had proposeda theoretical system which implied four evolutionary mechanisms: naturalselection, sexual selection, direct effects of the environment, and inheritanceof acquired characteristics. This pluralism of evolutionary mechanisms hadled by the end of the 19th century to a second phase that culminated in theestablishment of two major schools of Darwinism.


The neo-Darwinians, in particular August Weismann (1885), eliminatedLamarckian mechanisms and Darwin’s pangenesis, thus opening the avenueto a consequent selectionism. In details, however, Weismann’s approachwas hardly compatible with the mature version of Darwinism (i.e. the syn-thetic theory of evolution).1 The interpretation of such “details” of the var-ious principles and their relative importance also gave rise to alternative(anti-Darwinian) and rival theories of evolution. Incoherences within theDarwinian theories and unresolved puzzles left by classical Darwinism andneo-Darwinism even resulted in what became a temporary eclipse of selec-tionism in evolutionary biology (Bowler 1983).

Only with the third phase of Darwinian thought could the essentialdifficulties be overcome to a certain extent. This phase started with the break-through of the Synthetic Theory of Evolution in the late 1930s, the so-calledModern Synthesis prepared by the rise of Mendelism and population genetics.It was one more step to Darwinism as it is now: a logically coherent andempirically applicable research program. In this form, it is able to integrateclassical genetics, population genetics and molecular genetics, systematics,evolutionary morphology, developmental biology, palaeontology etc. on aselectionist basis while, at the same time, acknowledging the importance ofnon-selectionist factors of evolution, especially isolation, chance events, andpopulation size. This means that “. . . selection is regarded as important, butonly as one of several evolutionary factors” (Reif et al. 2000).

Integral to Classical Darwinism—the starting point of the longdevelopment—was Darwin’s and Wallace’s introduction of the “principle ofnatural selection” as a hypothetical mechanism of biological evolution (Darwinand Wallace 1858; Hossfeld and Olsson 2009). Darwin’s influential Germandisciple, Ernst Haeckel,2 argued that a scientific theory of transmutation(evolution) had already been formulated by Lamarck (a theory for whichHaeckel coined the very term “Lamarckism” as opposed to Cuvierism, atheory demanding constancy of species). According to Haeckel, this wasa general theory of descent aiming at a complete and harmonic pictureof evolution by reducing all its phenomena to “the only physiologicalprocess of nature, the transmutations of species” (Haeckel 1866, p. 167).The theory of natural selection, by contrast, reveals the exact machinery,the “mechanical causes” of transmutation (evolution), and explains itsdirectionality. Accordingly, Lamarck should be appreciated, Haeckel claimed,for promoting general evolutionism, while Darwin must be praised for hiscausal explanation of the theory of descent. Yet what Haeckel considered

1For example, Weismann (1893) claimed that the ultimate source of variation is always the effectof external influence. Rejecting Lamarckian mechanisms at the level of soma, he championed whatWinther (2001) called “inheritance of acquired germ-plasm variations”.2Since Haeckel’s name became a synonym for Darwinism among continental scholars, his inter-pretation – encouraged by Darwin himself – can be taken as representative for what “Darwinism”meant at these early times. It also reflects, of course, the major problems that classical Darwinismfaced.


“Darwinism” differed crucially from Weisman’s “Darwinism” and thedifferences rooted, first of all, in their interpretation of variation andinheritance (see Hossfeld 2010).

Darwin read the proof-sheet of Haeckel’s opus magnum and approvedHaeckel’s interpretation of his own theory, albeit expressed concerns aboutHaeckel’s boldness in stating the theory.3 Haeckel had indeed made somestrong claims. One was related to the progressive nature of evolution, where heargued that natural selection gradually, but permanently, works to perfection-ate the organisms’ organization. It is well known that Darwin to some extentsupported the idea of evolutionary progress. On the other hand, he was ex-tremely cautious in formulating this idea and stated that the naturalists were indisagreement about “what is meant by an advance in organisation”. Haeckel,who was influenced not only by Darwin but also by natural philosophy, knewthese doubts but did not take them seriously. Another of his bold statementsrelated to the idea of internal and external constraints in evolution, echoing theidea of orthogenesis.4 Darwin modestly supported the idea of “constraints”,but again, Haeckel made it a “strong argument”.

With his elaborations, Haeckel contributed, in a sense, to the sharpeningof controversial points arising from Darwin’s original concept. Haeckel, asother early Darwinians, was aware of the fact that selection, variation, andretention played a role (though at the time they were not spelled out asgeneral principles), but the exact mechanisms underlying them still needed tobe specified. In trying to do so, he remained speculative, supporting ideas ofnatural selection combined with the concept of environmental influences onthe organism’s heredity, evolutionary constraints, and the idea of progressiveevolution—all compatible with the abstract principles. Thus, it was not by theseprinciples that the further fate of selectionism was decided upon.

3 Darwinism eclipsed: alternative evolutionary theories and the spaceof logical possibilities

Due to the incomplete and sometimes even contradictory data of paleon-tology, anatomy/morphology, biogeography, systematics, and genetics, thereconstruction of evolutionary history and evolutionary mechanisms still had

3“I received a few days ago a sheet of your new work, & have read it with great interest. You conferon my book, the ‘Origin of Species’, the most magnificent eulogium which it has ever received, &I am most truly gratified, but I fear if this part of your work is ever criticized, your reviewer willsay that you have spoken much too strongly.” Letter 5193 - Darwin to Haeckel, 18 Aug [1866](Darwin’s Correspondence Project).4The concept of orthogenesis is often combined with selectionism and/or inheritance of acquiredcharacteristics. In contrast to the claim that natural selection operates on a very copious or eveninexhaustible material generated by an unconstrained variation, orthogenesis holds that variationis strongly constrained and proceeds only in one out of many potential directions within eachphylogenetic line.


to be provisional and even speculative (Gould 1977, 2002; Bowler 1983, 1992).Failure to answer convincingly the questions concerning heredity, variation,evolutionary progress, and orthogenetic series indicated that Darwinism as atheoretical system was still incomplete. This weakness meant water on the millsof theories of evolution that were considered rivals of Darwinism. Indeed, inthe heydays of these rivals in the first decades of the 20th century, a situationprevailed in evolutionary biology in which the Darwinian theory of naturalselection became just one of an entire set of more or less plausible theoriesof how evolution proceeds. In the majority of cases, these were complextheoretical systems that combined the elements of selectionism with non-selectionist ideas. The variety of theories was so large that it was judged toalmost exhaust the “space of logical possibilities” for explaining phylogenetichistory (Zavarzin 1979).

One of the theories was neo-Lamarckism, often seen as a major alternativeto selectionism. By claiming that acquired characteristics are heritable, neo-Lamarckians suggested a logically consistent explanation for the high rateof evolutionary change and the appearance of complex adaptive structures.(The Darwinians had difficulties with explaining these phenomena on thebasis of their core assumption of variation being random.) However, themajority of the 20th century Lamarckians did not reject completely the ideaof natural selection and other evolutionary mechanisms. To the contrary, theyoften combined the inheritance of acquired characteristics with the Darwinianidea of natural selection, orthogenesis, mutationism, etc. Furthermore, neo-Lamarckism itself never represented a monolithic theory, and the notion ofthe inheritance of acquired characteristics is not more than a kind of ‘umbrella-concept’ for several different explanations of the evolutionary mechanism.5

A second theory that became especially popular in the first half of the20th century, and that was partly rivaling Darwinism, was based on the ideaof directed evolution (orthogenesis). It came in more than twenty differentvariants (Levit and Olsson 2006). Although some of them included finalisticideas (see Mayr 1982, p. 959), the common denominator of all of them wasthe idea of morphological, molecular, or other constraints supposed to guidephylogeny in such a way that the mechanism of natural selection becomessuperfluous either for explaining evolution as a whole or for explaining certainperiods of phylogenetic history. The majority of advocates of directed evolu-tion proceeded from the assumption that organisms are predisposed to vary incertain directions and that such biases determine major transitions in evolu-tion. They were convinced that these directions are empirically observable in

5The differences related in particular to the explanation of the exact mechanism of inheritanceand of whether and how heritable features can be acquired. Of the two major neo-Lamarckianapproaches, the first approach—developed by Lamarck himself—claimed that an organ adapts orloses its adaptive value according to its actual functional value in everyday life. (Favorite example:the blindness of cave animals.) The second approach favored the idea of a direct environmentaleffect on the organism’s heredity, an approach known as Geoffroyism.


the paleontological record and clearly definable. Accordingly, it was assumedthat evolutionary events follow certain clearly defined laws and restrictions,and that evolution proceeds to a significant or predominant extent in a non-adaptive fashion. In other words, orthogenesis was strongly coupled with theidea of non-adaptive trends in evolution.6

A third major anti-Darwinian theory in evolutionary biology wassaltationism—which is still influential today. The advocates of saltationism(e.g. Bateson, Goldschmidt, Schindewolf) rejected the Darwinian idea of aslowly and gradually growing divergence of characteristics as the only sourceof evolutionary change. They did not necessarily deny gradual variation ornatural selection any relevance, but claimed that major changes in the “bodyplans” come into being as a result of sudden, discontinuous and uniquechanges, e.g., in a series of macro mutations. The latter were held responsiblefor the sudden appearance of new higher taxa, while the small variationsresulting from natural selection were supposed to be responsible for theadaptations below the species level, i.e. the “fine tuning” adjusting organismsto their environment. Hence, they were considered of secondary importance.7

Considering Darwinism and all its rivals of the time, an impression can begained of how rich are the logical possibilities for making sense of evolutionarychange in the biological domain. There is no logical restriction implied bythe three “Darwinian” principles—as can be seen from the fact that they areshared by all these variants. The relationships between the various explanatoryapproaches and their mutual (in-) compatibility are determined by a plethoraof other doctrines. Thus, pure neo-Lamarckism and strict selectionism arenot mutually compatible, because the inheritance of acquired characteristicsalready guarantees adaptive changes and natural selection loses its shapingrole (in the presence of adaptive mutations, natural selection becomes asecondary evolutionary mechanism—raising the question of what could beassumed to guide the inheritance of acquired variation). Saltationism andmutationism are incompatible with all historical forms of Darwinism, becauseevolution cannot be gradual and saltational at the same time. The latter twoapproaches are incompatible with orthogenesis, because evolutionary changescannot be random and directed at the same time. Orthogenesis, in turn, isthe more difficult to align with Darwinism, the more constrained the randomprocess of variation is imagined to be.

6Although they often used an own terminology, many scientists contributed to the orthogeneticapproach. Some (like Berg 1926) coupled the concept of orthogenesis with saltationism and theidea of direct environmental impact on the organism’s heredity (i.e. “Geoffroyism”, see Levit andHossfeld 2005).7Finally, there existed the theoretical movement of old-Darwinism (as opposed to neo-Darwinism). Insisting on Darwin’s original approach, it combined neo-Lamarckism, orthogenesis,mutationism, and selectionism. By that time, the proponents of old-Darwinism were trying tofigure out the exact role of all these mechanisms in the evolutionary process. See Levit andHossfeld (2006) for details.


4 Modern Darwinism: the synthetic theory and the expanded synthesis

In the first third of the 20th century, Darwinism and all its scientific rivalsappeared to be on equal terms regarding their plausibility. Accordingly, manyoutstanding evolutionary biologists of those times subscribed to some form ofa “synthetic” theory, combining various selectionist and non-selectionist argu-ments in a way that tried to explain the diversity of the living world by somemajor mechanisms of evolution combined with various auxiliary mechanisms.To some extent, this was in accord with Darwin’s own appeal to additionalhypotheses, but made the resulting theoretical systems awkward, sluggish, andsometimes even amorphous. In the understanding of the early “synthesizers”,certain principles (for example, the principle of natural selection) were validfor certain domains or levels of biological reality, but not for others.

Before this background the selectionist turn of evolutionary biologistsworldwide from the mid 1930s onwards came suddenly and was remarkablyradical (as if all of them were passionate Kuhnian revolutionaries). All of asudden it was recognized that the nature of inheritance and variation could bere-interpreted in the light of a new genetics. Following a re-interpretation, itwas realized that biological systematics could be incorporated into a generaltheory of evolution involving the fields of palaeontology and evolutionarymorphology. The paleontological data fit the new explanatory paradigm devel-oped in molecular genetics and microsystematics. Moreover, it became clearthat the theory not only covered the findings at the population level, butcould also explain many macroevolutionary phenomena without appealing tosaltationism, orthogenesis, or neo-Lamarckian mechanisms.8

The “synthetic theory of evolution” was based on three crucial assumptions:(1) natural selection is the major direction-giving factor in evolution; (2)random mutations and recombinations cause variation and therefore imply theincorporation of genetics into the theory; (3) geographic isolation is the mostimportant mechanism separating populations (Junker and Hossfeld 2009).The original “Origin-of-Species-Darwinism” (Darwin 1859, 1871) served as amotivation and a paradigmatic framework, but the scope and content of thenewly established theoretical system went far beyond the historical sourcesand corrected them in many respects. In forming an alliance with genetics, thesynthetic theory of evolution not only enriched itself, but also restricted thepanoply of concepts that previously appeared potentially or actually accept-able. The concepts of inheritance of acquired characteristics, orthogenesis, andsaltationism were rejected as incompatible with what established itself now asmainstream “Darwinism” in the form of the synthetic theory. Thus it were thearchitects of the “second Darwinian revolution” (Mayr 1991) who, for the firsttime, drew an ultimate and clearly defined line between “Darwinism” and itsactual and potential rivals.

8See Mayr (1982, p. 607) and Levit et al. (2008b).


However, the synthetic theory did not solve all problems and retrospectivelyappears to have been too simplistic. There were still a whole range of phenom-ena which remained unexplained. For example, the “phylogenetic explosions”that had been observed by Schindewolf and Beurlen in the paleontologicalrecord (Beurlen 1930) remained a puzzle. Eldredge and Gould (1972) tookthis problem up again and proposed the well known concept of punctuatedequilibria as a solution. This was first thought to represent an alternative tothe gradualism of the synthetic theory of evolution, since the authors initiallyintended to demonstrate that there are domains of evolution inaccessiblefor gradualism. However, recent accounts of the problem have shown thatevolution is both gradual and punctuated, since even accelerated evolutionarytransitions are gradual processes at the micro-evolutionary level (Bokma 2002;Gould 2002; Kutschera and Nicklas 2004). What is still an issue, though, is theexact role of species selection and group selection from the empirical and thetheoretical points of view (Reif et al. 2000).

Yet another problem posed to the synthetic theory that requires furtherclarification and possibilities for further expansions is the whole complex ofissues around developmental biology and its relations with other biologicaldisciplines (Hall et al. 2003). The most recent synthetic movement of an “eco-logical developmental biology” tries to integrate the whole range of relevantdisciplinary domains in a further extension of the classic synthetic theory ofevolution (see, e.g., Gilbert and Bolker 2003). In that light, for example, allelicvariation in the structural genes is no longer the only source of variation. Thereare two other sources of variation accessible for natural selection, namelyallelic variation in the regulatory regions of genes and “developmentally plasticvariation” (Gilbert and Epel 2009, p. 318). The latter can be crucial for macro-evolutionary processes producing the variations associated with phyla andclasses of species. This example shows that the occurrence of variation as suchdoes not mean much as to whether it is to be considered a Darwinian principleor not. Depending on how the nature of variation is interpreted, the resultsof its combination with the principle of selection will also vary. The “devilis in the details” (Vromen 2007) indeed. Amendments to a general conceptsuch as variation that seemed of only technical character turn out to make aradical difference with respect to whether biological evolution is explained ina Darwinian or a non-Darwinian way.9

9Contemporary mainstream “Darwinism” also reflects on whether, and to what extent, elementsof rivaling approaches could play a role within an expanded synthetic theory. For instance,there is a debate revolving around the role of developmental and evolutionary constraints (e.g.,Wimsatt and Schank 1988) which reminds one of the idea of orthogenesis. Neo-Lamarckian ideasare reconsidered in view of the debate concerning epigenetic phenomena (Jablonka and Lamb2006). There is still a principle difference, though, between neo-Lamarckian soft inheritance andepigenetics so that the selectionist foundations of the theory are not compromised. See Haig (2007)and Gilbert (Gilbert and Epel 2009, p. 449). Also, mutationist-saltationist conjectures draw newattention among geneticists.


It is not our purpose here to give a full account of the directions in whichthe synthetic theory currently expands.10 The examples are rather given todemonstrate that contemporary evolutionary theory development continuesto be driven by bottom-up modifications, syntheses, and enrichments. Itexpands partly by accounting for elements that were previously consideredanti-Darwinian, but that can now be made sense of by the discovery of yetother evolutionary mechanisms consistent with the general framework ofmodern Darwinism. This is not to say, though, that “anything goes” providedonly the principles of variation, selection, and retention are complied with.It is the specific combination of the particular evolutionary mechanisms andtheir “details” that matter for whether or not an extension of the complextheoretical system is “Darwinian” in nature.

5 How useful can generalized Darwinism be for evolutionary economics?

The brief review of the developments in evolutionary biology in the previoussections shows that different variants of the theory of variation, selection,and inheritance have played a role in almost all approaches that have beeninfluential over the past 150 years in biology, whether Darwinian, non-Darwinian, or even anti-Darwinian. For this very reason we have argued thatvariation, selection, and inheritance as such are no particularly “Darwinian”.However, the sketch of the history of evolutionary biology also shows thatexplanatory progress in this discipline was driven by the inquiries into de-tails of evolutionary processes that were not captured by abstract principles.These inquiries evoked more general conclusions which were scrutinized andeventually modified or rejected by further work on “the details”—what wehave referred to as a bottom-up research strategy. In comparison, GeneralizedDarwinism proposes an entirely different research strategy for evolutionaryeconomics. Starting from the abstract principles of variation, selection, andinheritance, it suggests deploying them in the search for auxiliary hypothesesthat explain the details of evolutionary processes in the economic domain. Thisproposal raises several questions which, in our view, warrant reflections thatlead up to a critical assessment of the usefulness of Generalized Darwinism forevolutionary economics.

A first point that, we think, should be taken into consideration is thecomplete absence of something comparable to the top-down strategy of Gen-eralized Darwinism from the history of evolutionary biology. The lesson thatcan be learned from the leading evolutionary science is that evolution canbe expected to be an extremely complex process the explanation of whichrequires a similarly complex network of specific hypotheses. In biology, this

10Nor does space suffice here to address the recent contributions of philosophers of biology to thedebate between selectionist and evolutionary developmentalists. See for example Sober (1994),Callebaut and Rasskin-Gutman (2005), Samson and Brandon (2007).


emerged from an extremely broad, rich, and controversial discourse runningthe whole gamut from the details of the paleontological record to the findingsof molecular biology. It was the attempt to give the complex network of specifichypotheses a coherent causal and functional structure that eventually led tothe formulation of general principles—not the other way round. One maywonder what evolutionary biology would look like today had it proceededin a top-down fashion, starting just from the abstract principles of variation,selection, and inheritance. Why should what appears a strange idea with regardto evolutionary biology be a promising idea when it comes to other domainssuch as that of evolutionary economics?

A second point arises from the fact that Generalized Darwinism is basedon the claim that the three abstract principles derived by isolating abstractionfrom findings in the biological domain govern “all complex population systems(in the biological and social worlds)” (Hodgson 2009). Hence, evolution isconceived of as a homologous process in different disciplinary domains—among them, in particular, the domain of evolutionary economics. However,the homology postulate is an empirical claim. It is far from being self-evident,since an abstraction arising from a bottom-up discourse in one particulardiscipline is influenced by the domain-specific (ontological) conditions of thatdiscipline which are not necessarily the same elsewhere. Hence, the claimneeds to be supported by empirical evidence for each disciplinary domain forwhich its validity is asserted.

As far as evolutionary economics is concerned, a proof is lacking and,as Nelson (2006) has argued, doubts may be raised as to whether it canbe expected to ever be delivered. The best way to confirm the assertedhomology with evolutionary biology would be the demonstration that theprinciples of variation, selection, and inheritance emerge identically fromisolating abstraction in a “bottom-up” discourse reflecting specific researchresults on evolutionary processes in the economy.11 However, such a bottom-up discourse is precisely the opposite of what the proponents of GeneralizedDarwinism have in mind. Their idea is to invoke variation, selection, andinheritance in a top-down fashion as principles governing economic evolutionand, assuming their validity, to search for suitable “auxiliary hypotheses” bywhich the abstract scheme can be filled with economic explanatory substance(see Aldrich et al. 2008).

A third problem we see relates to the dichotomy of abstract principles, onthe one hand, and “auxiliary hypotheses” on the other. (To recall, the latter

11On a priori grounds, there is no reason to expect that a bottom-up research strategy wouldindeed lead to the three principles suggested by Generalized Darwinism or any other onesdiscussed in evolutionary biology such as the principle of orthogenesis. Indeed, a glance at worksin economics that may be considered exemplars of a bottom-up strategy points at a large variety ofprinciples that depend on the context and cannot easily be generalized. See, for instance, Penrose(1953) for the firm context, Klepper (1997) for the industry context, Witt (2001) for the context ofconsumption, Metcalfe et al. (2006) for that of economic growth, and Richerson and Boyd (2005)or North et al. (2009) for the context of institutions.


are needed because, by its very nature, Generalized Darwinism lacks discipli-nary substance.) By taking the validity of its abstract principles for granted,Generalized Darwinism factually turns into a search heuristic designed tofind something equivalent to, or reminiscent of, variation, selection, andinheritance at the disciplinary level. On purely logical grounds, this is a possibleway to proceed—as much as it is possible to start from any other generalprinciples and to look for suitable auxiliary hypotheses on “the details”. Butthere is a non-negligible risk involved in such a top-down approach of inducingsystematic heuristic biases. The interpretation of reality can be seriously biasedby selectively recognizing only those elements that seem to fit the logic ofthe chosen principles and ignoring all others, or by constructing disciplinarycorrelates that actually misrepresent the economic conditions.

In view of the criticism raised against Generalized Darwinism, particularlyits top-down research strategy, it may be asked what evidence for fruitfulapplications there is in works so far published on Generalized Darwinism.Stoelhorst (2008) has argued that the three principles of Generalized Dar-winism can be seen as part of an (abstract) explanans of an evolutionaryexplanation that can be put to work in the economic domain only if it is clearwhat the explanandum is going to be. One may add that it is precisely inspecifying the explanandum that domain-specific “details” have to be enteredand elaborated via “auxiliary hypotheses”. But, as Stoelhorst notes, this hasnot been clarified in the writings on Generalized Darwinism. Indeed, thealready quite numerous publications on Generalized Darwinism offer little onthe crucial “auxiliary hypotheses”.12

In their manifesto in defense of Generalized Darwinism, Aldrich et al.(2008) do not mention any specific implications of their approach for explain-ing evolution in the economy. They posit that domain-specific substance canbe developed on the basis of the reflections about the unit of selection. Inthat context, they refer to the replicator–interactor dichotomy and discussit at an abstract level. An interactor is defined as in Hull (1988, p. 408) as“an entity that directly interacts as a cohesive whole with its environmentin such a way that this interaction causes replication to be differential”.Replicators are defined as entities that are replicated in a way that “involvesa causal relationship between two or more entities, where there is substantialsimilarity between the original and the replicated entities, and where infor-mation concerning adaptive solutions to survival problems is passed fromone set of entities to another”. But what are the replicators and interactorsat the concrete economic level and how precisely are they varied, selected,and inherited, and how does all this contribute to understanding man-madeevolution?

12Stoelhorst draws a different conclusion. He suggests specifying the explanandum in terms ofan abstract notion of evolution as a “process of change that leads to adaptive complexity”(ibid. p. 347). This is again a notion distilled from evolutionary biology by isolating abstraction.Stoelhorst does not point out in what kind of economic processes “adaptive complexity” couldfigure as explanandum.


In the extensive discussions of interactors and replicators in their mostrecent book, Hodgson and Knudsen (2010) do not explain in any detailhow replication and interaction work and drive evolutionary processes in theeconomy. In terms of concrete applications, their reflections basically comedown to suggesting certain pairings of interactors and replicators.13 It is leftopen, however, how the developed categorizations lead to explanations ofconcrete economic evolutionary phenomena. In a paper inquiring more deeplyinto the role of firms as interactors, Hodgson and Knudsen (2004) argue thathabits and routines, which are more or less specific to individual firms, repli-cate under the influence of selection forces. But again, neither the selectionforces nor the assumed selection and replication mechanisms are explained indetail. The authors mention the Price equation (Price 1970; see also Frank1995)—another very abstract concept. Yet, they do not demonstrate whatcausal explanations can be derived regarding specific evolutionary processes ineconomics. Hodgson and Knudsen repeatedly claim a broad agreement of theirapproach with the argumentation and formalizations in Nelson and Winter’s(1982) seminal work on the evolution of firms and industries. (See, however,the reservations Nelson 2006 makes regarding this claim.) Yet, they do notshow what explanatory conjectures their Generalized Darwinism is actuallyable to contribute beyond those obtained already by Nelson and Winter onthe basis of only a loose analogy to the notion of natural selection.

In view of the problems that have been pointed out in this section, the ratherthin outcome of the top-down approach of Generalized Darwinism in terms ofconcrete, new explanations offered for evolutionary phenomena in economicsmay not come as a surprise. By comparison with the successful bottom-upresearch practice in evolutionary biology, one may infer that the reason forthis outcome lies in the nature of heuristically proceeding top-down fromabstract principles with uncertain validity for economics to suitable auxiliaryhypotheses representing the domain-specific, economic explanatory substance.If so, the recommendation for evolutionary economics would be to focus onanalyzing the huge variety of specific evolutionary processes in the economyat a concrete level, and only when explanatory progress has been made atthat concrete level to engage in a (bottom-up) discourse of how the complexset of specific hypotheses can be organized into a more coherent causal andfunctional structure.

Such a discourse may eventually lead to the formulation of general princi-ples. Since all of the specific economic processes are instances of man-madeevolution, it may turn out that general principles akin to economic evolutionrelate to the core features of human adaptiveness. These are individual andcollective learning, experimentation, insight, inventiveness, and the restlessmotivations driving these human forms of adaptations. If so, to understand

13Examples that are given for pairings of interactors vs. replicators are, among others, {organiza-tions vs. routines/habits/genes}, {human groups vs. habits/genes}, {individuals vs. habits/genes}, and{scientific institutions vs. scientific/technological knowledge}, {states vs. laws}, {families/tribes vs.customs}, see ibid. Table 7.2 and Table 8.1.


and explain how these features shape the specific, evolutionary processes inthe economy may be more important than demonstrating that the abstract hullof Generalized Darwinism can be stretched to fit them.

6 Conclusions

In a recent series of publications, “Generalized Darwinism” has been proposedas a new overarching research strategy that is based on the assumption of afundamental homology between evolution in nature and the evolution of theeconomy. The principles of variation, selection, and retention that have beendistilled from evolutionary biology by isolating abstraction are claimed to begenerally valid. It is suggested to apply these abstract principles as a unifyingframework for all evolutionary theories. By a brief reconstruction of thedifferent historical forms of Darwinism, we have shown that the identificationof these abstract principles with Darwinism is misleading. Moreover, on apriori grounds, other principles—non-Darwinian or even anti-Darwinian onessuch as, e.g., orthogenesis, saltationism, or neo-Lamarckism—could claim asimilar plausibility in explaining economic evolution.

The crux with such allegedly unifying abstract principles derived by isolatingabstraction from findings in other domains is that they provide but an abstracthull. In order to become a useful heuristic device, they need to arrive atdomain-specific explanation which, in turn, would require adding substance byhypotheses on the disciplinary “details” of actual evolutionary processes, e.g.in the economy. This is, of course, what is done in the first place in a bottom-up research strategy as it has fruitfully been practiced in the development ofDarwinism in evolutionary biology. In the final section, doubts have thereforebeen raised as to whether the top-down approach propagated by GeneralizedDarwinism can overcome its inherent limitations and develop into a fruitfulresearch strategy for evolutionary economics.

Acknowledgements Support by the Deutsche Forschungsgemeinschaft (Ho 2143, 9–1) forHossfeld and Levit’s research on the history of evolutionary biology is gratefully acknowledged.The authors also wish to thank the Situating Science: Works in Progress discussion round at theUniversity of King’s College, Halifax (Levit) and G.Hodgson, J.-W. Stoelhorst, and Jack Vromenfor inspiring discussions on the controversy (Witt).

References

Aldrich HE, Hodgson GM, Hull DL, Knudsen T, Mokyr J, Vanberg VJ (2008) In defense ofgeneralized Darwinism. J Evol Econ 18(5):577–596

Andersen ES (2009) Schumpeter’s evolutionary economics. Anthem, LondonBerg LS (1926) Nomogenesis or evolution determined by law, 2nd edn. 1969 MIT Press,

CambridgeBeurlen K (1930) Vergleichende stammesgeschichte. Borntraeger, BerlinBokma F (2002) Detection of punctuated equilibrium from molecular phylogenies. J Evol Biol

15(6):1048–1056


Bowler PJ (1983) The eclipse of Darwinism. The John Hopkins University Press, BaltimoreBowler PJ (1992) The non-Darwinian revolution. The Johns Hopkins University Press, BaltimoreBuenstorf G (2006) How useful is generalized Darwinism as a framework to study competition

and industrial evolution? J Evol Econ 16(5):511–527Callebaut W, Rasskin-Gutman D (eds) (2005) Understanding the development and evolution of

natural complex systems. MIT Press, Cambridge, MACampbell DT (1965) Variation and selective retention in socio-cultural evolution. In: Barringer

HR, Blankenstein GI, Mack RW (eds) Social change in developing areas: a reinterpretationof evolutionary theory. Schenkman, Cambridge, MA, pp 19–49

Cordes C (2006) Darwinism in economics: from analogy to continuity. J Evol Econ 16(5):529–541Crozier GKD (2008) Reconsidering cultural selection theory. Brit J Philos Sci 59(3):455–479Dawkins R (1983) Universal Darwinism. In: Bendall DS (ed) Evolution from molecules to man.

Cambridge University Press, Cambridge, UK, pp 403–425Darwin C (1859) On the origin of species by means of natural selection, or the preservation of

favoured races in the struggle for life. Murray, LondonDarwin C (1871) The descent of man, and selection in relation to sex. Murray, LondonDarwin C, Wallace AR (1858) On the tendency of species to form varieties,and on the perpetua-

tion of varieties and species by natural means of selection. J Proc Linn Soc Lond 3:45–63Eldredge N, Gould SJ (1972) Punctuated equilibria: an alternative to phyletic gradualism. In:

Schopf TJM (ed) Models in paleobiology. Freeman, Cooper and Co., San Francisco, pp 82–115Frank SA (1995) George Price’s contributions to evolutionary genetics. J Theor Biol 175:373–

388Gilbert SF, Bolker JA (2003) Ecological developmental biology: preface to the symposium. Evol

Dev 5(1):3–8Gilbert SF, Epel D (2009) Ecological developmental biology. Sinauer, SunderlandGould SJ (1977) Ontogeny and phylogeny. The Belknap Press of Harvard University Press,

CambridgeGould SJ (2002) The structure of evolutionary theory. The Belknap Press of Harvard University

Press, CambridgeHaeckel E (1866) Generelle morphologie der organismen, 2 Bde. Georg Reimer, BerlinHaig D (2007) Weismann rules! OK? Epigenetics and the Lamarckian temptation. Biol Philos

22(3):415–428Hall BK, Pearson R, Müller GB (2003) Environment, evolution and development: toward a

synthesis. MIT Press, Cambridge, MAHallpike CR (1985) Social and biological evolution I. Darwinism and social evolution. J Soc Biol

Syst 8(2):129–146Hallpike CR (1986) Social and biological evolution II. Some basic principles of social evolution. J

Soc Biol Syst 9(1):5–31Hodgson GM (2002) Darwinism in economics: from analogy to ontology. J Evol Econ 12(3):259–

281Hodgson GM (2009) Agency, institutions, and Darwinism in evolutionary economic geography.

Econ Geogr 85(2):167–173Hodgson GM, Knudsen T (2004) The firm as an interactor: firms as vehicles for habits and

routines. J Evol Econ 14(3):281–307Hodgson GM, Knudsen T (2006a) Why we need a generalized Darwinism, and why generalized

Darwinism is not enough. J Econ Behav Organ 61(1):1–19Hodgson GM, Knudsen T (2006b) The nature and units of social selection. J Evol Econ 6(5):477–

489Hodgson GM, Knudsen T (2010) Darwin’s conjecture—the search for general principles of social

and economic evolution. University of Chicago Press, ChicagoHossfeld U (2010) Ernst Haeckel. Biographienreihe absolute, Orange, Freiburg i. Br.Hossfeld U, Olsson L [Hrsg.] (2009) Charles Darwin. Zur evolution der arten und zur entwicklung

der erde. Frühe schriften zur evolutionstheorie. Suhrkamp Studienbibliothek 13, Suhrkamp,Frankfurt a. M

Hull DL (1988) Science as a process: an evolutionary account of the social and conceptual devel-opment of science. University of Chicago Press, Chicago

Hull DL (2001) Science and selection: essays on biological evolution and the philosophy of science.Cambridge University Press, Cambridge


Jablonka E, Lamb M (2006) Evolution in four dimensions. MIT PressJunker T, Hossfeld U (2009) Die Entdeckung der evolution, 2nd edn. WBG, DarmstadtKlepper S (1997) Industry life cycles. Ind Corp Change 6(1):145–181Kutschera U, Nicklas KJ (2004) The modern theory of biological evolution: an expanded synthesis.

Naturwissenschaften 91:255–276Levit GS, Hossfeld U (2005) Die nomogenese: eine evolutionstheorie jenseits des Darwinismus

und Lamarckismus. Verhandlungen zur geschichte und theorie der biologie 11:367–388Levit GS, Hossfeld U (2006) The forgotten “Old Darwinian” synthesis: the evolutionary theory of

Ludwig H. Plate (1862–1937). Int J Hist and Ethics of Nat Sci, Technol Med 14(1):9–25Levit GS, Olsson L (2006) “Evolution on rails”: mechanisms and levels of orthogenesis. Ann Hist

Philos Biol 11:99–138Levit GS, Meister K, Hossfeld U (2008a) Alternative evolutionary theories from the historical

perspective. J Bioecon 10(1):71–96Levit GS, Simunek M, Hossfeld U (2008b) Psychoontogeny and psychophylogeny: the selectionist

turn of Bernhard Rensch (1900–1990) through the prism of panpsychistic identism. TheoryBiosci 127(4):297–322

Mayr E (1982) The growth of biological thought: diversity, evolution and inheritance. The BelknapPress of Harvard University Press, Cambridge, MA

Mayr E (1991) One long argument: Charles Darwin and the genesis of modern evolutionarythought. Harvard University Press, Cambridge, MA

Metcalfe JS (1998) evolutionary economics and creative destruction. Routledge, LondonMetcalfe JS, Foster J, Ramlogan R (2006) Adaptive economic growth. Camb J Econ 30(1):

7–32Nelson RR (2006) Evolutionary social science and universal Darwinism. J Evol Econ 16(5):491–

510Nelson RR, Winter SG (1982) An evolutionary theory of economic change. Harvard University

Press, Cambridge, MANorth DC, Wallis JJ, Weingast BR (2009) Violence and social orders—a conceptual framework

for interpreting recorded human history. Cambridge University Press, CambridgePenrose ET (1953) The theory of the growth of the firm. Blackwell, Oxford.Price GR (1970) Selection and covariance. Nature 277:520–521Reif W-E (2000) Darwinism, gradualism and uniformitarianism. N Jb Geol Paläont 11:669–680Reif W-E, Junker T, Hossfeld U (2000) The synthetic theory of evolution: general problems and

the German contribution to the synthesis. Theory Biosci 119(1):41–91Richerson P, Boyd R (2005) Not by the genes alone: how culture transformed human evolution.

University of Chicago Press, ChicagoSamson R, Brandon R (eds) (2007) Integrating development and evolution. MIT Press, Cam-

bridge, MASober E (ed) (1994) Conceptual issues in evolutionary biology. MIT Press, Cambridge, MAStoelhorst JW (2008) The explanatory logic and ontological commitments of generalized Darwin-

ism. J Econ Methodol 15(4):343–363Vromen J (2007) Generalized Darwinism in evolutionary economics—the devil is in the details.

Papers on Economics and Evolution # 0711, Max Planck Institute of Economics, JenaVromen J (2008) Ontological issues in evolutionary economics: the debate between generalized

Darwinism and the continuity hypothesis. Papers on Economics and Evolution # 0805, MaxPlanck Institute of Economics, Jena

Weismann A (1885) Die Kontinuität des Keimplasmas. Jena, FischerWeismann A (1893) The Germ.Plasm: a theory of heredity. Scribner, New YorkWilkins JS (1998) The evolutionary structure of scientific theories. Biol Philos 13(4):479–504Wimsatt WC (1999) Genes, memes and cultural heredity. Biol Philos 14(2):279–310Wimsatt WC, Schank JC (1988) Two constraints on the evolution of complex adaptations and the

means for their avoidance. In Nitecki M (ed.) Progress in evolution. The University of ChicagoPress, Chicago, pp 213–273

Windrum P (2007) Neo-Schumpeterian simulation models. In: Hanusch H, Pyka A (eds) Elgarcompanion to neo-schumpeterian economics. Edward Elgar, Aldershot, pp 405–439

Winther R (2001) August Weismann on germ-plasm variation. J Hist Biol 34:517–555Witt U (2001) Learning to consume—a theory of wants and the growth of demand. J Evol Econ

11(1):23–36


Witt U (2003) The evolving economy. Essays on the evolutionary approach to economics. Elgar,Cheltenham

Witt U (2004) On the proper interpretation of ‘evolution’ in economics and its implications forproduction theory. J Econ Methodol 11(2):125–146

Witt U (2008) What is specific about evolutionary economics? J Evol Econ 18(5):547–575Zavarzin GA (1979) The space of logical possibilities in the diversity of bacteria and their phylo-

genies. Priroda 6:9–19

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Can Darwinism be “Generalized” and of what use would this be