The (ongoing) evolution of evolutionary theory

41J0nOguz-L._SY344_BO1,204,203,200_by Massimo Pigliucci

Nature magazine recently ran a “point-counterpoint” entitled “Does evolutionary theory need a rethink?” [1] Arguing for the “Yes, urgently” side were Kevin Laland, Tobias Uller, Marc Feldman, Kim Sterelny, Gerd B. Müller, Armin Moczek, Eva Jablonka, and John Odling-Smee. Arguing for the “No, all is well” thesis were Gregory A. Wray, Hopi E. Hoekstra, Douglas J. Futuyma, Richard E. Lenski, Trudy F. C. Mackay, Dolph Schluter, and Joan E. Strassmann.

That’s a good number of top notch evolutionary biologists, colleagues that I very much respect, on both sides of the aisle. My own allegiances have been made clear in a number of papers [2] and a co-edited book [3]. I have been arguing for some time now for what I consider the moderate-yes side of the debate: yes, evolutionary theory does need (and is, in fact, getting) an update, but that update is yet another expansion along the same trajectory that has moved us from the original Darwinism to the so-called neo-Darwinism to the Modern Synthesis to the (ongoing) Extended Synthesis (more on all of these in a moment). This expansion has nothing to do with hyped claims of rejection of the Darwinian tenet of natural selection, resurgence of Lamarckism, and so forth. (Cue immediate snarky commentary by the Discovery Institute.) Indeed, I have chastised more than once some of my more, shall we say, enthusiastic, colleagues (including, among the above listed authors, Eva Jablonka) for unwittingly creating a backlash among “conservatives” by talking about Lamarck and the death of Darwinism.

That said, I wasn’t going to chime in about the Nature commentaries because I had made my case plenty of times before, and also because I think I see the tide (among the all-important young practitioners of the field) moving our way, so why bother. It is interesting, and both amusing and flattering, for instance, to see that in recent years I have almost without exceptions been invited to talk about these issues by groups of graduate students around the country, but rarely by older colleagues. (I guess Planck was right, if a bit harsh, when he famously said: “A new scientific truth does not triumph by convincing its opponents and making them see the light, but rather because its opponents eventually die, and a new generation grows up that is familiar with it.”)

Why, then, am I writing this essay? Because my friend Sean Carroll (the cosmologist, not the evolutionary developmental biologist) commented on the Nature articles, and took the wrong side! [4] I was surprised, because Sean is usually fair and open minded, and rarely comments on fields that are so far from his own. I must admit I was also slightly piqued by the fact that it didn’t occur to him to check with me before publishing his essay (it’s not hubris, it’s just that I often check with him whenever I write something about quantum mechanics or cosmology, because you know, they ain’t my field).

Be that as it may, let me start with what Sean gets wrong, then zoom out to explore the broader picture so that the controversy can be seen in its proper context.

Sean states, near the beginning of his commentary: “I’m a complete novice here, so my opinion should count for almost nothing. But from reading the two arguments, I tend to side with the gradualists on this one.” At least some of the basis of his position seems to be his impression that the “yes” side has engaged in ad hominem attacks on their opponents, an example of which (from the Nature exchange) is the following:

“Too often, vital discussions descend into acrimony, with accusations of muddle or misrepresentation. Perhaps haunted by the spectre of intelligent design, evolutionary biologists wish to show a united front to those hostile to science. Some might fear that they will receive less funding and recognition if outsiders — such as physiologists or developmental biologists — flood into their field.”

I must admit (and I’m certainly biased) that this hardly sounds like ad hominem. It is a reasonable, if unwelcome, analysis of the sociology and psychology underlying the debate. It is definitely the case that evolutionary biologists are worried by the specter of ID. I’m not so sure about the bit concerning funding, though with current levels of NSF approval of research grants as low as 6% that’s not entirely out of the question either.

Moreover, though Sean couldn’t possibly know this, the other side has also engaged in quite a bit of harsh commentary. Here, for instance, is what my colleague Jerry Coyne said to Nature back in 2008, commenting on a conference that I organized in Vienna about the Extended Evolutionary Synthesis [5]:

“The whole thing about natural selection being an insufficient paradigm seems grossly overblown. There are a lot of interesting new things coming out that will change our view of evolution. But to say the modern synthesis is incomplete or fatally flawed is fatuous. … People shouldn’t suppress their differences to placate creationists, but to suggest that neo-Darwinism has reached some kind of crisis point plays into creationists’ hands.”

To think that scientific controversies are just about the science, and that personalities, social allegiances, and of course money, don’t play any significant role in them, is a bit naive, and I take it that Sean was joking when he concluded his post with: “Fortunately physicists are never like this! It can be tough to live in a world of pure reason and unadulterated rationality, but someone’s got to do it.”

Let us now move back to what the actual scientific controversy is about. To do this, I will have to provide a very brief overview of the history of evolutionary theory. Bear with me, I think it’s worth it. [6]

It all began [7], as it is well known, in 1858, when Charles Darwin and Alfred Russel Wallace presented a joint paper on evolution and natural selection at a meeting of the Linnean Society [8]. The paper was swiftly followed by Darwin’s publication, the following year, of On The Origin of Species, a bestseller that quickly eclipsed Wallace’s own contribution.

The twin foundational concepts underlying the Darwinian theory are common descent and natural selection. The idea is that all living organisms are related to each other, and that the major (though, crucially, even then, not the only) mechanism that explains their diversification through time is natural selection. Moreover, natural selection is the mechanism responsible for adaptation, i.e., the functional match between organismal characteristics and the environments in which the relevant organisms live.

The first major improvement on the original Darwinism, which came to be known as neo-Darwinism (and which to this day even practitioners of the field confuse with the later Modern Synthesis!) came at the hands of Wallace himself, together with developmental biologist August Weismann. You see, the problem is that Darwin didn’t have a mechanism of heredity handy (despite the fact that, unbeknownst to him, Mendel had published his work on pea plants in 1866, while Darwin was alive and well). So Darwin flirted with a bit of Lamarckism (inheritance of acquired characteristics), and even proposed his own half baked theory of blended inheritance. Wallace and Weismann sought to get rid of the Lamarckian stigma once and for all, which they accomplished via Weismann’s famous doctrine of the separation between somatic (i.e., non reproductive) and germ (i.e., reproductive) cell lines: even if the environment influences the makeup of somatic cells, these have no way to pass that information down to new generations, so Lamarckism is ruled out. (In reality things are much more complex, for instance because plants and a number of other organisms do not obey the Weismannian doctrine.)

We then come to the Modern Synthesis (MS), the next improvement over the original version of evolutionary theory. The MS is actually divided into two major phases, one roughly extending from 1918 to the early 1930s, the other taking place from the late ‘30s through the early ‘50s.

To simplify quite a bit, by the turn of the 20th century the neo-Darwinian theory was in trouble, so much so that historians of science refer to that period as “Darwin’s eclipse” [9]. The problem — ironically — was partly created by the rediscovery of Mendel’s work, which initially was interpreted not as the much sought after theory of heredity necessary to complement Darwin’s insights, but rather as a threat to the edifice of evolutionary theory. That’s because Darwin had insisted that evolutionary change is always gradual (despite warnings to the contrary by his friend and champion, Thomas H. Huxley), while “Mendelism” seemed to show that the genetic material is inherited as a number of discrete units (the genes, whose chemical basis was, of course, still unknown). Moreover, the early geneticists had begun working on mutations, especially in the fruit fly, which seemed to bring more trouble for Darwin, since all known mutations appeared to produce radical changes in the phenotype (the appearance or behavior of organisms), again contradicting the idea of gradualism. The combination of Mendelism and mutationism, together with the resistance to Darwinian ideas by paleontologists (who saw long term trends in the fossil record, which were thought to be independent of the vagaries of environmental change), produced the widespread feeling that scientists were on the verge of dealing a death blow to neo-Darwinism.

A small group of brilliant mathematical biologists came to the rescue, chiefly Ronald Fisher, followed by J.B.S. Haldane and Sewall Wright. Fisher was almost single-handedly responsible for the reconciliation (i.e., the “synthesis”) between Mendelism, Mutationism and (neo)Darwinism: he showed how, assuming (as it turned out to be the case) that phenotypic characteristics are influenced by a relatively large number of genes, the statistical effects of those genes (and their interactions with the environment) creates precisely the sort of continuous distribution of characteristics that Darwin thought would allow natural selection to work. Indeed, Fisher formalized the Darwinian insight in mathematical form, arriving at what still today is a cardinal idea in population genetics, his Fundamental Theorem of Natural Selection [10].

The second, and equally crucial, phase of the MS was brought about by a more diverse group of biologists, most prominent among whom were Theodosius Dobzhansky, Julian Huxley, Ernst Mayr, George G. Simpson, and G. Leydard Stebbins. Indeed, it was Huxley (grandson of Thomas Henry) who coined the term “Modern Synthesis.”

What did the MS-phase 2 consist of? Of the application of the new principles of statistical genetics, cast in a neo-Darwinian fashion, to population genetics (Dobzhansky), natural history (Mayr), paleontology (Simpson), and botany (Stebbins). It was a synthesis in the sense (different from MS-1) that it broadened the scope of what could by then fairly be called the Darwin-Fisher theory of evolution to a good number of (but, crucially, not all!) other domains in biology.

The Modern Synthesis is, by and large, what is taught today in undergraduate and graduate textbooks, it represents the Standard Model of the biological sciences. But, just as the Standard Model in physics is known to be incomplete and there has been (pace Sean’s joke) quite a bit of controversy among physicists about how and when it is going to be replaced, so to has the Modern Synthesis been under increasingly severe stress, dating back from the 1960s and ‘70s, but much more intensely so in the ‘90s to the present.

Whence the stress, despite assurances by the “no” side that “all is well”? Here is a quick run down of the major points:

  • Developmental biology was famously left out of the MS, and needed to be brought in. Systematic attempts at doing so have resulted in the development of an entirely new field of research, colloquially known as “evo-devo” [11], which has been developing its own agenda and principles.
  • Ecology was also only marginal to the MS, and some recently developed ecological principles, such as niche construction theory, need to find an organic place in our way of thinking about evolution [12].
  • The possibility that selection acts at multiple levels, rather than just on organisms (Darwin) or genes (modern population genetics) has been raised several times. After a couple of false starts under the label of “group selection,” there is now a well developed multi-level theory of selection and its action [13].
  • Eldredge and Gould’s idea of punctuated equilibria [14], which maintains that some lineages evolve (geologically) very quickly and then remain unchanged through long periods of stasis.
  • Evolutionary genomics has yielded not only terabytes of new data on the genetic makeup of biological species, but new insights into the evolution of regulatory vs structural (“house keeping”) genes, and has especially highlighted the necessity of thinking in terms of gene networks and their interactions with the environment, rather than single genes with the environment treated as background information [15].
  • Much research has been devoted to a widespread phenomenon known as phenotypic plasticity, which allows us to model the interaction between genomes and environments in novel ways. Plasticity had been discovered in the early part of the 20th century, but the field has come into its own only in the last couple of decades [16]. A number of authors, chiefly Mary Jane West-Eberhard [17], have argued that plasticity represents a novel type of evolutionary mechanism, much under appreciated until recently.
  • A trio of interrelated new concepts has been proposed, and has begun to be investigated empirically, to further expand the known set of evolutionary mechanisms: evolvability, robustness, and modularity [18].
  • There has been increasing evidence that there exist a panoply of epigenetic (i.e., non-genetic) inheritance systems that are responsive to environmental stresses (though definitely not in a Lamarckian fashion!), that interact with the known genetic inheritance system, and that may play a role at the least in short-term evolutionary responses [19].
  • A degree of biological complexity can be obtained independently of natural selection. This has been known since D’Arcy Thompson’s classic study on “growth and form” [20], but has been elaborated theoretically and investigated empirically under the name of “facilitated variation” [21]. This is related to, but further builds on, Gould’s old idea [21] that so-called developmental constraints can actually play a positive role in evolutionary change.
  • There is increasing evidence — if you ask paleontologists — that macro-evolutionary phenomena (i.e., changes above the species level) are partially causally decoupled from micro-evolutionary ones (i.e., within species) [23]. This has nothing whatsoever to do with the specious arguments proposed by creationists, and everything to do with species selection and group-level properties.

I may be missing something, but you get the idea. Now, you could say, as the “no” camp maintains, that all of the above is just cherries on the cake of the Modern Synthesis. I, and a number of others, beg to differ. The above represents a huge set of empirical discoveries and theoretical advances that go well beyond, and are certainly not implied by (and yet do not contradict) the Modern Synthesis. This is stuff that needs to be further explored, better articulated, and explicitly integrated into the general framework of evolutionary theory — work, I might add, that has been well under way for years.

The eminently sensible idea behind a push for the recognition of an Extended Synthesis in evolutionary biology, then, is that the above constitutes at the very least the same amount of conceptual novelty in the biological sciences that was represented by the universally acknowledged break between neo-Darwinism and the Modern Synthesis, and certainly much more than the one separating the original Darwinism from the Wallace-Weismann version of the theory.

In the end, of course, it doesn’t matter what we call it. Phenotypic plasticity, evolvability, epigenetics, niche construction, facilitated variation and all the rest are here to stay. But, we do usually label different versions of scientific theories with different names, and for good reasons. They mark significant advances in our understanding of the world, and of course recognize the work that went into making those advances, as well as the people who did that work. There certainly is no need for antagonism, on either side of the divide, we can and should all work together to further biological research. But it is hard to see what could possibly justify — given all of the above and much, much more — this recalcitrance to recognize that biology is entering a new phase of its history. It’s a very exciting phase, and one that will, thankfully, soon be in the hands of todays’ graduate students and young researchers.

_____

Massimo Pigliucci is a biologist and philosopher at the City University of New York. His main interests are in the philosophy of science and pseudoscience. He is the editor-in-chief of Scientia Salon, and his latest book (co-edited with Maarten Boudry) is Philosophy of Pseudoscience: Reconsidering the Demarcation Problem (Chicago Press).

[1] Does evolutionary theory need a rethink?, Nature, 8 October 2014.

[2] An Extended Synthesis for Evolutionary Biology, by M. Pigliucci, Annals of the New York Academy of Science 1168:218-228 (2009). / The extended (evolutionary) synthesis debate: where science meets philosophy, by M. Pigliucci and L. Finkelman, BioScience (2014).

[3] Evolution – the Extended Synthesis, by M. Pigliucci and G.B. Müller, eds. (2010), MIT Press.

[4] The Evolution of Evolution: Gradualism, or Punctuated Equilibrium?, by S. Carroll, Preposterous Universe, 10 October 2014.

[5] Biological theory: Postmodern evolution?, by John Whitfield, Nature, 3 October 2008.

[6] A more in depth treatment of the history of evolutionary theory from a philosophical perspective can be found in: Biology’s last paradigm shift. The transition from natural theology to Darwinism, by M. Pigliucci, Paradigmi 2012 (3):45-58 (2012).

[7] It actually began much earlier: for a fascinating discussion of pre-Darwinian concepts of evolution, and how the whole field was actually considered pseudoscientific (!), see Michael Ruse’s “From pseudoscience to popular science, from popular science to professional science,” in Philosophy of Pseudoscience: Reconsidering the Demarcation Problem, by M. Pigliucci and M. Boudry, eds. (2013), Chicago Press.

[8] The 1858 Darwin-Wallace paper.

[9] The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900, by P.J. Bowler (1992), Johns Hopkins University Press.

[10] Fisher’s fundamental theorem of natural selection, Wiki entry.

[11] Evo-devo, by the Understanding Evolution team.

[12] Niche construction, an overview.

[13] Evolution and the Levels of Selection, by S. Okasha (2009), Oxford University Press.

[14] Punctuated equilibrium, Wiki entry.

[15] Here is an example of what evolutionary genomics labs do.

[16] Rather immodestly, I can say that I wrote the book on phenotypic plasticity a few years ago: Phenotypic Plasticity: Beyond Nature and Nurture, by M. Pigliucci (2001), Johns Hopkins University Press.

[17] Developmental Plasticity and Evolution, by M.J. West-Eberhard (2003), Oxford University Press.

[18] Is evolvability evolvable?, by M. Pigliucci, Nature Reviews Genetics 9:75-82 (2008). / Robustness and Evolvability in Living Systems, by A. Wagner (2007), Princeton University Press.

[19] What Role Does Heritable Epigenetic Variation Play in Phenotypic Evolution?, by C. Richards, O. Bossdorf, and M. Pigliucci, BioScience 60 (3):232-237 (2010). / Transgenerational epigenetics, Wiki entry.

[20] On Growth and Form, by D.W. Thompson (1945), Cambridge University Press.

[21] The theory of facilitated variation, by J. Gerhart and M. Kirschner, Proceedings of the National Academy of Sciences, 104:8582-8589 (2007).

[22] Ontogeny and Phylogeny, by S.J. Gould (1985), Belknap Press.

[23] See, for instance: Mass extinctions and macroevolution, by D. Jablonski, Paleobiology 3:192-210 (2005).

69 thoughts on “The (ongoing) evolution of evolutionary theory

  1. Thank you, thank you. Your summaries of the sequential advances give me a framework for understanding the biological vocabulary that I did not have before, and I hope I can fit new knowledge in and around the categories. As a physicist, I cannot help but think that the situation mimics (a little bit) the post-Newtonian period in math/physics in the late 1700s and early 1800s. Darwin plays the part of Newton, laying down the basic laws, then Laplace, LaGrange, Hamilton, Jacobi, Euler, and many, many more extend Newton’s work to model more complex and general systems. It will be interesting to see if the field of biology is headed toward its own “quantum revolution” which might discover the limits of Darwin’s basic ideas, but that is a question for the future.

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  2. Hi Massimo,
    Good to see some more science on SS! When I (as a non-biologist) read those articles as they came out in Nature I found myself more persuaded by the “no” article. Has yours now persuaded me to the “yes” camp? Well, not really, indeed your article could be read as support just as much for the “no” camp.

    First, we all recognise that the scientific reward system is all about novelty, and thus all scientists have an incentive to hype the extent to which they are over-turning old orthodoxies. You say:

    The above represents a huge set of empirical discoveries and theoretical advances that go well beyond, and are certainly not implied by (and yet do not contradict) the Modern Synthesis. This is stuff that needs to be further explored, better articulated, and explicitly integrated into the general framework of evolutionary theory — work, I might add, that has been well under way for years.

    I (as an interested non-biologist) am persuaded of all of that. A crucial phrase is “and yet do not contradict” the Modern Synthesis. When many of the items you list were uncovered, they were often accompanied by “and this overturns the Modern Synthesis” hype. Or, alteratively, by the claim that the new ideas are more important, having a bigger effect on evolution overall, than the older ones.

    As I see it, the “no” camp are saying, yes those new ideas are interesting and important and need to be synthesized into the overall picture, but they amount to perturbations on the basic neo-Darwinism Modern Synthesis. Thus they add to, rather than replacing, the older ideas.

    The “yes” camp then seems to be saying that the combined effect of all the items you list is overall more important for evolution than the ideas of the Modern Synthesis, which is thus relegated to semi-irrelevance. The “yes” camp article did not persuade me of that, whereas the “no” camp do seem happy to assimilate the new ideas, where appropriate and where supported by evidence that they have significant effects in nature (rather than being just theoretical possibilities).

    I could also be convinced that both sides are mostly agreed on the actual science, and that the kerfuffle is merely over-hyping on the one hand and reaction to that over-hyping on the other.

    PS “Planck” not “Plank”.
    PPS Yes, Sean Carroll’s last line was indeed an ironic joke!

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  3. This is an interesting topic in general (how do scientific models advance) and specifically for biology. I’m somewhat inclined to agree with Coel here in that I don’t see the transition being as strong if the extended synthesis is not contradicting the modern synthesis, why not just consider it the natural progression of the modern synthesis? What advantage is gained from labeling the new research something different?

    One explanation could be is that the models do contradict when we look at the details. Having read some debates from professional biologists online about group selection, it doesn’t seem like the old guard is embracing some aspects such as group selection as strongly as say David Sloan Wilson and others. However, I’m not knowledgeable enough in evolutionary biology to know if one or the other side is exaggerating their claims. Massimo perhaps you can clarify that point.

    The other explanation could be is that if even if it’s not a major change, research areas covered under the extended synthesis get marginalized in terms of funding and prestige, which may be a good reason to promote the term extended synthesis. This maybe more sociology of science but some of those advances can be important if they move the research forward.

    I’ve personally become more conservative on claiming new models in science as I’m in the field of psychology, where every few years someone claims to have the absolute new model and it’s usually the same old hat with a few added bells and whistles. That doesn’t seem to be the case here but does make me skeptical when I hear of similar suggestions in other fields.

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  4. Thanks for the useful account of the history and state of evolutionary theory. But on the name question…

    “In the end, of course, it doesn’t matter what we call it.”

    Good. We agree this is just a matter of terminology, and there’s no substantive question at stake, except perhaps a question of degree about just how much evolutionary theory has changed. That question is a very fuzzy matter of judgement, though that’s not to say it isn’t a judgement worth making.

    “But, we do usually label different versions of scientific theories with different names, and for good reasons.”

    We adopt new names when there’s a specific identifiable difference. But when it’s lots of differences adding up to a matter of degree? That seems confusing to me. We’re not talking here about two distinct versions of a theory. We’re talking about a theory which has developed continuously over time, and you’re not identifying any specific transition. If you define the term “Extended Synthesis” to refer just to a snapshot at an arbitrary time–evolutionary theory as it stands right now–then the term will soon come to refer to an old version of evolutionary theory, not the latest version.

    My impression is that the adoption of the terms “Neo-Darwinism” and “Modern Synthesis” was not as totally arbitrary as this, but corresponded to a somewhat specific (if somewhat fuzzy) transition. And whether that’s so or not, the names have turned out to be something of a source of confusion, because of their non-specific names. Every theory is new and modern when it’s first invented! I suppose “Synthesis” meant something in the “Modern Synthesis”. But “Extended” isn’t very helpful. Will Evolutionary Theory 2024 be called the “Further Extended Synthesis”?

    Why can’t we just talk about “current evolutionary theory”? If we want to make the point that evolutionary theory has changed enormously over recent years, we can just say that in so many words. We shouldn’t adopt confusing language just for the sake of making a point.

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  5. Where would the Baldwin effect fit within your list of on-going directions? I am tempted to push it under phenotypic plasticity, but I am not sure if that is the most natural place. It also seems like a tension that has been around for a long time; since 1902 for Baldwin, and revived in 1953 by G.G. Simpson. It has survived all the transformations since the birth neo-Darwinism and yet still feels like a piece that doesn’t fit.

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  6. Hmm. Color me somewhat bemused. On one hand I thoroughly agree that there have been revolutionary developments in evolutionary theory over the past few years. The discovery that noncoding regions of DNA play at least as important a role in evolution as coding regions is particularly important, and really wasn’t anticipated by the Modern Synthesis. The so-called “Red Queen” (the realization that evolution usually occurs against a rapidly changing background) is also a major shift. But the problem is that I’m unable to see anything in the “extended synthesis” that actually looks like a global synthesis. It looks more like a program of research into developments which will eventually produce a new synthesis.

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  7. I don’t see this so much as topics to include as to the roles entities play. One common perception is that organisms are at the whim of the environment, they are the objects that selection acts on. I think one reason there is so much pushback from conservative religion is that they simply believe scientists conceived natural selection as an attempt to replace their God. When in reality organisms and the environment are in a feedback system – changing each other. Natural selection is not all powerful. The relative recent idea of ecosystem engineer is a case in point, but it not restricted to a few groups – every organism/population just by interacting with the environment changes the environment and themselves. We can see some of the same themes in the resistance to climate change – the belief that the environment is off-limits to change by mere organisms – only God can control the environment.

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  8. Without continuous attention and improvement (reality checks) the theatre of ideas risks becoming a worn out parody of itself. To let this happen is a kind of hubris – because it implies that there is nothing more to be learned, no refinements could possibly be introduced. It defies the principle of all science – that every hypothesis – every causal explanation – is provisional.

    That does not mean that the work of Dobzhansky, Mayr, G. Gaylord Simpson – as well as Haldane and Fisher and others on the modern synthesis are no longer material covered in our physical anthropology 101 classes, as they were 40 years ago, at the University of Toronto. (I have well thumbed and coffee stained copies of their major papers and books cluttering my office even today, for they are those things that no one in their right mind would send off to recycling EVER, not even with the ease of access in the digital age. As for copies of the Charles Darwin the Origin of Species, I have at least three different editions, one in all rooms where I regularly sit and work.)

    Our comprehension the rich inter-digitation of living forms; the complexity of each species’ plastic responses to larger ecosystems, is still unfolding. Taking the depth of developmental plasticity, including epigenetic phenomena, and the regulatory functions of the ecology, into which each life form is sunk, into account, is normal science, surely?

    From there, it is possible to venture towards an understanding of learned behavioural suites. The realization, that these can be manifestations of a kind of plasticity that goes beyond strictly genetic replication, is not to break with standard evolutionary models. Indeed, it is not to put forward ideas about plasticity that are in any causal sense antievolutionary or nonbiological, but rather to add depth and rigour to these hypotheses – we can move closer to an understanding of each life form as subject to natural selection via multiple tiers of potential resiliency and constraints.

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  9. Hi Massimo, nice essay with some good historical overview. I had read Sean Carroll’s piece earlier and did not feel put out by his conclusion, especially given the caveats he’d made. From an outsider’s perspective it may not seem like enough changes have occurred to merit changing the name on the billboard and so appear to be more about in-fighting.

    My own perspective is that a change is due. The overall picture of evolution is significantly different, given the changes in our understanding of biology at so many different levels. Punctuated Equilibrium alone is a pretty important change in perspective regarding rates of change.

    To add to the list of new ideas above, there was Margulis’s theory of endosymbiosis which radically adds to methods of evolution and was so outside the framework that her struggle to get it recognized was formidable. More recently the discovery of “jumping genes” or “transposons” show how viral interactions may have (and may continue) to alter the genomes of species.

    This is no longer your grand dad’s evolutionary theory (caveat: unless your family multiplies very quickly).

    That said, I’m not certain if I’d qualify a change in name as an “urgent” need. That word drives the question, urgent for what? If it is not done immediately what will happen? The honest answer is, nothing, really. So why then is it urgent?

    Your arguments here explain why it makes sense, but not so much the urgency. You rightly question the “recalcitrance” of those not wanting to do it, but do you have specific arguments why it should be done sooner rather than later (when more of these new ideas get better fleshed out)?

    The only reasons I can give are that it would be useful (to recognize the degree of changes to those within or entering the field) and that it would be exciting. It would build interest and enthusiasm in considering what our current model really is. I have to admit that doesn’t raise to the level of a crisis. My only point then is why do we have to wait for a crisis to do something exciting if it is in fact relevant?

    Richardwein makes a valid point that we could stop naming each new iteration and just talk about “current evolutionary theory”, noting the changes as they come. But that itself is a change from what we have now and would stand against the “no” group, who are arguing that we are still working under a specific model (the modern synthesis) without significantly relevant changes. What would you feel about having an open/generic name with planks added as they come in?

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  10. I enjoyed and appreciated the write-up of the history of evolutionary theory, as well as the current state of the art, so I think this was a great article, Massimo.

    However I agree with the points made by Coel, imzasirf and Richard Wein. I can’t get very exercised about what we call the theory of evolution, but am inclined to side with the gradualists. The core idea Darwin had seems to stand, and we are only refining and adding to it. I don’t see any significant paradigm shifts happening, as happened when Einstein overturned Newtonian mechanics (though Newtonian mechanics still work in everyday scenarios as an approximation to relativity, they are fundamentally flawed).

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  11. Hi DM-

    In one sense isn’t there also a parallel conceptually between Newton mechanics/relativity and the new emphasis on updates to evolution theory. Relativity unified previously separate concepts (space, time, gravity), the background was no longer a separate static thing. In some sense to me it seems like a lot of the updates Massimo outlined recognize a greater emphasis on plasticity in the feedback between levels (genes, organisms, groups, larger environmental ecologies ). We don’t have a single unifying systemic theory that successfully unifies all of this, but it does seem the progress taking place involves small unifications. If my laypersons understanding is correct I do think some formal acknowledgement of this trend in emphasis might be productive.

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  12. Hi DM,

    “The core idea Darwin had seems to stand, and we are only refining and adding to it.”

    That is not exactly true, and I’ll explain why using the three ideas I mentioned in my reply. The core idea of Darwin was gradual change with forces making selections on changes organic to the organisms themselves.

    Punctuated Equilibrium recognizes that change is not, or at least is not required to be gradual. Changing conditions can lead to abrupt changes in species.

    Endosymbiosis recognizes that it is not merely forces working on singular organisms (through selection) that creates evolutionary changes. That eukaryotes likely emerged from extended symbiotic (cooperative and internal) behavior between prokaryotes means that one of the most profound historical evolutionary events did not involve the main evolutionary mechanism endorsed by Darwinian theory.

    Finally, with transposons we see that a mechanism for change can come from direct interaction between viruses and the genetic code of a species, with possible lingering effects from then on down the line.

    I won’t go into the other points Massimo mentioned (as I think he will) but I don’t see how those three are “only refining and adding to” the core of Darwinian theory. Or maybe I should put it another way, if that is all these did, then I don’t see how any of the other changes merited identification with a name change. These seem just as, if not more significant.

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  13. I find Massimo’s list of ten major points lacking some very important topics, like:

    1. Transponsable elements (TE) are mobile repetitive sequences that make up large fractions of mammalian genomes, including at least 45% of the human genome. They were first described by Barbara McClintock already in the late 1940s but her results were so outside mainstream biology that were ignored quite some time (eventuelly she got Nobel Prize for this discovery). Intriguing species of TE are endogenous retroviruses, which is virus’s genetoc material incorporated in host genome (like human’s). Comprehensive overview is in book Evolution: A View from 21st Century.

    2. Talking about viruses: virology is in recent years in a kind of renaissance due to discovery of giant viruses, comparable in size and genetic material to small bacteria (see for example Hints of Life’s Start Found in a Giant Virus in Quanta Magazine). There are many fascinated proposals: after Carl Woese incorporated Archaea domain in phylogenetic Tree of Life (besides Bacteria and Eucarya), viruses could become the fourth domain. Of course, first official line about their non-living status should be changed. This is still minority position, but it’s not difficult to see viruses as live entities: they have dormant phase as virions and active phase, when they engage/hijack sell’s replicative machinery. Then there is big gene pool in viruses which is unique to them and then there is possibility (the most speculative) they were active participants in RNA world. Not to forget to mention Sputnik: other viruses attacking virus.

    3. Modern Synthesis talks about natural selection that acts on novelties, but it let unspecified how this new features arise. In theory should be small modification, but in reality that’s not always the case. Well known example is hemoglobin, of which they are many types and they were made by duplication event, not minor modification. (nice exposition is in American Scientis, no. 2, year 1999: The Evolution of Hemoglobin). Look from different angle is recent book by Andreas Wagner Arrival of the Fiitest.

    All in all, my opinion is this might be Kuhnian moment for biology (Thomas Kuhn is being criticized for building his case [paradigm shifts] mostly on history of physics).

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  14. How on earth are the simpler minded among us ever going to understand and accept a theory of evolution with so many moving parts? Most folks want things simple, certain and static (aka creationism) or GTFO! I would say that issue is far more pressing than polishing the science. Hence the quarrel I suppose…

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  15. Darwinism is Lamarckism, according to Wallace, and not denied by Darwin (to Darwin’s daughter’s reprobation).

    Evolution by artificial selection was known for millennia, and practiced for tens of millennia (the oldest dog known is Belgian, and around 35,000 year old).

    Fossils were known in Ancient Greece. They caused confusion. To remedy this, Aristotle (PBUH), sent his students to study and report on life forms, thus founding, de facto, biology. (That the universe was not in a steady state was shown by the landing of a giant meteorite in northern Greece; it was visited for centuries.)

    By 1766, after proposing that the Solar System had been accreted from a cloud of debris, Buffon proposed that animals changed: they evolved. African and Asian elephants had evolved from Siberian mammoths, due to migrations changing their environment, he claimed. The details are unimportant: the evolutionary horse is out of the barn.

    The full blown theory of evolution was proposed by Lamarck. This was a great conceptual breakthrough.

    Lamarck’s book, “Philosophie Zoologique” was published in 1809. The year Darwin was born.

    Lamarck, employed as one of the world’s first research professors, demonstrated both the immense age of the Earth, and natural evolution, by studying fossilized mollusks (while coining the word “biology”).

    Then Lamarck suggested that the way animals lived could directly affect their genetics. A scientifically confirmed way to get this effect is now called “epigenetics” (“above genetics”).

    Lamarck got hated for all this by the forces of Christianity. The idea that the living creature, could, by the way it lived, CREATE its own features was revolting to those who promoted the Christian god. Lamarck was outlawed in British (religious) universities (Oxford, Cambridge, etc.)

    Lamarck proposed that the long necks of giraffes evolved as generations of giraffes reached for ever higher leaves. The Church made fun of that (some still do, following the Church!).

    But Lamarck did not just propose that evolutionary was driven by behavior. Following Buffon, Lamarck believed life started with spontaneous creation (the present view). Lamarck proposed that, insensibly, each baby was more complex than the preceding baby, so evolution would be characterized by an increase in complexity (as it indeed is).
    Lamarck suggested birds descended from reptiles.
    Lamarck suggested that there are flying squirrels, because squirrels tried to fly for generations (natural selection does not contradict this).

    After 1815, reaction came over Europe. Jews got discriminated against by the Middle European dictatorship (they could not be doctors, lawyers, etc.). Lamarck, being an enemy of god, was made destitute.

    Lamarck being French, some feel more appropriate to attribute the discovery of evolution to the English Darwin, who, besides, was the heir of a financier, and not a vulgar professor, like Lamarck.

    Ideas are hard, especially when revolutionary. Parodying Lamarck’s ideas the way the Church did means that no meta-lesson was learned. Those who introduce the greatest new ideas, like evolution, deserve the greatest respect. Not showing respect for geniuses such as Lamarck is not to show respect for what makes civilization advance.

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  16. Thanks for your “huge set of empirical discoveries and theoretical advances that go well beyond, and are certainly not implied by (and yet do not contradict) the Modern Synthesis.”

    I am trying to get a grip on what all the fuss is about extended, expanded, and new syntheses and this helps enormously!

    There are many issues I don’t yet understand and I’d like to seize the opportunity to ask for your views.

    * In your overview of the development of evolutionary theory you do not mention the early attempts to work out the implications of Mendelian genetics for evolutionary theory and the development of population genetics by Bateson, Hardy, Weinberg, Morgan, Johannsen, Nilsson-Ehle, Tammes, and Punnett in 1902–1916. It seems to me that this work (including such things as Johanssen’s distinction between genotype and phenotype, Morgan’s origin-fixation model, and Bateson, Johannssen, Tammes and Nillsson-Ehle’s development of a probabilistic theory of the evolution of quantitative characteristics) is much more important than Weismann’s germ-plasm theory (which applies only to a very limited set of organisms). Do you agree?

    * Your set of important developments doesn’t include molecular evolution, genetic drift, the neutral theory and the re-introduction of the origin-fixation model in the 1960s. Don’t these developments contradict certain tendencies of the modern synthesis? If so isn’t this a much more revolutionary development than the ones you do mention (after all, you say that the latter “do not contradict the Modern Synthesis”)?

    * You do mention punctuated equilibrium and macro-evolution. Don’t these ideas contradict the view of the modern synthesis that macroevolution is just microevolution over millions of years?

    * According to S.J. Gould the modern synthesis was already “effectively dead” in 1980. Was he correct? If so, why all the fuss about extending the synthesis? Isn’t the no-side defending a dead horse, while the yes-side tries to kill a horse that is already dead?

    * It seems to me that there is a difference between on the one hand applying a theory to new area’s (as happened in what you call the second phase of the synthesis) and extending it to more complex cases and on the other hand modifying a theory. The ‘yesies’ clearly think that evo-devo and niche construction are of the second kind. Do you agree with them? If so doesn’t that contradict your statement dat the new developments do not contradict the modern synthesis?

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  17. Massimo,

    Always a pleasure to read and I agree with your assessment.

    It seems to me the ‘no side’ in the Nature article is not really against change, they acknowledge the yes sides points but seem worried any change would be “diluting” or would “de-emphasize the most powerfully predictive, broadly applicable and empirically validated component of evolutionary theory”, or that changes would be like writing in “upper case” and giving “special advocacy” to more recent developments in evo theory. They also seem to feel uncomfortable with the level of some of the evidence on some of the more or less recent advancements in evo theory.

    At the end of the article they invite “Laland and colleagues to join us in a more expansive extension” to evo theory, so I would like to be positive and assume they are sincere, their fears can be easily addressed, and things can move forward.

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  18. Reviewing the major points,I see neutral mutation is omitted. It seems without a genuine acceptance of this, the importance of genetic drift is minimized. In popularizations, it may be entirely omitted. Not so long ago, Carl Zimmer, a very highly respected science writer commonly praised for skill in presenting evolution, wrote “Surely, if my blood type has endured for millions of years, it must be providing me with some obvious biological benefit.” If this were true, he wouldn’t have had an article to write (http://carlzimmer.com/articles/index.php?subaction=showfull&id=1406307212&archive=&start_from=&ucat=17&). The possibility that blood groups rose more or less by accident, a more or less neutral variation that natural selection could not distinguish, much less erase, was not even deemed worthy of discussion. And the obvious utility of blood cells was assumed to be irrelevant to the issue. What is it that makes even such a highly respected writer as Zimmer make what appears to a layman to appear to be a gross lapse in elementary judgment?

    It seems to me to be a consequence of the emphasis on gene selection. So far as I can tell, gene selectionists tend to state that the gene is the unit of heredity, which no one I know of has denied, then leave it at that. Indeed, it is not common for gene selectionists to clearly state what’s wrong with seeing the organism as the primary unit of selection. But the real problem seems to be the logical conclusion that, genes being the units of selection, reproduction of alleles is prima facie proof they have been selected. Even more insidiously, the mere existence of alleles being selected means they simply must have causal efficacy in the consitution, development and behavior of the organism. I’ve seen formal acknowledgement of the discoveries of neutral mutation and even genetic drift, but in practice? In practice we seem to get Mr. Zimmer’s “surely…” I don’t think it means much to deny believing in “genetic determinism,” then try to explain pretty much everything by gene selection. I think that’s why most of the points listed in the OP are evidence against the de facto genetic determinism, or against an exclusive fixation on gene selection.

    If you look at the popular presentation of evolutionary theory, you see evolutionary psychology, and practically nothing else. The ardent defenders of the overwhelming importance of gene selection and the power of natural selection to make all traits adaptive seem to be totally committed to this, seeing EP =Modern Synthesis. A disagreement with the one is an attack on the other. Well, it is not a popular opinion, but I believe studies of society can be conducted on a scientific basis as well. Frankly, as near as I can tell, what the EP/standard Modern Synthesis people sneer at as the Standard Social Science Model (SSSM)…and yes, some do write it that way!…is in fact the gold standard. This is far too complex to go into into detail. But I would suggest that it is a fairly fundamental and intuitive premise that there is only one nature, so although we find it practical to divide our investigations into more manageable subjects, the unity of nature means that the premises and conclusions of each meet each other, in the real world so to speak.

    Personally I would add that there is an awful lot of confusion about the question of when natural selection acts to conserve an organism’s morphology as opposed to when it serves as a driver for the emergence of evolutionary novelties. I should think an extended Modern Synthesis needs to highlight the centrality of answering this question for individual studies. On a terminological level, I wish either Modern Synthesis, today’s or the putative Extended, would sharpen its vocabulary so that it was much easier to tell when we are talking about the change of allele frequencies in populations, or about the multiplication of species.

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  19. I assume that part of the animus on the “no”-side comes from the feeling of having to deal at least partly with people who go for hyperbole in the hope of being recognised as the next Mayr or Dobzhansky – without themselves necessarily contributing anything beyond that hyperbole to the body of knowledge. Seriously, why not just do some practical research and then let our descendants in fifty years decide if the early 21st century saw the rise of something that turned out to be as important a break as the Modern Synthesis? The “end of history” guy was also a bit premature in his declaration.

    And yes, I think some of the individual points are seriously overblown. Epigenetics is really interesting but as long as it doesn’t change the DNA sequence it is irrelevant in the long run; and if it does, it is just standard mutation. I have never understood what is supposed to be so special about punctuated equilibria either. Unless some serious saltationism and hopeful monsterism were surprisingly found to be true (cf. the Creationist caricature of the Crocoduck), this is just normal Modern Synthesis evolution with genetic bottlenecks and stabilising selection. Gould himself pointed out that people misinterpreted it because they didn’t think on a geological scale.

    And macroevolution =/= microevolution? That seems to boil down to the observation that mass extinctions have a strong stochastic element and entail very unusual one-off selection events. Do we really need to slap a new label on the theory of evolution because we realised that “being restricted to North America while that continent is hit by a meteorite” and “fortuitously having an unusually efficient respiratory system when encountering a once-in-100-million-years oxygen crisis” are not character traits under the same kind of selection as wing span or leaf surface area?

    Etc.

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  20. Hey, Massimo… I agree with your assessment: “but that update is yet another expansion along the same trajectory that has moved us from the original Darwinism to the so-called neo-Darwinism to the Modern Synthesis to the (ongoing) Extended Synthesis”. It does seem that sciences have never been static monoliths, and they’ve always been revisionist from the time of Bacon on. I’ll agree that as you state, whether it is called by this specific term or not doesn’t matter, what matters is the inclusive acceptance of these other various sciences and conceptual tools within the ongoing Modern Synthesis. It’s almost as if we’re back to bias and category mistakes. It’s not an either/or situation, more of an both/and inclusion of new data sets and categories left out of the MS. In that sense your path seems the better of the two both philosophically and scientifically. Almost a Modern Synthesis + … additions to the existing synthesis: which from what you’ve mentioned would need a revamp of the mathematical formula underpinning the original MS.

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  21. Massimo: “I have been arguing for some time now for what I consider the moderate-yes side of the debate: yes, … as the “no” camp maintains, that all of the above is just cherries on the cake of the Modern Synthesis. I, and a number of others, beg to differ.”

    Amen! Go, Massimo Go!

    With the 10 bullet points, if the ‘no’ group still says no, we can just let Planck’s proverb take its course.

    Yet, I would like to discuss this issue from a different perspective.

    I am a diehard physics-ism-ist. That is, every (every, …, and each) thing (matter, non-matter, psychic or spiritual, etc.) in ‘this’ universe is ‘based’ totally on the bottom of the physics-laws. Of course, every ‘bit’ of human is just an ‘expression’ of those physics laws. The two most obvious novelties of human are ‘intelligence and consciousness’, and they must be evolved as ‘expressions’ tier by tier to reach the ‘human state’. That is, they can be written as a ‘set’ of function equation. Let F as the ‘function’ of intelligence, then,
    F (human) = intelligence (human) = {math, poetry, … air plane, iphone, etc.}

    F (dog) = intelligence (dog) = {… best friend of man}

    F (virus) = intelligence (virus) = Ebola (let the human intelligence sh- its pants just recently)

    What is intelligence? I wrote a long article for it, at http://www.prebabel.info/aintel.htm . For here, one short criterion should be enough. All (any) intelligences must rely on a counting device (wherever it sits) as its ‘information-processor’. For the diehard physics-ism, that computing device must sit at the bottom of the physics, such as in ‘proton or neutron’. That is,

    F (proton) = intelligence (proton) = {at least having a counting device, such as Turing Computer)

    If ‘proton intelligence’ is not the case, then all the talk above is simply nonsense. Obviously, this is a bit long story, needs 500 word space of its own to show that it is not nonsense.

    However, this issue can be viewed in a more traditional ways.

    One, semantically:
    ‘Selection’ is selecting from some ‘existing’ contestants, that is, nothing new (novelty) comes out after the selection. Selection is ‘external’.

    ‘Adaptation’ is facing off a challenge by something new, different from the old school. This ‘new’ can be never used packet change or newly ‘acquired’ skill or matter. Adaptation is powered ‘internally’.

    By all means, selection, sieving and filtering is not adaptation.

    Two, from facts:
    A. No single life or even a single species can evolve ‘alone’. It needs a platform which is definitely not constructed with Darwin-mechanism (DM).

    B. Genetics is a well-defined internal ‘dynamics’ while the external ‘selection’ is at best acting as a ‘boundary condition’ for this dynamics.

    C. Etc.

    I have showed some these at https://scientiasalon.wordpress.com/2014/10/28/the-varieties-of-denialism/comment-page-1/#comment-9225 and https://scientiasalon.wordpress.com/2014/11/05/back-to-square-one-toward-a-post-intentional-future/comment-page-3/#comment-9503 .

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  22. “[W]hat could possibly justify … this recalcitrance to recognize that biology is entering a new phase of its history”. Which it didn’t do with the rise of molecular biology in the 1960’s? I don’t see where there is a paradigm shift in the relevant foundations: group selection was always floating around, adaptiveness and plasticity too (otherwise behaviour genetics would be a non sequitur), speed and limits of evolution (Haldane 1957 “The cost of natural selection” shows him thinking across several of these domains), canalization is not exacly a recent concept either. Specifically, plasticity and imprinting can always be modelled and relevantly (from the long term evolutionary standpoint) understood as GxE. The great quote (from Bonner 1961 about D’Arcy Thomson) in the introduction to the revised “The Causes of Evolution”:

    “If any gene combination produces a structure of good mechanical design, it may, because of this remain in the population through favourable selection. In other instances, there is clearly a direct effect of the environment which causes, by mechanical or physical forces, the forms of a living structure. In the case of these direct adaptations, we may assume that such responsiveness to the environment is adaptively advantageous and therefore the gene complement that favours responsiveness is maintained by selection”

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  23. I’m not a biologist, so I’m not going to comment on the technical issues here.

    My understanding is that the “yes” (EES) faction is looking to develop a broader, more inclusive theory of evolution than is currently accepted as standard. This has academic-political implications (concerning what research gets grants and valorization in journals), and these are by no means negative motivations, it’s how the academy operates. But it also has educational implications. Right now the Modern Synthesis is taught; is this what we want students to learn, or do wish broader horizons for their futures?

    Given that, I am sympathetic with the “yes” faction’s desire to broaden horizons, but also agree with the “no” faction that the empirical research must be there to validate it, and encourage further work.

    But let me shift gears here, and consider Patrice Ayme’s comment here. It presents us with some of the risks of this discussion as it moves beyond the discipline of biology.

    First, some of what Patrice writes is simply wrong. While Lamarck has left us a “full blown evolutionary theory,” it is a theory lacking in essential knowledge that we’ve accumulated since; and it includes conjectures that Darwin’s more mechanistic view has discredited. It is simply not the case that giraffes grew longer necks by reaching for leaves on higher branches. Also, a principle weakness of Lamarck’s theory, that even a non-biologist can see, is that it provides no adequate explanation for extinction; Lamarckian theory, taken to its logical limits, suggests that, barring environmental catastrophe, all species should be able to adapt and survive, and that hasn’t been the case. There have been catastrophes, but not all extinctions have occurred because of these, some simply occurred because a species could not develop adaptation to non-catastrophic environmental change.

    Also, Patrice argues that bias against Lamarck repeats the criticism of Christians when Lamarck’s theory first appeared; in fact, Lamarck’s theories were adopted by some Christians in the late 19th century because they are amenable to what we now call ‘intelligent design.’

    It should be noted here also that “spontaneous creation” is, first, not truly Lamarck’s theory (it’s not his own, and he seems to have pushed it further, to spontaneous creation of individual species, an ID hypothesis). And it is not the “currently accepted theory.” evolution has no theory on the origin of life, but a bag of chemicals achieving self-replication would not be “spontaneous creation,” since it could be explained by chemistry.

    Finally, Patrice suggests that the bias against Lamarck is culturally determined (because he was French). This is simply and only post-modernism, in the least convincing sense. Need we really come to this?

    Patrice’s comment, rather than making a strong case for Lamarck, evidence the murky waters this discussion faces when brought before the public.

    I don’t have a solution to that, only a reflection on the problem.

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  24. Massimo, I’m not convinced by your argument, mainly because you didn’t deal enough with the middle party in your trinity of “Mendelism, Mutationism and (neo)Darwinism”. Blind mutation to get a specific 3 billion sequence? I would need to see your work on all the mutations required to get to a human or any complex life from a single cell. I have never seen work where calculations of likelihood are made on the basis that each point mutation or deletion/insertion is purely blind. However, I have heard calls for such work to be actually done by biologists, so if it exists I would like to know. I’m with Hoyle and his 747 in a whirlwind. Until you can produce the stats, I will explore developments in Epigenetics, which leave open the influence of specific environmental conditions to gene expression based on “unknown” “flexibility”.

    You need to look at heat loss in the environment, ultimately. You have not factored that all that life “is”, is economy in compounding on a planetary surface. Heat is lost as atoms bond into anatomies, in Homeostasis with an environment. Homeostasis is a useful concept, also not mentioned in your essay. Look at convective chemistry in bonding to release heat. Remember, mutations are the gene expressions that release heat, and are important in Homeostasis, if only for that reason. Much more to be done, and far too soon to rest on unestablished laurels like “100 random mutation”.

    You can also go beyond Darwinian evolution at an overall level. DNA merely engineers pre existing non-living chemicals into anatomies. The next area you need to work on is the pre existing chemical capacities, liquids, gasses, solids, and so on, and how the behave naturally on Earth, and how that “flow”, “puff”, and “drop” can be used for anatomies, as for circulation, respiration, or digestion, basic stuff that has been overlooked. You might find that anatomies directly apply all natural chemical capacities to function, and we are bound to what is available in pre existing layers. Far more work to be done on that, rather than looking at what DNA has already engineered and making up loose environmental narratives for its ongoing existence.

    I’m far from convinced about Recombination, 100% Blind Mutation, and ex-post (neo)Darwinism (after the event to see if it can manage to survive somehow on the blind luck of having mutated). I need more work from biology on all these matters before deciding. I have no idea where “intelligent Design” fits into your argument – that’s not science and if you are arguing against spiritualists then I bow out as that is futile. But if it is science that interests you, then the point is to explore Epigenetics and whether it influences mutation and furthers expressions using pre-existing chemical availability (in natural layers for anatomical functions). For the time being I side with the writer of a piece linked at Dr Novella’s websitehttp://1drv.ms/1tnKM6f

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  25. Massimo makes a strong case.
    Darko Mulej has an interesting point:

    All in all, my opinion is this might be Kuhnian moment [paradigm shift] for biology” (http://bit.ly/QMhpM8)

    First consider that paradigm shifts can be micro, minor, moderate, major and complete.
    Then consider that human attitudes/behaviour/beliefs can be characterised as punctuated equilibria where the punctuations are some kind of paradigm shift, varying from micro through to major. In the world of mechanics we call that slip-stick friction(http://bit.ly/1EnII1d).

    As an example, in the management world we see
    1) The older manager come under increasing attack from his younger colleagues as he fails to respond to newer ideas and conditions. He fails to respond because he is too busy defending his own, one time novel, contributions from criticism.
    2) Crisis ensues and a Young Turk replaces the older manager, often a traumatic event.
    3) The Young Turk changes and reorganises things, implementing his new ideas.
    4) He consolidates his gains, making his revolution permanent.
    5) He defends his vision from attacks by the new generation of Young Turks. He becomes a worn out Old Turk.
    6) He is displaced by a new generation Young Turk and discarded.
    All of this happens in a space of 5 to 7 years. The management world is tough. Only exceptional managers survive more than two such cycles. One could call these micro organisational paradigm shifts. A small number of people have that remarkable property of self insight that allows them to renew themselves and thus transition between paradigm shifts.

    On a personal level it is sometimes called a mid-life crisis. Massimo admits to undergoing decadal shifts, which I consider a very healthy thing. That should be every person’s personal goal in life, regular self-renewal.

    My point is that paradigm shifts(from micro to major) are an important feature of the way society works. It is a form of adjustment and renewal. When there is a generation of Young Turks clamouring for change and a generation of Old Turks protecting their turf and the status quo, it is a warning sign of an impending paradigm shift(of whatever size). Is that where we are today with evolutionary theory? The signs are there. The science world is not as brutal as the management world so the cherished beliefs of the Old Turks will be quietly displaced as they die off(Massimo’s point).

    @richardwein

    If we want to make the point that evolutionary theory has changed enormously over recent years, we can just say that in so many words. We shouldn’t adopt confusing language just for the sake of making a point.

    Labels are important. They are a form of baptism that fixes the new concept(s) in the cognitive landscape, making them recognisable and a feature of our discourse. Or another way to put it. The label is like the handle on a suitcase, containing a group of related concepts. Without the handle, the suitcase is difficult to manage, in discourse, dissemination and in teaching.

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  26. Maybe we could be more focused and first try to agree on some essential tenets of Modern Synthesis. If some of them are found shaky then (for the purpose of this article) MS loses.

    My proposition is:

    1. Gradual evolution results from small genetic changes that are acted upon by natural selection.

    2. The origin of species and higher taxa, or macroevolution, can be explained in terms of natural selection acting on individuals, or microevolution.

    (Taken from Freeman & Herron as quoted in http://campus.udayton.edu/~hume/EvolSynth/evolsynth.htm.)

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  27. Coel,

    “Well, not really, indeed your article could be read as support just as much for the “no” camp”

    I’m glad to see that physicists are quick to formulate opinions about technical matters outside of their expertise. No worries, you are in good company: Sean didn’t change his mind either, despite a private correspondence with me.

    “A crucial phrase is “and yet do not contradict” the Modern Synthesis. When many of the items you list were uncovered, they were often accompanied by “and this overturns the Modern Synthesis” hype.”

    But the Modern Synthesis itself didn’t contradict neo-Darwinism, and yet even then there were hyped claims. If my no-colleagues were even slightly more familiar with the history of biology they would realize that there is no difference btw the two cases.

    “the “no” camp are saying, yes those new ideas are interesting and important and need to be synthesized into the overall picture, but they amount to perturbations on the basic neo-Darwinism Modern Synthesis”

    You may have gotten that impression from the Nature exchange. In fact, if you look at the literature over the past decade, the no-camp has been in denial about a number of new phenomena and ideas, and has only grudgingly acknowledged part of them eventually. See Dawkins’ dismissal of the entire field of epigenetics in a single footnote in one of his recent books.

    “The “yes” camp then seems to be saying that the combined effect of all the items you list is overall more important for evolution than the ideas of the Modern Synthesis”

    No, the yes-camp is saying that a number of new discoveries and conceptual advances have changed the very way we look at evolution, particularly in terms of abandoning the gene-centric paradigm that has characterized the MS since its inception.

    “I could also be convinced that both sides are mostly agreed on the actual science”

    They don’t. The no-camp had dismissed phenotypic plasticity for decades before it finally established itself as a major biological phenomenon out of the sheer volume of published papers. And see my comment about epigenetics above.

    imzasirf,

    “I don’t see the transition being as strong if the extended synthesis is not contradicting the modern synthesis, why not just consider it the natural progression of the modern synthesis?”

    See my comment about gene centrism above.

    “it doesn’t seem like the old guard is embracing some aspects such as group selection as strongly as say David Sloan Wilson and others. However, I’m not knowledgeable enough in evolutionary biology to know if one or the other side is exaggerating their claims”

    My take is that David is indeed exaggerating, while some of his detractors are minimizing. The best take on multi-level selection is provided in the book by Samir Okasha that I cited in the notes.

    “where every few years someone claims to have the absolute new model and it’s usually the same old hat with a few added bells and whistles”

    Tell it to the physicists… 😉

    richardwein,

    “We agree this is just a matter of terminology, and there’s no substantive question at stake”

    No, we absolutely don’t. I think there are very many substantial matters at stake. I was simply saying that the crucial point is to recognize the impending end of MS-type gene centrism, whether one wishes to baptize the new way of looking at things or not.

    “We adopt new names when there’s a specific identifiable difference. But when it’s lots of differences adding up to a matter of degree?”

    See above, gene centrism.

    “We’re not talking here about two distinct versions of a theory. We’re talking about a theory which has developed continuously over time”

    Again, same as the neo-Darwinism to MS transition, which none of the no-people has any trouble whatsoever in recognizing.

    “My impression is that the adoption of the terms “Neo-Darwinism” and “Modern Synthesis” was not as totally arbitrary as this, but corresponded to a somewhat specific (if somewhat fuzzy) transition”

    No more or less than the current situation, in my opinion.

    “whether that’s so or not, the names have turned out to be something of a source of confusion, because of their non-specific names”

    I can assure you that no professional biologist is confused about what the MS stands for.

    Artem,

    “Where would the Baldwin effect fit within your list of on-going directions? I am tempted to push it under phenotypic plasticity, but I am not sure if that is the most natural place”

    It is, see my book on plasticity, cited in the notes. There too goes Waddington’s genetic assimilation.

    Bill,

    “The discovery that noncoding regions of DNA play at least as important a role in evolution as coding regions is particularly important, and really wasn’t anticipated by the Modern Synthesis”

    It wasn’t anticipated, but it has had little conceptual / theoretical effect. Unlike, say, phenotypic plasticity and epigenetic inheritance.

    “It looks more like a program of research into developments which will eventually produce a new synthesis.”

    Pretty much the same could have been said of the MS when Huxley published his book giving it that name.

    michaelfugate,

    “When in reality organisms and the environment are in a feedback system – changing each other. Natural selection is not all powerful.”

    Precisely. Gould has done an excellent (if baroquely presented) job in his last book of analyzing the changing role of natural selection in evolutionary theory. Needless to say, the no-camp positively hates that book.

    brandholm,

    “Punctuated Equilibrium alone is a pretty important change in perspective regarding rates of change.”

    Correct. It was funny when a few years ago I asked two colleagues about puncteq, Doug Futuyma (a population biologist, and one of the co-authors of the no-article) and David Jablonski (a paleontologist). Doug was absolutely sure the idea had been debunked. David was equally adamant that it is now accepted in mainstream evolutionary thinking…

    “Margulis’s theory of endosymbiosis which radically adds to methods of evolution”

    Indeed. Under the Extended Synthesis that one falls under the rubrics of both evolvability and multi-level selection, as a special (if very important!) case.

    “That said, I’m not certain if I’d qualify a change in name as an “urgent” need. That word drives the question, urgent for what?”

    That’s when we go from science to sociology of science: urgent to trigger funding and to reorient research programs, which is precisely what the no-camp wants to clearly to avoid.

    “I have to admit that doesn’t raise to the level of a crisis.”

    It doesn’t. Notice that I never used the word, unlike some of my more extreme yes-colleagues, like Eva Jablonka.

    Like

  28. DM,

    “The core idea Darwin had seems to stand, and we are only refining and adding to it”

    As I already said: precisely the same happened when the MS was ushered in, so why not now?

    “I don’t see any significant paradigm shifts happening”

    I have argued in print (see my “Paradigmi” paper cited in the notes) that biology has never undergone a paradigm shift, and likely never will. But science doesn’t always advance in Kuhnian fashion. And, again, there was no paradigm shift when the MS came on stage.

    Darko,

    yes, my list is definitely incomplete, and it leans toward the conceptual rather than provide a list of empirical discoveries:

    “Transponsable elements”

    Indeed, although that one changed things in molecular genetics more than in evolutionary biology, because the latter is mostly an organismal discipline, which can treat much (though not all — see epigenetics) of what happens below as a semi-black box.

    “virology is in recent years in a kind of renaissance due to discovery of giant viruses”

    True, but my comment would be along the same lines as the previous example.

    “Modern Synthesis talks about natural selection that acts on novelties, but it let unspecified how this new features arise”

    Yes! The ES brings in facilitated variation and evolvability as some of the new tools to deal with evolutionary novelties.

    Patrice,

    “The full blown theory of evolution was proposed by Lamarck. This was a great conceptual breakthrough”

    It was also the wrong theory, unfortunately.

    “Lamarck got hated for all this by the forces of Christianity”

    Maybe. In this he certainly had something in common with Darwin.

    “Lamarck being French, some feel more appropriate to attribute the discovery of evolution to the English Darwin”

    Nah. In fact quite the opposite: almost the only people (with the exception of the above mentioned Jablonka) who still insist on Lamarck today are, ahem, French. And I suspect that nationalism has a lot to do with it.

    Arno,

    “the development of population genetics by Bateson, Hardy, Weinberg, Morgan, Johannsen, Nilsson-Ehle, Tammes, and Punnett … seems to me that this work … is much more important than Weismann’s germ-plasm theory … Do you agree?”

    It depends how you measure importance. Most of those people are recognized in standard textbooks, but it was Weismans’ ideas that provided the first clear rejection of Lamarckism, which was conceptually crucial.

    “Your set of important developments doesn’t include molecular evolution, genetic drift, the neutral theory”

    Partly because this was a 3000-word essay, not a textbook (though see my co-edited book on the ES), partly because I picked the aspects of the ES that I think are conceptually most relevant. Also see my comment above about much of the molecular stuff being invisible from the perspective of evolutionary theory.

    “Don’t these developments contradict certain tendencies of the modern synthesis?”

    Not really, I don’t see how.

    “punctuated equilibrium and macro-evolution. Don’t these ideas contradict the view of the modern synthesis that macroevolution is just microevolution over millions of years?”

    Yes, but that idea was never really central to Darwin’s insights, as Huxley himself pointed out to him.

    “According to S.J. Gould the modern synthesis was already “effectively dead” in 1980”

    The MS is not dead, but it is transforming…

    “The ‘yesies’ clearly think that evo-devo and niche construction are of the second kind. Do you agree with them? If so doesn’t that contradict your statement dat the new developments do not contradict the modern synthesis?”

    I disagree, there are no contradictions of the fundamental ideas of natural selection, common descent, and basic population genetics. But there is so much that those ideas don’t even come close to encompassing.

    marclevesque,

    “At the end of the article they invite “Laland and colleagues to join us in a more expansive extension” to evo theory, so I would like to be positive and assume they are sincere”

    I’m sure some of them are. But I’ve seen too much backstabbing at the level of peer review of papers and grants to fully believe it. And no, I’m not going to name names. And yes, the backstabbing is from both sides.

    stevenjohnson,

    “I see neutral mutation is omitted”

    See comment above about molecular vs evolutionary biology.

    “the importance of genetic drift is minimized”

    Indeed, the issue of genetic drift — both in terms of its theoretical status and of how to study it empirically — is a big one, but it’s rather technical, and I didn’t want to go that direction in the essay (I have published technical papers on this topic, though).

    “What is it that makes even such a highly respected writer as Zimmer make what appears to a layman to appear to be a gross lapse in elementary judgment?”

    What Gould and Lewontin called the adaptationism program, which should have been greatly diminished by their famous 1979 critique of it. And yet…

    “It seems to me to be a consequence of the emphasis on gene selection”

    Exactly, gene centrism.

    “If you look at the popular presentation of evolutionary theory, you see evolutionary psychology, and practically nothing else.”

    And that, I maintain, is highly unfortunate. Talk about adaptationism gone wild.

    “there is an awful lot of confusion about the question of when natural selection acts to conserve an organism’s morphology as opposed to when it serves as a driver for the emergence of evolutionary novelties”

    Again, yes, see my comment above about additional mechanisms, such as facilitated variation, and phenomena, such as evolvability.

    Like

  29. Thanks Massimo. I’d love to see more articles on philosophical issues in biology – especially one dealing with the role of functional explanations in biology (which has been touched upon in the ongoing discussions about reductionism, but I don’t recall ever being focused on in-depth.)

    Like

  30. Massimo,

    Re my earlier comment and your reply: The importance of non-coding DNA has not yet been fully absorbed, but it really is a high-level game-changer, because it provides an improved explanation of gradualism. Mutations to coding DNA, if they cause any change in fitness at all, often cause a large change and usually a harmful one. Mutations to non-coding DNA, however, don’t change the structure of proteins, they change the amount of a protein that is produced. That’s intrinsically a much less “violent” sort of change — a quantitative rather than qualitative change, if you will. It’s much easier to understand gradual evolution in terms of an accumulation of quantitative changes than in terms of a superposition of qualitative changes.

    I’ll add to my earlier comment by saying that in my view the most important feature of a “new synthesis” will be a vastly improved account of evolution at the DNA level. In particular, not very long ago most people thought that evolution works mainly by changing the structure of proteins. It is now clear that substantial changes in protein structure are rare, and that the bulk of evolution involves changes in the amounts of proteins that are expressed.

    Best regards, Bill

    Like

  31. I think that there is enough to say that that there is a major shift, and we need to call it such. My reasoning is as much sociological as it is biological; Massimo himself mentions sociology of science.

    Besides Brandholm’s mention of transposons as material that Massimo didn’t include at first (but did in a comment), and other things, there’s also prions (mentioned by nobody yet). We still have relatively little handle on just how much they may mean, or not, to the bigger picture, and I remember how hugely deep-seated the reaction of much of the evolutionary biology community was to Prusiner’s ideas — and I suspect a certain amount of that community feels that way still today.

    I have no idea if prions have an effect on heredity. But, I don’t know that they don’t, either. And, even without that, they seem important enough. That said, while prions may challenge some aspects of older evolutionary theory, they are NOT support for any neo-Lamarckian ideas. (That said, both seriously and jokingly, I’m surprised Patrice didn’t mention them as such.)

    And, that ties in with the resistance to the … Kuhn time, hinted at by others.

    Re the idea of a paradigm shift, I think that the amount of heels being dug in is precisely another reason why we actually do need to say this list of things Massimo has, plus further material as noted. Massimo also notes that this may be partially a younger-vs-older split, which also tracks with bringing Kuhn, or a stereotypical version, into the picture.

    Related to that, if a paradigm is being shifted, the whole of “what’s new” gets addressed seriously as a group.

    Per Labnut and his quote, I’d put this at moderate as far as a level, maybe between moderate and major.

    I think Alexander addresses the negative side, or potentially negative side, of this. But, my take is the flip side of his — some “last ditchers” are resisting the idea that this is actually a real difference mainly because of fears of the likes that Alexander mentions.

    StevenJohnson has a good point vis-à-vis the old way of thinking that one gene = one protein, too.

    John Smith Hoyle’s 747? Meh, it’s just Paley’s watch, gussied up and modernized. Epigenetics is part of a neo-neo-Darwininan synthesis, not something apart from it. On a quick read, it sounds to me like you’re looking for creationist or ID turtles all the way down.

    Like

  32. EJ Winner to accuse me of misquoting the traditional distortion of what Lamarck said on giraffes. What Lamarck truly said on giraffes is compatible with the most mechanistic “Darwinism”.

    Gould said: “[they] re-read Lamarck, cast aside the guts of it…”

    When one quote Planck, one quotes what he wrote in 1900 (right), not what he wrote in 1899 (not quite right). Lamarck’s ideas clearly changed between 1801 and 1815 (when he did not believe anymore in continuous spontaneous creation).

    Science is subtle. So is vocabulary: “principal” is not “principle”.

    Massimo:
    “The full blown theory of evolution was proposed by Lamarck (Patrice)… It was also the wrong theory, unfortunately.” (Massimo). Massimo, do you mean that evolution theory is the wrong theory?

    Lamarck’s main body of work is just about establishing “evolution” as Darwin’s personal teacher named it.
    Lamarck also suggested several evolution mechanisms jointly operating.

    When Copernic, copying Buridan and Aristarchus, proclaimed heliocentrism, nobody asked him for a mechanism. Still, one attributes heliocentrism to Copernic. While Copernic did not discover General Relativity, one still do not attribute heliocentrism to Einstein.

    If Darwin’s teacher taught Darwin in 1825 that Lamarck had established “evolution”, why should we say now that “evolution” is the wrong theory? Because, being in the Anglo-Saxon realm, we have to be Anglo nationalists? Is it all about tribalism?

    The lies about Lamarck were deliberately crafted by a Christian fanatic, the biologist Cuvier who totally believed that God had created all the species. Cuvier’s argument was that, as there no evidence of recent evolution, there could be none. Lamarck retorted that the argument was mathematically stupid and proved, instead, that the Earth was very old.

    In any case, attributing “evolution” to Darwin, instead of Lamarck, constitutes scientific, and philosophical fraud. For want of a nicer way to put it.

    Now to elevate the debate.

    It is clear that life on Earth did not start with DNA. It is also clear that there are non-DNA inheritance mechanisms (prions, cell division). Thus life cannot be reduced to the inheritance of “genes” (or then the notion of “genes” has to be freed from DNA).

    So what is life? Pretty much the inheritance of more or less reproducible hyper complex geometrical structures. Not just reproduction of DNA.

    When one goes from pure nanotechnology to “life” is an open, philosophical problem: are viruses alive?

    There is a basic flaw in the strictest “Darwinist” idea of purely mechanical selection by greater aptitude of reproduction of superior genes. First, as I said, although that is observed, it’s not all what is observed. Lying by omission is still lying.

    What is evolved is enormously complex, reproducing systems. To reproduce better, it would help evolution if it had eyes. Now, true, species evolved eyes. But evolution itself evolves, it is a sort of meta-organism. Did evolution evolve eyes? Did evolution evolves teleonomy?

    Teleonomy is what full blown “Darwinism” would stumble on, given enough time. So did it have enough time? Was Lamarck even more right than is assumed today?

    Like

  33. Hi Massimo

    I’m glad to see that physicists are quick to formulate opinions about technical matters outside of their expertise.

    Well, on this one we can’t bow to the biologists’ consensus, since there are plenty of notable biologists on both sides. [Of course no non-physicists would ever have an opinion on, say, string theory, would they? :-)]

    The substantive issue seems to be over whether new developments are adding to the Modern Synthesis or over-turning parts of it. You seem to be saying that a lot of gene-centric ideas are being over-turned. Can you give some more commentary on which of those ideas are wrong and why? (Some of the stuff I’ve read on that topic seems to have a large element of strawmanning.)

    Hi imzasirf,

    Having read some debates from professional biologists online about group selection, it doesn’t seem like the old guard is embracing some aspects such as group selection as strongly as say David Sloan Wilson and others.

    It would be wrong to see that issue as young upstarts versus old-guard fuddy-duddies. The idea of group selection is pretty old and many of the leading advocates (e.g. D.S. Wilson, E.O Wilson, Nowak) are not exactly spring chickens themselves. It could be that the majority are correct in seeing group-selection as a minor part of the picture.

    Hi Darko Mulej,

    Well known example is hemoglobin, of which they are many types and they were made by duplication event, not minor modification.

    If you consider the issue in terms of information content, then a duplication is indeed a minor change of information content.

    Hi brandholm,

    Punctuated Equilibrium recognizes that change is not, or at least is not required to be gradual. Changing conditions can lead to abrupt changes in species.

    It’s important to be clear on timescales in such declarations. Change that is abrupt on a geological timescale is still gradual on a generational timescale, and thus still Darwinian gradualism. On which point:

    Massimo,

    I asked two colleagues about puncteq, Doug Futuyma (a population biologist, and one of the co-authors of the no-article) and David Jablonski (a paleontologist). Doug was absolutely sure the idea had been debunked. David was equally adamant that it is now accepted in mainstream evolutionary thinking…

    So the population biologist was thinking about generational timescales, and the paleontologist about geological timescales?

    Hi Arno Wouters,

    You do mention punctuated equilibrium and macro-evolution. Don’t these ideas contradict the view of the modern synthesis that macroevolution is just microevolution over millions of years? According to S.J. Gould the modern synthesis was already `effectively dead’ in 1980. Was he correct?

    No, and no. This is the problem. People such as Gould over-hyped their ideas by declaring that they’re overturning the Modern Synthesis, thus forcing others to play the fuddy-duddies of the “no” camp. (Even Massimo, who supports the “yes” camp, admits that some in that camp over-do the rhethoric.)

    Hi Patrice Ayme,

    You are right that history is unkind to Lamarck, in remembering him mostly by one thing he got wrong, and not his many major advances.

    Like

  34. IMHO this debate is confused and poorly framed (http://www.molevol.org/the-great-non-debate-on-evolutionary-theory-nature-oct-2014).

    The disputants agree that we need a Grand Unifying Theory of Evolution (GUTE), and that the Modern Synthesis (MS) was it; they disagree if EES should be the new GUTE.

    When did “theory of evolution” become “list of topics”? Scientific theories have structure and coherence. They are falsifiable. This debate seems to be mainly about something more sociological, like a “community of practice.”

    Originally, the MS *was* a GUTE, unified by Darwinian doctrines of gradualism, the creativity of selection, and the denial of internal forces. It began to fall apart immediately when protein comparisons evoked a mutationist view that Mayr, et al had rejected, spilling out into a new sub-discipline (“molecular” evolution) and evoking a novel theory (the Neutral Theory). Then we rejected gradualism. Then we found that biases in variation are important. And so on.

    The doctrinal basis of the MS rotted away, leaving only the hard bits– findings and models we still trust. Bizarrely, even opponents now treat the MS as an open-ended framework consisting of these hard bits, leading to perverse conclusions, e.g., the MS now *includes* the Neutral Theory, on the grounds that both invoke population genetics! Fisher is rolling over in his grave.

    This bait-and-switch is sustained by a lot of bad historiography– not Pigliucci’s fault because 90 % of available historiography was written under the influence of Mayr’s MS kool-aid. The story about Fisher coming to the rescue is nonsense. As Gayon explains in _Darwinism’s Struggle for Survival_ (1998), “the fundamental doctrines of quantitative genetics were developed early in the century, long before the publication of Fisher’s canonical article of 1918 which is often credited with having laid the foundations of the discipline” (p. 316).

    Mendelism brought down Darwin’s theory, not because Mendelians were too stupid to synthesize mutation, heredity and selection, but because Darwin’s original theory was non-Mendelian. The Mendelians had a pluralistic view like ours: they welcomed selection and neutrality; they allowed (superficially) smooth change and discrete jumps; they rejected Lamarckism yet welcomed mutation biases. Their view is labeled “anti-Darwinian” or “non-Darwinian” because it denies _natura non facit saltum_ and awards individual mutations an important role as a source of initiative, direction and creativity.

    There *was* an eclipse of Darwinism. We are indoctrinated to see it as an eclipse of reason. It was not. It was a period when leading thinkers doubted the idea that selection is creative and, though remaining interested in selection, were also interested in other ideas.

    To the extent that the MS is a genuine GUTE, it is wrong. To the extent that it has devolved into an open-ended framework, we should refer to it as “Mendelian-Mutationism”, admit that it is non-Darwinian, and credit it to early geneticists.

    And we don’t need a GUTE. We haven’t had one since 1959.

    Like

  35. Patrice Ayme,

    Craig Holdrege has an interesting discussion on the giraffe issue, wherein he discusses both Lamarck’s and Darwin’s views on the matter. An importment moment in his quote from Darwin is when Darwin, while noting that long-necked giraffes would have an advantage in eating higher leaves, also remarks “the individuals, less favoured in the same respects will have been the most liable to perish….” The death/ non-reproducibility of animals (including whole species extinction), that have no traits adaptive to environmental change is crucial to Darwinian theory. It actually provides a stronger foundation to the development of complexity than seemingly purposive tendencies to progress. Holdrege makes the point that the giraffe’s neck, not only being long enough for eating, is almost too short for drinking. So why does this ungainly animal with such a difficulty reaching down to water sources have the neck it has? As Holdrege notes, the neck serves several purposes; but the simplest answer is that it survived, and its predecessors did not.

    Michael T. Ghiselin also believes that Darwin depended more on Lamarck than many think, that Lamarck is poorly represented in history. But he makes a telling remark:

    “Lamarck’s approach to evolution was that of a metaphysician rather than a natural scientist. It invoked a mystical assumption (the notion that organisms sought “perfection” and tended to become increasingly complex and man-like) which could not be treated scientifically (…).” “Darwin’s concept was a well articulated body of scientific thought that could be, and was, tested by recourse to facts. Lamarck’s was not.”

    Unsurprisingly, Ghiselin remains a committed Darwinian.

    Without belittling Lamarck, the fact remains that Darwin addressed evolution in a manner that articulated known facts and made possible further research and discovery of new data, and that is necessary for a scientific model that is not purely speculative.

    As to the Lamarck- Cuvier conflict, it should be remembered that these men came to prominence in revolutionary times. That doesn’t excuse Cuvier, but neither does it make him out to be the ‘Christian fanatic’ you paint him as. Anyway, the history of ideas, while embedded with their undeniable contributions, moved elsewhere.

    “When Copernic, copying Buridan and Aristarchus, proclaimed heliocentrism, nobody asked him for a mechanism.”

    It is the mechanism, not the model, that makes science.

    “Because, being in the Anglo-Saxon realm, we have to be Anglo nationalists? Is it all about tribalism?”

    This is the post-modernism of which I complained. Insisting on purely ideological motivations for scientific theory and research, and suggesting such motivations thus delegitimate the foundations of a science, seems dangerous. At which points are the data of the sciences allowed to speak for themselves? At what point can we reach agreement on a theory that adequately explains the data?

    Your final remarks seem to wander into some sort of pantheist teleology (not teleonomy); if so, I can’t agree.

    Holdredge on Giraffes: http://natureinstitute.org/pub/ic/ic10/giraffe.htm

    Ghiselin on Lamarck and Darwin: http://www.textbookleague.org/54marck.htm

    Like

  36. Hi Coel, I don’t mind if people from other fields express an opinion on biology, but to be truthful I actually don’t weigh in on things like string theory… usually because I can’t figure out what they are talking about 🙂

    In your response I think you were being a bit unfair to me (and Massimo) by stressing punctuated equilibrium (PE) to the exclusion of all the rest of the points mentioned. It is the combination, not each point singularly that suggest an update is necessary.

    Regarding PE itself, I understand the difference between generational and geological timescales. And I am telling you I still believe PE represents a significant change in viewing how evolution occurs. Rather than species gradually accommodating to a stable environment, PE suggests that change is or may primarily be the result of species adapting to a changing environment (whether they move into a new one or the environment itself changes). This suggests a shift in both the driver and timeframe of change that should be expected from theory… and makes separate predictions of what one can expect from accumulating fossil evidence.

    To this (and I am restricting myself from discussing the mountain of other changes Massimo noted) I also mentioned two completely novel evolutionary mechanisms: endosymbiosis and transposons (aka jumping genes aka transposable elements). You appear to have ignored that and asked Massimo:

    “You seem to be saying that a lot of gene-centric ideas are being over-turned. Can you give some more commentary on which of those ideas are wrong and why?”

    Well lets consider endosymbiosis. Here is a case of (at least) two separate individuals merging over time into one individual with two separate genomes. They are now inseparable yet there was no acting on any single set of genes to produce that outcome. It was not a single genome splitting in two, due to external pressures, to create a sub-individual inside. This was arguably a cellular level event, and certainly did not (need to) involve external selection pressures on inherited variations.

    And how about transposons? This adds a completely unique form of species interaction, directly editing the other’s genome. And it provides a novel mechanism for future change within a genome (beyond random mutation). Inheritance (as it is normally conceived of) is not the only source of raw material for selection.

    The familiar “tree of life” image with all of its divergent evolutionary branches is arguably a product of the MS. These last two mechanisms I mentioned would demand branches (from potentially vast distances on the tree) merge to form new branches.

    If that is not a significant alteration to the picture of evolution, I do not know what is.

    If you feel that integration of separate species at the cellular and genomic level to form new species is something predicted by or amenable to MS, and traditional “gene-centric” mechanisms, I would like to know how.

    Hope I didn’t come off combative.

    Like

  37. Standard Model of physics is a phenomenology hodge-podge, without any theoretical base {how do these SM particles rise?}, and M-string theory tries to play that theoretical role. While at LHC, the SM is all there is, but all physicists accept that SM is not complete as dark matter and dark energy are not explained by SM.

    The current evolutionary biology is also a phenomenology hodge-podge, with the MES (modern evolutionary synthesis) tries to play its theoretical role. Yet, those ‘no’ camp people do not accept that MES is not incomplete while every street walking people knows that the most obvious empirical human attributes (intelligence and consciousness) are not explained by that MES. There are two very simple (simple,…) questions.

    Q1, is ‘intelligence’ an empirical fact as a human attribute?

    Q2, is ‘intelligence’ a result of evolution or just pops out from the ‘blue’? [Note, if it pops out from the blue, I would like to know what the essence is for that ‘blue’.]

    If the ‘intelligence’ is the result of evolution, there is no reason of any kind for anyone to worry about that biology-phenomenology-hodge-podge. The question is very simple, show me {what is the evolution process for the intelligence?} and nothing else. If MES can do it, it will be the happy time for everyone. But, MES is far removed from answering this simple question.

    For the physics-ism, the intelligence must arise from the bottom of physics laws, and I have set the criterion {proton (or neutron) must carry a counting device, such as Turing computer}. With this criterion, the ‘evolution’ is no longer a biology issue but an issue of physics. If nothing can meet this criterion, the only option will be to take the ‘blue’ for answer, as MES is far removed from the issue.

    The sole mission for M-string theory is the ‘string-unification’ {describing SM particles with the string-‘language’}. While M-string theory fails on its mission dismally, the following G-string could be the rescue.

    One G-string (a, b, c) has eight (8) strings.
    String 1 = (V, A, A 1) = {1st , red, 2/3 e, ½ ħ} = red up quark.
    String 2 = (A, V, A 1) = {1st , yellow, 2/3 e, ½ ħ} = yellow up quark.
    String 3 = (A, A, V 1) = {1st , blue, 2/3 e, ½ ħ} = blue up quark.

    See the entire description at https://scientiasalon.wordpress.com/2014/05/22/my-philosophy-so-far-part-ii/comment-page-2/#comment-2432

    Is this G-string correct?

    Well, G-string is not a theory, not a model as it has no prediction nor any test point. It is simply a god damn ‘language’ which ‘spell’ out the SM particles. Its validity depends only on the {cross the t’s and dot the i’s}.

    Then what? So what?

    With this G-string language, both proton and neutron are in fact ‘Turing computer’ (see http://www.prequark.org/Biolife.htm ).

    No, extended MES won’t cut it, sorry.

    Like

  38. Coel, on reflection I think that gene duplication might not be news, on some estimates more than half genes are due to this event. But there are many examples, when new genes acquire new function, like hemoglobin – oxygen transport) vs. myoglobin – oxygen storage (see http://tinyurl.com/meqf427)

    Massimo, my point about tranponsable elements would be to see them as enablers of novelties: one can see them as help from foreign organisms (mostly micro). Or can give label like “junk DNA” though this view is recently seriously challenged (https://www.encodeproject.org/). Similar for viruses (http://tinyurl.com/qac7sta).
    To continue with microorganimsms, our gut flora got some exposure lately, being renamed (microbiota or microbiome), counted (there are more of them than our body has cells), weighted (about 1 kg) and sorted (bacteria, arhaea and fungi). Or consider Mastotermes darwiniensis, a termite: how can a worker termite be considered individual when it is the hive that is the reproductive unit of the species, and the worker cannot even digest cellulose without its gut symbiont, Mixotricha paradoxa (see http://www.jstor.org/stable/full/10.1086/668166).
    Here are lurking two interesting philosophical questions: what is individual and what is level natural selection is acting on.

    Like

  39. Regarding Kuhn and this topic, I think the more interesting figure is Georges Canguilhem (who is, sadly, lesser-known in the Anglosphere). Canguilhem was also concerned with epistemological “breaks” in knowledge and concepts, but unlike Kuhn he was, as a trained physician, concerned with knowledge and concepts from the standpoints of medicine and biology. The breaks that Canguilhem proposed are “milder” than the rather radical paradigm shifts in Kuhn, in that knowledge/concepts remain continuous with the previous paradigm while also representing a discontinuity.

    This is closely connected with how he understood the nature of living orgnaisms, which centered on the question of how we can have a concept of pathology in the life sciences. This leads to an account of normality that turns on the possibility of dysfunction. The organism creates norms and meanings in its responses to and engagement with its outer surroundings. This account develops into an information-theoretical stance, wherein the conjunction of information about the developing organism, found in the structure of genes (etc.), and the matter that composes the living organism, in which that information is “inscribed”, can be understood as a way of (re)introducing form and function back into the life sciences.

    In the context of Canguilhem’s epistemological story, biology can quite happily accept these notions without exactly bringing us back to the pre-modern preoccupation with Aristotelian formal and final causes. If we’re to take the shift in evolutionary thinking in this direction, then we can see that there is a change in the way of thinking which is both a break from and continuity with the Modern Synthesis. The knowledge and the concepts employed are both changed (and I’d think rather substantively if we’re recognizing that form and purpose do mean more than a “mechanistic” view can allow) but in the end, we haven’t made the same radical departure from the content of our theories (which you’d find with Kuhn).

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  40. Patrice Ayme,

    “When Copernic, copying Buridan and Aristarchus, proclaimed heliocentrism, nobody asked him for a mechanism”…

    Hmm, what do you mean by copying? Are you claiming that copying is synonymous of plagiarism? If it is so, I’m afraid that you are misguided. Can you tell us a book written by Aristarchus proving the heliocentric theory? This book, if ever existed, is lost and wasn’t known by the Polish astronomer.
    Did Aristarchus copy Heraklides of Ponticus? Did Heraklides copy a former astronomer? Did Kepler the same regarding Copernicus? Did Newton copy Euclid? Did Einstein copy Euclid, Newton and Riemann? According to your opinion, it seems that the history of science is a sequence of plagiarists. Who buys it?

    On the other hand, you refer to Christianism in an unpolite and confusing way. Luther, that was Christian though Protestant, attacked Copernicus (that was Catholic), and called him insane and heretic. Regarding Cuvier, he based in the Bible his attack on Lamarck, don’t forget that the Bible is not a Christian but a Jewish book. The Christian books are the four canonical Gospels, the Acts of the Apostles, 21 Epistles, St. John’s Apocalypse and the apocryphal Gospels.

    In other words, if you like mentioning the religious bigotry as a deplorable fact, take into account that the Bible wasn’t written down by Christians. Obviously, there are many examples of dogmatic Christians, scientific and laymen, that have undertaken harmful actions against science, but don’t forget that the Bible is a Jewish, non-Christian book.

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  41. I don’t think I could really adjudicate one way or another, but it was certainly an interesting overview of the state of the science.

    And I learned something about a matter which has puzzled me for a while, how something like a beak can adapt in such a short time when it would seem to involve lots of different mutations to make the upper and lower beak fit and still attach to the face etc.

    As for punk eek, I have always thought that something like this would turn out to be the case, maybe not quite as Gould envisioned, because mathematical models of random mutation and natural selection seem to nearly always progress in this manner (unless you cheat and hint it to the path).

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  42. First, I don’t understand what the issue is. According to Massimo “the yes-camp is saying that a number of new discoveries and conceptual advances have changed the very way we look at evolution, particularly in terms of abandoning the gene-centric paradigm that has characterized the MS since its inception.” I can see that this is what the ‘yesies’ are arguing in the *Nature* article.

    However, according to the title of that paper, the question at stake is not “did evo-devo, niche construction, phenotypic plasticity, and epigenetic inheritance etc. change our view of evolution?” but “does evolutionary theory need a rethink?”

    I would think that the ‘yes-answer’ to this latter question implies at least the existence of phenomena that are superficially at odds with current evolutionary theory, if not phenomena that contradict current evolutionary theory. The nay-sayers in the *Nature* paper seem to argue that this isn’t the case.

    Perhaps the parties are talking past each other?

    Second, I wonder how many parties there are.

    I cannot set aside the impression that the views of people like S.J. Gould, Richard Lewontin, Michael Lynch, Eugene Koonin, and Andreas Wagner as well as those of Bill Skaggs, Darko Mulej, and astolzfus in preceding comments are quite different from those of both the ‘yesies’ (whether moderate or extreme) and the nay-sayers. Unlike Massimo and the nay-sayers they seem to reject the modern synthesis. They seem to do so not because of its gene-centrism (as the yesies do according to Massimo) but because of its Darwinism, that is because of its view of evolution as adaptation to changing circumstances by means of natural selection. In his reply to earlier comments, Massimo repeats his belief that most of the molecular stuff is invisible from the perspective of evolutionary theory. This is obviously not the view of the people I just mentioned.

    Third, it seems to me that to address the question whether evo-devo, phenotypic plasticity, niche construction and epigenetic inheritance have fundamentally changed evolutionary theory, we need a theory of what counts as a theory, what as a change of theory, what as an extension of the theory, and of how to asses the importance of certain changes etc. Such a theory is lacking in Massimo’s account.

    Several commentators refer to the alleged extended synthesis as a Kuhnian revolution. As long as they do not substantiate their view with a list of the anomalies that confront the modern synthesis and an explanation of the way these are solved in the new theory, I see no reason to take this view seriously.

    I don’t understand Massimo’s intuitions about what changes are important. For example, I don’t see why De Vries & Bateson’s distinction between variation and fluctuation and Morgan’s origin-fixation model are less important conceptual innovations than Weismann’s invention and rejection of ‘Lamarckism’.

    Mayr, of course, maintained that Weismann was the most important evolutionary biologist after Darwin, but I don’t think that Mayr’s judgments and the distorted view of the history of evolutionary thought with which he supports them are an acceptable criterion for importance.

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  43. Unfortunately, the following, introductory paragraph got lost when I copied my previous comment from my editor to my browser:

    The more I think and the more I read about modern, new, extended and expanded syntheses, the less I understand what is going on.

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  44. Alexander,

    “Seriously, why not just do some practical research and then let our descendants in fifty years decide”

    Seriously, let’s not insult a large number of people who have, in fact, done precisely what you are asking, and for decades. Besides, shall we retroactively apply your rule to the Modern Synthesis too? Huxley named it while he was in the midst of it, not fifty years later…

    “Epigenetics is really interesting but as long as it doesn’t change the DNA sequence it is irrelevant in the long run”

    Thinking of evolution only “in the long run” is a serious mistake. Evolution happens from one generation to the next, and natural selection works at that level too.

    “I have never understood what is supposed to be so special about punctuated equilibria either … this is just normal Modern Synthesis evolution with genetic bottlenecks and stabilising selection”

    Right, and that’s why so many people have so ferociously attacked Eldridge and Gould for decades now.

    “And macroevolution =/= microevolution? That seems to boil down to the observation that mass extinctions have a strong stochastic element and entail very unusual one-off selection events”

    Not really, you may want to read some of the literature on species selection, I recommend Jablonski’s papers in particular.

    Hickman,

    “Almost a Modern Synthesis + … additions to the existing synthesis: which from what you’ve mentioned would need a revamp of the mathematical formula underpinning the original MS.”

    Very much so. Standard population genetics theory doesn’t really know what to do with most of the stuff I mentioned. Not exactly incompatible with it, simply insufficient.

    david,

    “Which it didn’t do with the rise of molecular biology in the 1960’s? I don’t see where there is a paradigm shift in the relevant foundations”

    Molecular biology is related to, but not the same as evolutionary biology. The revolution in the first field hardly touched the second one for decades. And, once again, I’m not talking about paradigm shifts.

    “group selection was always floating around, adaptiveness and plasticity too”

    Yes, but the first one had a number of false starts, and the latter was pretty much ignored for almost a century. If by adaptiveness you mean evolvability, no, that’s a novel concept.

    ej,

    “the “yes” (EES) faction is looking to develop a broader, more inclusive theory of evolution than is currently accepted as standard. This has academic-political implications (concerning what research gets grants and valorization in journals), and these are by no means negative motivations, it’s how the academy operates”

    Precisely.

    “also agree with the “no” faction that the empirical research must be there to validate it, and encourage further work”

    It has been there for a while, and it keeps getting better.

    John,

    “you didn’t deal enough with the middle party in your trinity of “Mendelism, Mutationism and (neo)Darwinism”. Blind mutation to get a specific 3 billion sequence? … I’m with Hoyle and his 747 in a whirlwind.”

    All I can say there is that no serious biologist is. If you want, google a review I did of one of Hoyle’s books in Skeptic magazine, a number of years ago, I present a more extensive argument there.

    Bill,

    “Mutations to non-coding DNA, however, don’t change the structure of proteins, they change the amount of a protein that is produced.”

    I know, this is part of the ongoing discussion about structural vs regulatory genes, where the defenders of the MS (e.g., Coyne) find themselves on what I see the (eventually) loosing side (as opposed to, say, Sean Carroll, the evodevo guy, not the physicist).

    “It is now clear that substantial changes in protein structure are rare, and that the bulk of evolution involves changes in the amounts of proteins that are expressed.”

    Completely agreed. I said this much in my first book, co-authored with Carl Schlichting back in 1998. Time flies…

    Patrice,

    “Massimo, do you mean that evolution theory is the wrong theory?”

    No, I mean that Lamarck’s theory of evolution was the wrong theory of evolution.

    “Lamarck’s main body of work is just about establishing “evolution””

    No, if it were just that then we could definitely credit the pre-Socratic atomists with the idea.

    “Because, being in the Anglo-Saxon realm, we have to be Anglo nationalists? Is it all about tribalism?”

    As others have said, I tend to consider this a type of unfounded postmodernism and I don’t buy it. Sure, national rivalries do play a role in science, but as far as I can see from my understanding of the history of the field, it’s a minor one, and it gets rapidly overwhelmed by actual progress, regardless of the nationality of the scientist that makes that progress possible.

    “attributing “evolution” to Darwin, instead of Lamarck, constitutes scientific, and philosophical fraud. For want of a nicer way to put it.”

    Actually, there are plenty of nicer way to put it. But the accusation is obviously false.

    Coel,

    “Well, on this one we can’t bow to the biologists’ consensus, since there are plenty of notable biologists on both sides. [Of course no non-physicists would ever have an opinion on, say, string theory, would they? :-)”

    As you know, I regard philosophers of science as in the business of having opinions about scientific theories, so…

    “The substantive issue seems to be over whether new developments are adding to the Modern Synthesis or over-turning parts of it.”

    No, that’s not it at all. I’m clearly in the “adding” camp, but gene centrism is being cut to size to its proper importance, and that, in my mind, is a very significant change.

    “Can you give some more commentary on which of those ideas are wrong and why?”

    Sure, I wrote two books (Phenotypic Evolution: A Reaction Norm Perspective, and Phenotypic Plasticity: Beyond Nature and Nurture) about it, and co-edited the one about the extended synthesis with Gerd Muller.

    “So the population biologist was thinking about generational timescales, and the paleontologist about geological timescales?”

    Obviously yes, but it’s also, I think, that people like Doug simply don’t take, and have never taken, paleontology seriously. They consider population genetics to provide the fundamental structure of evolutionary theory, a structure that I think is woefully incomplete.

    Darko,

    “my point about tranponsable elements would be to see them as enablers of novelties”

    Sure, but I think it’s more helpful to frame that within the general debate on regulatory evolution.

    “Here are lurking two interesting philosophical questions: what is individual and what is level natural selection is acting on”

    Yep, and sure enough there is a huge philosophical literature on both.

    astoltzfus,

    “IMHO this debate is confused and poorly framed”

    Well, I guess we’ll have to agree to disagree on that.

    “Scientific theories have structure and coherence. They are falsifiable.”

    No, they aren’t. Falsifiability has gone out the window since Kuhn, and it has really never been in the game at the theory level since at least Duhem.

    “This bait-and-switch is sustained by a lot of bad historiography– not Pigliucci’s fault because 90 % of available historiography was written under the influence of Mayr’s MS kool-aid.”

    I’d like to get a bit more credit than that. I have read plenty of non-Mayr based historiographies, and I have been pretty critical of Mayr when he was still alive. (I think I have the distinction of having been the last one to negatively review a book of his, which he published when he turned 100.)

    “Mendelism brought down Darwin’s theory, not because Mendelians were too stupid to synthesize mutation, heredity and selection, but because Darwin’s original theory was non-Mendelian”

    That is simply incorrect. Darwin’s theory made no claim at all about the mechanism of heredity.

    Arno,

    “I would think that the ‘yes-answer’ to this latter question implies at least the existence of phenomena that are superficially at odds with current evolutionary theory”

    Well, it depends on what you mean by “superficially.” Group selection, plasticity, evolvability, epigenetic inheritance and niche construction, to name a few, are indeed *superficially* at odds with the MS. But, I claim, that is due to the incompleteness of the latter, not to any fundamental incompatibility. Similar case to the superficial (as it turned out) incompatibility btw the Darwinism and Mendelism that brought about the MS itself.

    “I wonder how many parties there are”

    There is indeed a spectrum, as should be clear from my distancing from Jablonka. Incidentally, one of the co-authors of the no-paper in Nature, Greg Wray, was actually present at my workshop on the Extended Synthesis a few years ago, and contributed to the resulting volume for MIT Press.

    “Massimo repeats his belief that most of the molecular stuff is invisible from the perspective of evolutionary theory. This is obviously not the view of the people I just mentioned.”

    Actually, I would distinguish further. For instance, Lynch is, in my mind, one the most clear no-sayers, so much so that I took him to task in print for a scathing commentary he wrote for PNAS. And Gould certainly wasn’t interested in the molecular stuff, at the least not in any way like Lynch.

    But you are correct that a reduced scope of natural selection is another prong of the ES, which I should have made more clear in my responses.

    “we need a theory of what counts as a theory, what as a change of theory, what as an extension of the theory, and of how to asses the importance of certain changes etc. Such a theory is lacking in Massimo’s account.”

    Indeed, I didn’t write a full book on this. Actually I did: with Jonathan Kaplan, Making Sense of Evolution (Chicago Press). And of course there are very interesting accounts of these matters in the philosophical literature (which people like Gould and Lewontin do read).

    “Several commentators refer to the alleged extended synthesis as a Kuhnian revolution. … I see no reason to take this view seriously.”

    Neither do I, and I have argued this at length in print.

    “I don’t understand Massimo’s intuitions about what changes are important.”

    I was basing my judgment on who and what has made an impact on textbooks. By the way, a good analysis of how textbooks in evolutionary biology have (or have not) changed over time, tracking theory change and attitudes, is provided by Alan Love in my co-edited volume for MIT Press.

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  45. Massimo

    My take is that David is indeed exaggerating, while some of his detractors are minimizing. The best take on multi-level selection is provided in the book by Samir Okasha that I cited in the notes.

    Ah, thanks for the clarification. I will have to check out Samir’s book, I’m just about finished reading David’s.

    Coel

    It would be wrong to see that issue as young upstarts versus old-guard fuddy-duddies. The idea of group selection is pretty old and many of the leading advocates (e.g. D.S. Wilson, E.O Wilson, Nowak) are not exactly spring chickens themselves. It could be that the majority are correct in seeing group-selection as a minor part of the picture.

    I was not actually referring to the “old guard” actually being old but as the “conservative” group resistance to change. However, judging from Massimo’s comments in his essay (that most people inviting him to talk about ES are graduate students) I would be willing to bet that age does have a lot to do with it. Sure there are exceptions (Wilsons and others) but that doesn’t mean the majority of the individuals supporting ES are older. Of course I’m just guessing here based on my general impression in how change occurs in my field and from my interactions with colleagues in other fields. I’d be open to evidence to the contrary.

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  46. Besides, shall we retroactively apply your rule to the Modern Synthesis too? Huxley named it while he was in the midst of it, not fifty years later…

    Yes, I guess if I had been in the midst of it I would also have taken a “get on with the work already, who cares about labels” approach to it.

    Thinking of evolution only “in the long run” is a serious mistake. Evolution happens from one generation to the next, and natural selection works at that level too.

    That is right, and in fact I would say that selection mostly works at that level, but it is not quite the point. If the stuff doesn’t get fixed in the genome then it cannot have a hereditary influence over long time periods, so it is more of an accident than anything else. Conversely, if it is about genes providing the ability to have epigenetics happen, then it is just a normal trait under normal selection. Or in other words, it doesn’t seem revolutionary enough to warrant a New Evolutionary Theory ™.

    Right, and that’s why so many people have so ferociously attacked Eldridge and Gould for decades now.

    As far as I can tell much of the controversy was based on misunderstandings, poor communication and conflation with saltationism. The fact remains that evolution appears to work glacially slowly through changes in allele frequencies in populations, and things only seem sudden because we look at a woefully incomplete fossil record across very deep time.

    Then again, I don’t claim to be an expert on this issue, I mostly don’t see what would be so big about it even if it applied to most lineages most of the time. It seems quite logical that most selection would be stabilising / purifying most of the time anyway.

    Not really, you may want to read some of the literature on species selection, I recommend Jablonski’s papers in particular.

    I looked into the paper you gave as a reference, and otherwise relied on what I got out of discussion with colleagues. While I am somewhat agnostic, not everybody is convinced that species selection and group selection actually make sense.

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