Abstract
Inferences from dependencies in development, or from conserved traits, structures, or genes, have become a central and ubiquitous tool in evolutionary developmental biology. An element with many downstream consequences of its operation in an adaptive system should be conserved because its deletion or change would likely have massively deleterious effects. Thus, substantial observed dependencies should predict conservation, and vice versa. Dependencies and conservation are both matters of degree, generating an ordering of rates of evolutionary change. I trace four independent sources for this idea with subtle differences in their application. The origins of the use of genetic pleiotropy to predict conservation is complex and progressed in multiple stages, with partial awareness emerging in the 1960s, and grew erratically in use without becoming a major theme until the 1990s. Since 2000 it has grown to become an essential tool and applied in an expanding number of ways. I distinguish and illustrate “top-down” and “bottom up” approaches, though people now working from one direction often recruit collaborators from the other. Other approaches have had diverse inspirations and points of initial application. Rupert Riedl’s notion of “burden” was applied primarily to morphological traits, with an elaborated account of organizational factors modulating entrenchment. Wallace Arthur explored dependency in “morphogenetic trees,” utilizing as his inspiration cellular descent trees in metazoans. Bill Wimsatt looked for “generative entrenchment” in factors both internal and external to the developing system to analyze innate behavior. Entrenchment was later extended to analyze development in Drosophila and to construct multi-locus population genetic models of gene control networks, yielding new results that illuminate relations between micro and meso-evolution. Applications of generative entrenchment to cultural and technological evolution are also sketched, where development of individuals and other social entities must play a central role.
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Notes
- 1.
This was a remolding of the old committee on mathematical biology under the influence of Richard Lewontin and leadership (as newly recruited chair) of Jack Cowan. Under Rashevsky’s leadership that committee has had a far if sometimes eccentric reach in the university and the world. Such diverse figures as linguist Eric Hamp and sociologist-cognitive-computer-scientist-organization-theorist-economist Herbert Simon took or sat in on courses there. The new department was both more focused and more biologically based (though still with a broad reach), and development was one of the key elements of that focus.
- 2.
Indeed, even earlier: as an undergraduate at Cornell, I took Frank Rosenblatt’s course (1962) in “Brain Models and Mechanisms.” Rosenblatt (1962) created the first well-developed connectionist theories of brain organization. His was both a populational and developmental theory in which he aimed to model systems that would, with experiential feedback, learn to recognize patterns. Rosenblatt argued that there wasn’t sufficient genetic information to specify neural connections completely, so rather than seeking to model adult abilities with networks specified in detail, he sought to model classes of randomly connected networks (specified only at a molar level, characterized by probability distributions for connections in a multi-level framework) that could develop the relevant capacities through reinforcement learning. In this he anticipated Kauffman’s approach to looking for generic properties of classes of randomly organized gene-control networks, but I was thus also primed for the evolutionary relevance of development. In graduate school at the University of Pittsburgh (Pitt) I took a historical and modern course on developmental biology taught by Stanley Shostak in 1967. I first met and established ongoing contact with Steve Gould in the summer of 1968 at the wedding of his undergrad roommate, Carl Putz, who was my roommate at Pitt. Before the 1981 Dahlem conference, I had also interacted with John Bonner, and benefitted from Dave Raup as a colleague. Rudy Raff, Günter Wagner, Gerd Müller, and Steve Stearns have all influenced my perspectives in different ways since.
- 3.
I feared earlier that I had “explained” a rather crude recapitulation with the “developmental lock”—a problem that to my mind made it unpublishable, and was delighted to realize that instead I had explained von Baer’s laws. Gould’s paper for this 1974 conference at the American Academy of Arts and Sciences (convened by sociologist Talcott Parsons) was an early sketch of the view developed later in Ontogeny and Phylogeny (1977). My series-parallel reliability model anticipated that of Oster and Wilson (1977) for ant colonies. Wilson, Lewontin, and Mayr were also there. Unfortunately, the conference papers were not published.
- 4.
- 5.
Aside from the work delineated below on entrenchment, this more interactionist idea emerged simultaneously with but independently from “developmental systems theory” [DST] (Oyama 1985), with which it shares many assumptions. But DST needs an engine to drive it: DST urges that we take the whole developmental system into account, without privileging genes, but includes no mechanisms capable of driving evolutionary processes (Wimsatt 2001). Self-organization (widely cited in this literature) and generative entrenchment can both do this, and should ultimately serve complementary roles in doing so (as argued in Wimsatt 1986, 2001).
- 6.
This is extensively documented in Frietson Galis’s work, discussed at length below.
- 7.
One might say that it was almost an embarrassment to population genetics, since pleiotropy, like epistasis would have seemed to reduce the likelihood of additive (and thus heritable) gene effects. Wright (1968) was unusual in noting pleiotropy’s probable role in the severity of deleterious mutations involving polydactyly in his guinea pigs.
- 8.
Morgan et al. (1925) review the properties and expression of over 500 mutations. See Kohler (1994) for a broader discussion of their methodology. Dobzhansky’s immensely influential (1941), in which he used different patterning of giant salivary gland chromosomes indicating inversions and translocations to distinguish Drosophila species in nature, together with the fact that hybridization between species differing by inversions was usually severely deleterious, reinforced hostility to views like those of Goldschmidt who sought the (very occasional) hopeful monster as a product of major mutational changes.
- 9.
Curiously, no one appeared to juxtapose Darwin’s abhorrence of macromutations with the simultaneous effects of multiple mutational changes to get a pleiotropy or entrenchment based account of Dollo’s law.
- 10.
- 11.
Hadorn introduced the distinction between highly interactive and pleiotropic, which became important in distinguishing the impact of mutations acting at the phylotypic stage (Sander 1983) with others that were just very pleiotropic. The distinction (as developed, for example, by Galis) is between a mutation which has multiple but characteristic specific effects, each with relatively high penetrance, and one which may have a multiplicity of substantial effects of diverse kinds with substantial variation in specific effects from case to case.
- 12.
Williams’ very influential (1966) uses the severity of fitness reduction of bithorax mutants to scoff at Waddington’s experiments with genetic assimilation of bithorax as indicating a possible mechanism to explain the evolutionary incorporation of larger innovations.
- 13.
The idea of a phylotypic stage as being a “neck” in an “hourglass” with greater variation both earlier and later in development dates back to Seidel (1960), but did not become a matter of focus until Sander (1983) and Raff (1996). This is somewhat of a puzzle. Both are frequently cited, Sander is cited more than three times as frequently as Seidel. This could be due to the greater visibility of Sander’s paper: in English, rather than German, and in a very visible conference volume when interest in the relation between evolution and development was blossoming.
- 14.
The phylotypic stage attracted attention first, but then the “hourglass pattern” itself became a topic of dispute with those who expected a more cone-like pattern, as well as those who saw neither (e.g., Richardson et al. 1997). Increasing evidence for the “hourglass” has emerged over the past 15 years (see Raff 1996; Galis and Metz 2001; Kalinka et al. 2010; Kalinka and Tomancek 2012).
- 15.
This is more complicated because to understand the earliest divergence of the “hourglass” one needs to look at the variability of the niche of the zygote, or to the egg, a (possibly much more divergent) property of the reproductive adult, in response to different ecological situations, or to a more fully protective maternal effect stabilizing and facilitating a supportive environment as in placental mammals. Furthermore, unless some canalization of the phylotypic stage occurs, either the apparent similarities at that stage among diverse phyla are illusory (Richardson et al. 1997), or there must be some other cause for the convergence (e.g., Newman 2013).
- 16.
- 17.
I read this article when it came out in April of 1972, and used it in my biology course for the next 5 years. I think it influenced my developing ideas of generative entrenchment, and particularly the idea—not emphasized by Wallace Arthur at first—that entrenchment applied “all the way down.”
- 18.
- 19.
“General systems theory” should not be confused with systems biology. Ironically, the rise of systems biology has reinvigorated “systems” talk, as well as cybernetic language and intuitions.
- 20.
- 21.
It appears that population geneticists tended to treat a relatively improbable event as impossible—the same kind of mistake that emerges in ruling out (rare, positive) larger mutations. The surprising feature of their simulations was how many trajectories ending in positive solutions (~40 %) had gone through local minima of fitness (a possibility normally ignored by population geneticists), and many of the optimal solutions embodied suboptimal elements as necessary components.
- 22.
I first learned of Raff’s direct-developing echinoderms that apparently deleted the larval stage with relative impunity in his talk at the Field Museum in the spring of 1989. His richly characterized and analyzed counter-example to von Baer’s laws fascinated me, and helped to convince me that I should look to a systematic investigation of organizational features that could reduce entrenchment, or otherwise facilitate change of deeply entrenched features.
- 23.
Although written in the first person (appropriate from 1972 to 1984), this research in the 1980s and later involved close collaboration with two graduate students, Nick Rasmussen and especially Jeff Schank, who each made important contributions to how it developed, and later with Jim Griesemer (in the late 1980s and again after 2003).
- 24.
- 25.
Kauffman picked a high mutation rate so that he could get results in a reasonable amount of time with population sizes of 100 for the smaller circuits. He supposed that the inversely proportional relationship between circuit size increase and mutation rate decrease would preserve the qualitative conclusions. But the result was deeply paradoxical because then current estimates had suggested genome sizes of 100,000 genes, and data on amounts of heterozygosity in populations suggested that ~95 % of these should be preserved over time spans much longer than 1,000 generations. This was much larger than Kauffman’s results would suggest unless the vast majority were selectively neutral. But then how could they maintain adaptive organization against drift?
- 26.
- 27.
We are not yet in a position to say generally how often entrenchment is built on top of robustly determined features, or whether deep entrenchment acts to stimulate selection for increased robustness. Specific analyses to determine whether “deep” architectural features of circuits are maintained by robustness or entrenchment favor entrenchment (see, e.g., Galis et al. 2006), but these don’t answer the question in general.
- 28.
See Wimsatt and Schank (2004) for a more complete list of changes.
- 29.
This allows an important move towards realism, but still embodies a problematic simplification because the additional loss due to a gene will depend on what other losses occur. The idea of having fitness classes of different sizes was to allow for different numbers of downstream consequences, but with building this into the initially assigned fitnesses, the changed topology of the circuit after a mutation is no longer taken into account because we are no longer working directly from that topology to compute fitness contributions. Tracking the changing topologies was in principle possible, but immensely more demanding in computational and memory capacity and beyond what we could do in 1987–1989. Aldana et al. (2007), with far greater computational power, utilized the changing topologies.
- 30.
In 1987–1989, our simulations were limited to a maximum total data of 32 K bytes. So increasing the number of loci required that we avoid large populations.
- 31.
Mutations were always to connections with another gene in the genome. This definition made “back mutation” possible, with calculable rates. Connections on a predefined list were “good,” and all others were “bad.” One gets credit for a good connection, but additional identical good connections didn’t count. The crosshatched connections in the three cases have the same relative fitnesses (Fig. 17.8).
- 32.
Wagner and Zhang (2011) review evidence that pleiotropy, though widespread, is rarely massive, suggesting a usable degree of variational modularity for evolution.
- 33.
This argument supposes that the absolute fitness contributions of the alleles are constant. This is not necessarily the case, but is presumably true for a subclass of alleles.
- 34.
The metaphor analogizing genic selection to switching rowers in a shell (Dawkins 1976) to pick out the fittest in effect ignores the fact that the rowers must use a common shell. Analogously, genes are always embedded in an interactive genome, and must bear the consequences of their collective activity, so Dawkins’ metaphor is crucially flawed. He never considers the role of the boat in making good times when he switches the rowers!
- 35.
This was made possible by looking at circuits with 33, 65, 130, and 260 connections, with proportional distributions scaled to produce analogous results, so that only the relative position in the distribution of a connection class with given fitnesses has changed.
- 36.
Culture is in some ways like an ecosystem in which reproduction for most species is so dependent on rich, articulated structures in the environment that the notion of independent species breaks down. I came to appreciate this through economist Kenneth Boulding’s striking remark that, “A car is just an organism with an exceedingly complicated sex-life.”
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Acknowledgments
In addition to the formative influence of many of the people in this, or in the original 1981 Dahlem volume, I would like to acknowledge the help and influence of Alan Love for extended and penetrating editorial commentary, and the pleasures of co-teaching Evo-devo and working together. Günter Wagner pointed me to the work of Frietson Galis when I was in a quandary as to how to trace the growing influence of pleiotropy in assessments of evolutionary conservatism in the last 15 years. Thanks to Frietson herself for many illuminating articles and conversations. Scott Gilbert confirmed and amplified my intuitions on C.H. Waddington. The remarkably constructive and open atmosphere of the “Dahlem revisited” conference in Berlin in July 2010 owes a great deal to the participants, to the charge of its organizers to us, and to the hospitality of the Max Planck Institute for History of Science, and the support of the Minnesota Center for Philosophy of Science and the Konrad Lorenz Institute for the Study of Evolution, Development and Cognition. Finally, the Minnesota Center for Philosophy of Science and the support of my Winton Professorship in the College of Liberal Arts at the University of Minnesota during this period have been invaluable, and the environment as challenging and stimulating as anyone could wish.
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Wimsatt, W.C. (2015). Entrenchment as a Theoretical Tool in Evolutionary Developmental Biology. In: Love, A. (eds) Conceptual Change in Biology. Boston Studies in the Philosophy and History of Science, vol 307. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9412-1_17
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