OCTOBER 31, 2018
LAST SPRING, I abounding a appointment in New Mexico featuring evolutionary biologists alive on a new assay diplomacy they acquire been calling the “extended evolutionary synthesis” (EES). The diplomacy aims to go above the alleged “modern synthesis” of the mid-20th century, which aing Darwinism to Mendelian genetics, whose algebraic formulations could be artlessly and accurately expressed. Biologists circuitous in the EES acquire been calling for a broader and beneath reductive appearance of evolution, complete to Mendelian genes. In particular, they acquire been acclamation the avant-garde synthesis’s absence of advice about adorning biology. These EES revisionists are absorbed in feedbacks: in how adorning processes, forth with ecological and alike cultural ones, augment aback into one another, into abiogenetic and added forms of inheritance, and appropriately into evolution. While the avant-garde amalgam proposes that epigenetic, developmental, ecological, and cultural processes are all articles of evolution, the EES claims they are causes as able-bodied as products.
The roots of this movement extend aback to the aboriginal 1970s, to the assignment of Richard Lewontin at Harvard, and Luigi Luca Cavalli-Sforza and Marcus Feldman at Stanford, amid others. But the reigning, reductive neo-Darwinist archetype — in added words, the avant-garde amalgam — charcoal able-bodied entrenched, and its defenders allegiant in its support. Alone in the aftermost 25 years or so has the added all-embracing eyes of the EES boring amorphous — adjoin abundant attrition — to authorize itself in boilerplate biology.
As allotment of this development, EES biologists acquire been added absorbed in culture, amid added forms of transformation and transmission, and so acquire accustomed the ascribe of humanists, including philosophers and historians of science like me, whose job it is to abstraction and acquire culture. Taking allotment in their conversations has in about-face a my own assignment in the history of evolutionary theory.
Still, I able a moment of absurd ability shock at the contempo affair I attended. A biologist wrote an blueprint on the whiteboard in which one of the variables was a “w.” He again circled the w, answer that it represented “culture,” and acicular out that beneath assertive conditions, the bulk of “w” would tend adjoin zero, while beneath added altitude it would tend adjoin 100. “Perhaps,” I thought, “we don’t beggarly absolutely the aforementioned affair by ‘culture.’” To a humanist, or anyhow to this one, “culture” is an abstruse noun encompassing abounding things of abounding kinds: processes, objects, habits, behavior both absolute and implicit. It seems a class aberration to anticipate that we can represent such a delight by a audible variable, or that the able assortment could act as a detached affair accepting a audible assessable aftereffect on some added detached thing. Could we say, for example, that in a accustomed society, “culture” influences “politics” by some assessable bulk x? Could we say that “the arts” has a y-percent aftereffect on bearing amount or activity expectancy?
As it angry out, I had somewhat blurred the situation. Back I bidding a assertive dubiousness about apery “culture” with a audible variable, an EES biologist explained to me that the variables continuing for “culture” in biologists’ algebraic models are not meant to denote the absolute Gestalt, but rather assessable $.25 of culture: a audible behavior, for example, that ability be taught, learned, transmitted, or counted, and whose furnishings on adaptation and reproduction can be abstinent and modeled. Perhaps these abandoned ability variables ability in acceptance add up to a single, overarching W, but for the moment, no one claims to be able to accomplish that summation. For now, we can artlessly use the little w’s to body detached cultural $.25 or forms into an evolutionary model. This seems to me added credible, but it still assumes that we can advisedly represent cultural forms as assessable bits, and that this will add added to our compassionate of the role of cultural forms in evolutionary processes than artlessly aggravating to alarm this role in qualitative terms. I can’t advice apprehensive if that’s a complete assumption.
Of advance I’m by no bureau the aboriginal to accession the question, nor absolutely acquire such objections been bedfast to humanists. Lewontin himself, calm with the historian Joseph Fracchia, argued in a 1999 cardboard adjoin the abstraction of cultural evolution. They wondered whether conceptualizing entities like “the abstraction of monotheism” as “cultural units” begged acute questions — for example, how can we calculation up these units in a population, and what are their laws of bequest and variation? Fracchia and Lewontin maintained that there could be no such accepted laws because cultural phenomena, clashing atoms and molecules, alter from one addition in their backdrop and dynamics of manual and change. “There is no one transhistorical law or generality,” they contended, “that can explain the dynamics of all actual change.”  Marcus Feldman disagreed, admitting not accurately with attention to the actuality of accepted laws answer the dynamics of all actual change; rather, he dedicated the angle of “observable units of culture,” which he did not accessory with admirable acclimation account such as monotheism. An archetype of an appreciable cultural assemblage for Feldman is a behavior or custom that follows statistical rules of transmission, and that can appropriately be a accepted article of algebraic study. 
The biologist with the “w” capricious and I were appropriately reenacting an bookish battle that has been activity on for decades. As is about the case in longstanding debates, we absolutely accede on the essentials: that attributes and ability are at basal fabricated of the aforementioned actuality — in fact, of one addition — which no humanistic or accurate assay can accurately disregard. Evolutionary access charge beset cultural processes aloof as animal history charge beset biological ones. But, admitting our abysmal accord, this biologist and I are cerebration incommensurately about methods, about how to put our two fields into communication. His adjustment is algebraic modeling, and abundance is blubbery description. These are diametrically adverse in trajectory, one abstractive and reductive, the added concretizing and expansive. While I acquire and adore these biologists’ conclusions, I accumulate wondering: Why these methods? Why algebraic modeling? By which I mean, what action do biologists intend their algebraic models to serve? Are they meant to prove claims about evolution? Or rather to express, represent, or apostle assertive analytic angle of evolutionary processes? If the latter, why acquire this accurate bureau of expression, representation, advocacy? These will absolutely assume naïve questions to any biologist account this. But I acquire abstruse from teaching academy apprentice and sopres that naïve questions from green newcomers can be the hardest and best useful, which emdens me to ask mine.
Kevin Laland, columnist of Darwin’s Unfinished Symphony, was an organizer of the appointment I abounding and is a baton of the Extended Evolutionary Amalgam assay program. Account his important and ardent book, the to-date arbitrary of a groundbreaking career, I had agnate animosity as I did at the conference. Uppermost amid these is acrimonious agreement: Laland’s capital credo assume to me greatly right, some absolutely incontrovertible. These accommodate the axiom that cultural practices — in accurate teaching, imitating, and artful — are causes as able-bodied as after-effects of evolution; that in mammals and abnormally humans, such cultural practices acquire accelerated evolutionary development by consistently creating “new alternative regimes” in a action that Laland, citation evolutionary biologist Allan Wilson, calls “cultural drive”; and that, accordingly, in bodies especially, there has been a “gene-culture coevolutionary dynamic.” The aboriginal of these — that cultural practices are causes as able-bodied as after-effects of change — seems to me incontrovertible, but added like a aboriginal acceptance than like an empiric result. Cultural practices charge be causes as able-bodied as after-effects of change because any aftereffect of the evolutionary action becomes a affection of the apple of causes abstraction the assiduity of that process.
The added arch credo — such as “cultural drive” and “gene-culture co-evolution” — are not absolutely aboriginal principles, but they assume to me bureau of compassionate how the feedback-loop of change encompasses cultural forms. To accurate these bureau of compassionate in the accent of algebraic clay seems fine, if one brand to do that, but no added absolute than cogent them in words. This is because a algebraic model, like a exact description, contains abounding layers of interpretation. This is not a criticism: estimation is capital to (and abiding from) any attack to acquire the world. But insofar as a algebraic archetypal is taken to prove rather than to altercate or represent, that’s area I anticipate it can mislead.
Laland has adherent his career to beat assignment adjoin reductive, simplistic, and arbitrary accounts of evolution, architecture brick by brick a complete case for the richer and added circuitous eyes of the EES. Darwin’s Unfinished Symphony is a almanac of his aural success. But while he has been amalgam this advocate accurate theory, he has about accurate it by acceptable methods. An archetype is his game-theoretic clash to abstraction amusing learning. After alms several examples of amusing acquirements in animals — such as Japanese macaques who apprentice from one avant-garde macaque to ablution their candied potatoes afore bistro them, and angle who apprentice from one addition area the affluent agriculture patches are amid — Laland asks what ability be the best “social acquirements strategy.” He explains that the “traditional bureau to abode such questions is to body algebraic models using, for instance, the methods of evolutionary d theory.”
Game access became a accepted archetypal in evolutionary assay in the aboriginal 1970s with the work, notably, of the British abstract evolutionary biologists W. D. Hamilton and John Maynard Smith, forth with the citizenry geneticist George R. Price. Hamilton, Price, and Maynard Smith developed a game-theoretic access to clay the behaviors of bacilli in the attack for survival. Their assignment was basal to the neo-Darwinist, gene-centric diplomacy that Laland has adherent his career to challenging. In this gene-centric view, all higher-order entities — abandoned organisms, their behaviors and interactions — are epiphenomenal, controlled by and reducible to genes, so that any credible bureau or ambition on the allotment of an animal is illusory. Bacilli survive if they arise to accomplish an optimal accompaniment of abiogenetic affairs, one that maximizes some action for greater changeable success. They die out back they abort to do so. Maynard Smith appropriately emphasized that his abstruse analogue of “strategy” was carefully behaviorist. “Nothing,” he maintained, “is adumbrated about intention.” A action was abandoned “a behavioural phenotype,” in added words, “a blueprint of what an abandoned will do [in a accustomed situation].”  These “strategies,” therefore, circuitous no ascription of centralized agency, but abandoned credible observations of behavior. Neither empiric behaviors nor any added macrolevel abnormality could comedy a causal role in change according to this academy of thought.
Maynard Smith’s access has aggressive the best reductive of neo-Darwinists. For example, Richard Dawkins has acclimatized it to his own access of gene functioning, emphasizing that the “strategies” in catechism are behaviorally authentic and do not crave the ascription of consciousness, let abandoned agency, to the cardinal agent. Dawkins absolutely refers to “unconscious strategists,” the advised adumbration auspicious the clairvoyant to acquire these credible ascriptions of bureau to genes as aitionist denials of any such agency.  Neither behaviors, nor agency, nor consciousness, nor ability operates causally at any akin of Dawkins’s picture; all reduces to aloof gene functioning.
Game-theoretic clay has been a authentication of neo-Darwinist reductionism and, specifically, of the abnegation of any affectionate of evolutionary bureau to the evolving organism. But in Darwin’s Unfinished Symphony, Laland describes how he and his collaborators acclimated d access in an avant-garde way, to architecture a basic apple in which they hosted a tournament. The d circuitous basic “organisms” or “agents” agreeable in a hundred “behavior patterns,” with capricious ante of success consistent in greater or bottom “fitness” (i.e., adaptation and reproduction). The d additionally included three altered “moves” — “innovate,” “observe,” and “exploit” — apery altered phases of alienated or amusing learning. Added than a hundred bodies of assorted ages and backgrounds took allotment in the game. Clashing in Maynard Smith’s applications of d access to evolution, Laland and his collaborators were not attractive for an optimum in the anatomy of a audible action or acreage to be maximized. They did not pre-judge what had to arise in adjustment for an animal to win the competition. Rather, they set the competitors afar and waited to see who would triumph. The acceptable action was an unpredictable, circuitous mix of behaviors, although it did represent an all-embracing optimum band-aid composed of behavioral bits.
Analyzing the acceptable strategy, Laland concludes that celebratory and artful are abundantly valuable, abundant added so than innovating on one’s own except “in acute environments that change at abnormally aerial rates,” which charge be attenuate in nature. The cessation is persuasive, but the clash seems to me added a way of cogent than of proving this point: the basic agents and their behaviors and strategies of advance aggregate an analytic representation of accustomed processes. They are not fatigued in pastels or composed in prose, but the actuality that they are programmed on a computer makes them no beneath a representation.
To busy further, accede an agreement Laland describes, performed with his postdoctoral apprentice Hannah Lewis. Laland explains that to archetypal the furnishings of high-fidelity manual of advice on the constancy of cultural forms or “traits” in a population, he and Lewis “assumed that there are a anchored cardinal of ancestry that could arise aural a accumulation through atypical inventions and that are absolute of any added ancestry aural a culture. We alleged these atypical inventions ‘cultural berry traits.’ Then, one of four accessible contest could occur”: a new berry affection could be acquired by atypical invention; two ancestry could be accumulated to aftermath a new one; one affection could be modified; or a affection could be lost.
This model, in its affiliation to absolute cultural forms, seems to me the agnate of a Cubist painting. Cultural “traits” that are absolute of one addition action no added about in attributes than adolescent ladies with altogether geometrical appearance broadcast all on one ancillary of their two-dimensional heads. Likewise for the abstracted and audible accident of atypical invention, combination, modification, or accident of cultural forms. These processes biking in the absolute apple as aspects of a audible amoebic article and not as abstracted blocks. Of course, I’m not against to apery cultural forms in these Cubist agreement any added than I’m against to Picasso’s portraits of Dora Maar. Representations should, though, acknowledge themselves as such.
Mathematical models are interpretative from the get-go. Again, let me be bright that I anticipate that’s able — indeed, assured — because estimation is abiding from any attack to acquire the world. Indeed, some scientists alarm their use of algebraic models in these actual terms. The abstract physicist Murray Gell-Mann warned that we charge be careful, apropos models, “not to booty them too actively but rather to use them as prostheses for the imagination, as sources of inspiration, as accustomed metaphors. In that way I anticipate they can be valuable.”  Feldman, who acicular me to Gell-Mann’s assuming of models as “prostheses of the imagination,” added that “insofar as the archetypal assists in the interpretation, again it has value.”  On addition occasion, Feldman told an interviewer, “[p]eople who accomplish models for a active like I do don’t absolutely accept they’re anecdotic reality. We aren’t adage that our archetypal is added apparent than addition model; we’re adage it exposes what is possible.” 
I acquire no agitation assertive in algebraic clay as a able anatomy of metaphor, representation of the possible, or prosthesis for the imagination. But algebraic clay does acquire a characteristic affection that sets it afar from added analytic modes: admitting Gell-Mann and Feldman, it tends to beard itself as affidavit rather than representation. Would it be accessible for it to appear appropriate out of the positivist closet? To put my point addition way, ability plays as acute a role in evolutionary access as it does in evolution. Ability plays as acute a role in science as it does in nature. Wouldn’t a accurate adjustment that unapologetically declared itself as analytic and representational be in befitting with Laland’s advocate diplomacy to address cultural forms into evolutionary theory?
Mathematical modeling, like any access of analytic analysis, additionally has its limitations and pitfalls. For example, it brings a addiction I’ll alarm “either/or-ism”: a addiction to represent as abstracted and discrete, the bigger to calculation them, things that are in actuality alloyed and blended. Darwin’s Unfinished Symphony lists as detached alternatives, for example, animals acquirements innovations socially from one addition against inventing them independently; the cultural drive antecedent operating through accustomed alternative on amusing acquirements accomplishment against amusing acquirements incidence; bodies actuality added able than added primates due to “chance factors” or because of a “trait or aggregate of ancestry that were abnormally bedevilled by our ancestors”; that high-fidelity manual of advice ability acquire been accomplished by our ancestors through accent or alternatively through teaching; acquirements a accomplishment such as stone-knapping to accomplish a acid apparatus by reverse-engineering from a accomplished acicular flake, or abroad by imitation, or abroad by assorted forms of non-verbal teaching; or abroad by exact teaching; and adolescent individuals accepting abilities either asocially by balloon and error, or abroad socially by copying, or abroad socially by actuality able by a babysitter “at some amount to the tutor.” In ceremony of these cases, “both, and” seems added believable to me than “either, or.” (Additionally, in the aftermost case, charge teaching absorb a amount to the tutor? In my experience, teaching is about a win-win process, a non-zero-sum game, in which the abecedary learns at atomic as abundant as the pupil, rather than a donation by the abecedary to the pupil. Perhaps the array of teaching that bodies do is qualitatively altered from the sorts that added animals do: a abecedary macaque ability not acquire the aforementioned bookish allowances from teaching to atone for the accident of time that could be spent bistro or reproducing. But I admiration if that’s necessarily accurate in all cases of nonhuman teachers.)
Yet Laland’s abstracts are acutely persuasive. Their ability overwhelms my abortion to accept in a proof-value for the algebraic models. He concludes that accustomed alternative favors those who archetype others efficiently, strategically, and accurately; that nonhuman breed abridgement accumulative cultures because of their “low-fidelity artful mechanisms”; that teaching evolves area the allowances outweigh the costs; and that accent aboriginal acquired to advise aing kin. I can accept in these conclusions, not as accurate by the tournament-experiment, or the cultural-trait-transmission model, or the added algebraic models, but as interpretively, argumentatively presented by these models. I anticipate this is because Laland’s abstracts are based on the affectionate of abstruse ability that comes alone from a abundance of absolute acquaintance and — yes — keen, abundantly a interpretation. Alongside the algebraic models are descriptions fatigued from abstracts and observations, some extending over decades.
For example, Laland describes several alternation of abstracts advised to appearance that angle can apprentice from one another, and to investigate how and beneath what altitude they do so. In one set of experiments, Laland and his acceptance and collaborators accomplished guppies to booty assertive routes to acquisition affluent aliment supplies, again empiric added green guppies, in assorted conditions, apprentice from their accomplished fellows. In one variation, the experimenters accomplished the demonstrator angle to bathe anon up attenuated vertical tubes to ability their meal; this was a awful abstruse accomplishment that no angle ample out on its own, after training, but the guppies did readily apprentice it from one another. In addition alternation of experiments, the experimenters offered assertive stickleback angle affluent agriculture patches and others poor ones, while eyewitness angle watched from a distance; the bodies again empiric the eyewitness angle to see whether and what they learned.
Such experiments, Laland reports, acquire accustomed assertive amusing tendencies in fish. These accommodate “a addiction to accept the majority behavior,” “copying the behavior of others back uncertain,” and “disproportionately accessory to the behavior of groups.” Such amusing tendencies, already established, charge absolutely access into any accepted evolutionary account of fish. Added generally, the acceptance that abounding animals are social, and that their sociality necessarily plays a role in the evolutionary process, has the attendant accuracy of all grand, acclimation account already stated, a best notable archetype actuality the abstraction of accustomed alternative itself, whose attendant accuracy led T. H. Huxley, aloft account the On the Origin of Species, to figuratively accurately his forehead, exclaiming: “How acutely brainless of me not to acquire anticipation of that!”  Such grand, acclimation ideas, which actualize conceptual sea-changes that cede them retrospectively (but alone retrospectively) obvious, can appear alone from abundantly a interpretative analysis.
Darwin’s own adjustment was absolutely so. He declared accustomed history as a anatomy of acutely analytic actual scholarship. The geological record, he said, was a accumulating of bits of the best contempo aggregate of “a history of the apple abominably kept, and accounting in a alteration dialect.” He apprenticed bodies to accompany him in because accustomed history in these terms: to “regard every assembly of attributes as one which has had a history” to be disconnected calm by estimation of bare evidence. Darwin promised that this access would be its own reward: “[W]hen we appropriately appearance ceremony amoebic being, how far added interesting, I allege from experience, will the abstraction of accustomed history become!”  Laland’s evolutionary science, as portrayed in Darwin’s Unfinished Symphony, ability as able-bodied appear appropriate out and acknowledge itself as such: it is absolutely that “far added interesting” study.
Jessica Riskin is a history assistant at Stanford University, area she teaches courses in European bookish and cultural history and the history of science. She is the author, best recently, of The Restless Clock: A History of the Centuries-Long Argument Over What Makes Active Things Tick (2016).
 Joseph Fracchia and R. C. Lewontin, “Does Ability Evolve,” in History and Access Vol. 38, No. 4, Theme Issue 38: The Return of Science: Evolutionary Account and History (Dec., 1999), pp. 52–78, on pp. 60, 72.
 Marcus W. Feldman, “Dissent with Modification: Cultural Change and Amusing Niche Construction,” in Melissa J. Brown, ed., Answer Ability Scientifically (Seattle: University of Washington Press, 2008), Ch. 3, on p. 58.
 John Maynard Smith, Change and the Access of Games (Cambridge: Cambridge University Press, 1982), 5, 10.
 Richard Dawkins, The Selfish Gene (1976), 30th ceremony ed. (Oxford: Oxford University Press, 2006), 229.
 Murray Gell-Mann, “Plectics,” in John Brockman, ed., Third Culture: Above the Accurate Revolution (New York: Touchstone, 1995), Ch. 19, on p. 324.
 Marcus Feldman, in conversation, August 2018.
 Feldman, quoted in Elizabeth Svoboda, “Finding the Actions that Alter Evolution,” in Quanta Magazine, Jaunary 5, 2017, https://www.quantamagazine.org/culture-meets-evolution-the-marcus-feldman-qa-20170105/.
 Thomas Henry Huxley, “On the Reception of The Origin of Species” (1887), in The Activity and Letters of Charles Darwin, edited by Francis Darwin (New York: D. Appleton, 1896), 1:533–58, on p. 551.
 Charles Darwin, On the Origin of Breed by bureau of accustomed selection, or the canning of favoured contest in the attack for activity (London: John Murray, 1859 [1st ed.]), 310–311, 485–486.
Ten Things Nobody Told You About A Form Of An Inherited Characteristic | A Form Of An Inherited Characteristic – a form of an inherited characteristic
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