With Carl Gegenbaur and Ernst Haeckel, inspiredby Darwin and the cell theory, comparativeanatomy and embryology became established andflourished in Jena. This tradition wascontinued and developed further with new ideasand methods devised by some of Haeckelsstudents. This first period of innovative workin evolutionary morphology was followed byperiods of crisis and even a disintegration ofthe discipline in the early twentieth century.This stagnation was caused by a lack ofinterest among morphologists in Mendeliangenetics, and uncertainty about the mechanismsof evolution. Idealistic morphology was stillinfluental in (...) Germany, which prevented a fullappreciation of the importance of Darwinstheory of natural selection for comparativemorphology. Evolutionary morphology andembryology failed to contribute significantlyto the modern synthesis of evolutionarybiology, thereby probably delaying theintegration of developmental biology intomodern evolutionary biology. However, Haeckelsstudent Oscar Hertwig, as well as Victor Franzand Alexej N. Sewertzoff from a youngergeneration, all tried to forge their ownsynthetic approaches in which (inspired byHaeckels work) embryology played an importantrole. Important for all three researchers wereattempts to refine, and sometimes redefine, thebiogenetic law, and to find new scientificexplanations for it (and for the manyexceptions to it). Their research was latermore or less forgotten, and had littleinfluence on the architects of the modernsynthesis. As the relationship betweenevolutionary and developmental biology is nowagain rising in importance in the form ofEvo-Devo, we would like to draw attention tohow this earlier research tradition grappledwith similar questions to those now on theagenda, albeit from sometimes quite differentperspectives. (shrink)
Evo-Devo exhibits a plurality of scientific “cultures” of practice and theory. When are the cultures acting—individually or collectively—in ways that actually move research forward, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? This chapter identifies six cultures – three /styles/ (mathematical modeling, mechanism, and history) and three /paradigms/ (adaptationism, structuralism, and cladism). The key assumptions standing behind, under, or within each of these cultures are explored. Characterizing the internal structure (...) of the cultures is necessary for understanding how they collaborate or compete, and how they are fragmented or integrated, in the rich interdisciplinary /trading zone/ (Galison 1997) of Evo-Devo. Evo-Devo is an important example of how science can progress through a radical plurality of perspectives and cultures. (shrink)
_The emerging discipline of evolutionary developmental biology has opened up many new _ _lines of investigation into morphological evolution. Here I explore how two of the core _ _theoretical concepts in ‘evo-devo’ – modularity and homology – apply to evolutionary _ _psychology. I distinguish three sorts of module - developmental, functional and mental _ _modules and argue that mental modules need only be ‘virtual’ functional modules. _ _Evolutionary psychologists have argued that separate mental modules are solutions to _ _separate (...) evolutionary problems. I argue that the structure of developmental modules in _ _an organism helps determine what counts as a separate evolutionary problem for that _ _organism. I suggest that homology as an organizing principle for research in _ _evolutionary psychology, has been severely neglected in favor of analogy (adaptive _ _function). I consider some arguments suggesting that determining homology is less _ _epistemically demanding than determining adaptive function and argue that _ _psychological categories defined by homology are, in fact, more suitable objects of _ _psychological - and particularly neuropsychological - investigation than categories _ _defined by analogy. _. (shrink)
This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about fins and limbs compare with (...) modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for fins as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty. (shrink)
The study of evolutionary developmental biology (“evo‐devo”) has recently experienced a dramatic surge in popularity among researchers and theorists concerned with evolution. However, some biologists and philosophers remain skeptical of the claims of evo‐devo. This paper discusses and responds to the recent high profile criticisms of evo‐devo presented by biologists Hopi E. Hoekstra and Jerry A. Coyne. I argue that their objections are unconvincing. Indeed, empirical research supports the main tenets of evo‐devo, including the claim that morphological evolution is the (...) result of cis ‐regulatory change and the distinction that evo‐devo draws between morphological and physiological traits. *Received January 2008; revised March 2009. †To contact the author, please write to: Department of Philosophy, University of Cincinnati, Cincinnati, OH 45221; e‐mail: email@example.com. (shrink)
Evolutionary developmental biology (“evo-devo”) may provide insights and new methods for studies of cognition and cultural evolution. For example, I propose using cultural selection and individual learning to examine constraints on cultural evolution. Modularity, the idea that traits vary independently, can facilitate evolution (increase “evolvability”), because evolution can act on one trait without disrupting another. I explore links between cognitive modularity, evolutionary modularity, and cultural evolvability. (Published Online November 9 2006).
Sound comparative psychology and modern evolutionary and developmental biology (often called evo-devo) emphasize powerful effects of developmental conditions on the expression of genetic endowment. Both demand that evolutionary theorists recognize these effects. Instead, Tomasello et al. compares studies of normal human children with studies of chimpanzees reared and maintained in cognitively deprived conditions, while ignoring studies of chimpanzees in cognitively appropriate environments.
Although the construction of neo-Darwinism grew out of Thomas Hunt Morgan's melding of Darwinism and Mendelism, his evidence did not soley support a model of gradual change. To the contrary, he was confronted with observations that could have led him to a more "evo-devo" understanding of the emergence of novel features. Indeed, since Morgan was an embryologist before he became a fruit-fly geneticist, one would have predicted that the combination of these two lines of research would have resulted in (...) early formulations of concepts relevant to evolutionary developmental biology. It is thus of interest to review Morgan's thought processes and arguments for at first rejecting both Darwinism and Mendelism, and then for later dismissing data that would have yielded a model of rapid morphological change in favor of a model of change based on the accumulation of minor mutations and their morphological consequences. (shrink)
In the evolutionary biology of the Modern Synthesis the study of patterns refers to how to identify and systematise order in lineages, looking for hierarchies or for branching/splitting events in the tree of life, whereas the resulting order is supposed to be due to underlying processes or mechanisms. But patterns and processes play distinct roles in evo-devo: four different views on the role of patterns and processes in descriptions and explanations of development and evolution: A) transformational; B) generative; C) (...) processual; and D) complex are reviewed in this paper. Then, this discussion is related to two issues in evo-devo: homology and variation. (shrink)
This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about ﬁns and limbs compare with (...) modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for ﬁns as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty. (shrink)
Ever since Darwin a great deal of the conceptual history of biology may be read as a struggle between two philosophical positions: reductionism and holism. On the one hand, we have the reductionist claim that evolution has to be understood in terms of changes at the fundamental causal level of the gene. As Richard Dawkins famously put it, organisms are just ‘lumbering robots’ in the service of their genetic masters. On the other hand, there is a long holistic tradition that (...) focuses on the complexity of developmental systems, on the non-linearity of gene– environment interactions, and on multi-level selective processes to argue that the full story of biology is a bit more complicated than that. Reductionism can marshal on its behalf the spectacular successes of genetics and molecular biology throughout the 20th and 21st centuries. Holism has built on the development of entirely new disciplines and conceptual frameworks over the past few decades, including evo-devo and phenotypic plasticity. Yet, a number of biologists are still actively looking for a way out of the reductionism–holism counterposition, often mentioning the word ‘emergence’ as a way to deal with the conundrum. This paper briefly examines the philosophical history of the concept of emergence, distinguishes between epistemic and ontological accounts of it, and comments on conceptions of emergence that can actually be useful for practising evolutionary biologists. (shrink)
Like Laland et al., I think Mayr’s distinction is problematic, but I identify a further problem with it. I argue that Mayr’s distinction is a false dichotomy, and obscures an important question about evolutionary change. I show how this question, once revealed, sheds light on some debates in evo-devo that Laland et al.’s analysis cannot, and suggest that it provides a different view about how future integration between biological disciplines might proceed.
In the Modern Synthesis the study of patterns refers to how to identify and systematize order in lineages (description), attributed to underlying processes or mechanisms (explanation). But patterns and processes play distinct roles in evodevo. In this paper we (1) distinguish three different views (the transformational, the morphogenetic and the process approach) according to the role they play in the description and explanation of development and evolution, and (2) relate this discussion to the issues of homology and variation.
In the fall of 1990 I had just began my doc- toral studies at the University of Connecticut. Freshly arrived from Italy, I came to the United States to work with Carl Schlichting on something to do with phenotypic plastic- ity. I spent most of that semester discussing with other graduate students what I thought was a momentous paper by Mary Jane West- Eberhard (1989) in the Annual Review of Ecol- ogy and Systematics. That paper, entitled Phe- notypic Plasticity and (...) the Origins of Diversity, was a (quite lengthy) forerunner of the (also quite bulky) book I am reviewing now. Like the paper, this volume has the potential to be momentous in the development of our ideas on phenotypic evolution. (shrink)
In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary questions, a belief that has (1) hindered (...) progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation. (shrink)
Many natural and biological phenomena can be depicted as networks. Theoretical and empirical analyses of networks have become prevalent. I discuss theoretical biases involved in the delineation of biological networks. The network perspective is shown to dissolve the distinction between regulatory architecture and regulatory state, consistent with the theoretical impossibility of distinguishing a priori between “program” and “data”. The evolutionary significance of the dynamics of trans-generational and inter-organism regulatory networks is explored and implications are presented for understanding the evolution of (...) the biological categories development-heredity; plasticity-evolvability; and epigenetic-genetic. (shrink)
Traditional accounts of the role of learning in evolution have concentrated upon its capacity as a source of fitness to individuals. In this paper I use a case study from invasive species biology—the role of conditioned taste aversion in mitigating the impact of cane toads on the native species of Northern Australia—to highlight a role for learning beyond this—as a source of evolvability to populations. This has two benefits. First, it highlights an otherwise under-appreciated role for learning in evolution that (...) does not rely on social learning as an inheritance channel nor “special” evolutionary processes such as genetic accommodation (both of which many are skeptical about). Second, and more significantly, it makes clear important and interesting parallels between learning and exploratory behaviour in development. These parallels motivate the applicability of results from existing research into learning and learning evolution to our understanding of the evolution of evolvability more generally. (shrink)
In September 2008, 10 years after the untimely death of Pere Alberch (1954–1998), the 20th Altenberg Workshop in Theoretical Biology gathered a group of Pere’s students, col- laborators, and colleagues (Figure 1) to celebrate his contribu- tions to the origins of EvoDevo. Hosted by the Konrad Lorenz Institute for Evolution and Cognition Research (KLI) outside Vienna, the group met for two days of discussion. The meeting was organized in tandem with a congress held in May 2008 at the Cavanilles Institute (...) for Biodiversity and Evolutionary Biology (ICBiBE) in Valencia, Spain. The talks at the KLI were equal parts: nostalgic remembrance, excitement over new ways of thinking about old problems, and an unrepressed vitriol against the resurgence of reductionist thinking in EvoDevo. Here we highlight some of the key aspects of Pere’s life and work that informed and infused the talks. (shrink)
The science of genetics is undergoing a paradigm shift. Recent discoveries, including the activity of retrotransposons, the extent of copy number variations, somatic and chromosomal mosaicism, and the nature of the epigenome as a regulator of DNA expressivity, are challenging a series of dogmas concerning the nature of the genome and the relationship between genotype and phenotype. According to three widely held dogmas, DNA is the unchanging template of heredity, is identical in all the cells and tissues of the body, (...) and is the sole agent of inheritance. Rather than being an unchanging template, DNA appears subject to a good deal of environmentally induced change. Instead of identical DNA in all the cells of the body, somatic mosaicism appears to be the normal human condition. And DNA can no longer be considered the sole agent of inheritance. We now know that the epigenome, which regulates gene expressivity, can be inherited via the germline. These developments are particularly significant for behavior genetics for at least three reasons: First, epigenetic regulation, DNA variability, and somatic mosaicism appear to be particularly prevalent in the human brain and probably are involved in much of human behavior; second, they have important implications for the validity of heritability and gene association studies, the methodologies that largely define the discipline of behavior genetics; and third, they appear to play a critical role in development during the perinatal period and, in particular, in enabling phenotypic plasticity in offspring. I examine one of the central claims to emerge from the use of heritability studies in the behavioral sciences, the principle of minimal shared maternal effects, in light of the growing awareness that the maternal perinatal environment is a critical venue for the exercise of adaptive phenotypic plasticity. This consideration has important implications for both developmental and evolutionary biology. (shrink)
American biologists in the late nineteenth century pioneered the descriptive-comparative study of all cell divisions from zygote to gastrulation -- the cell lineage. Data from cell lineages were crucial to evolutionary and developmental questions of the day. One of the main questions was the ultimate causation of developmental patterns -- historical or mechanical. E. B. Wilson's groundbreaking lineage work on the polychaete worm Nereis in 1892 set the stage for (1) an attack on Haeckel's phylogenetic-historical notion of recapitulation and (2) (...) support for mechanistic explanations of cleavage patterns. As more lineage work -- especially Lillie's work on "Unio" and Conklin's on "Crepidula" -- became available in the mid-late 1890s, mechanism was tempered with more evolutionary, homology-based views. However, as I show by focusing on three major issues -- homology, body plans and life history -- these views were primarily based on the precocious segregation and prospective significance -- what the cell became not what it was. Even on issues like adaptation, most lineagists argued teleologically from the adult backward. Most cell lineage workers, by 1900, were to varying degrees mechanist/experimentalist and recapitulationist simultaneously. The exception was E. G. Conklin, whose views were more akin to a Darwinian evolutionist than either mechanist or recapitulationist. Lineage work eventually declined and by 1907 published accounts of new lineages had basically stopped. I argue that established workers and younger researchers stopped wanting to take on cell lineage projects because the general patterns were the same for all the spiralians while the specifics showed too much variation. It was hard to theoretically encompass or analyze the minutiae of variation in a recapitulationist or mechanist framework. The only established worker who continued to do comparative lineage studies was E. G. Conklin, perhaps because the variation could best be accommodated by Darwinian evolution. (shrink)
Evolutionary theory is undergoing an intense period of discussion and reevaluation. This, contrary to the misleading claims of creationists and other pseudoscientists, is no harbinger of a crisis but rather the opposite: the field is expanding dramatically in terms of both empirical discoveries and new ideas. In this essay I briefly trace the conceptual history of evolutionary theory from Darwinism to neo-Darwinism, and from the Modern Synthesis to what I refer to as the Extended Synthesis, a more inclusive conceptual framework (...) containing among others evo–devo, an expanded theory of heredity, elements of complexity theory, ideas about evolvability, and a reevaluation of levels of selection. I argue that evolutionary biology has never seen a paradigm shift, in the philosophical sense of the term, except when it moved from natural theology to empirical science in the middle of the 19th century. The Extended Synthesis, accordingly, is an expansion of the Modern Synthesis of the 1930s and 1940s, and one that—like its predecessor—will probably take decades to complete. (shrink)
Evolutionary developmental biology (Evo-Devo) is a new and rapidly developing field of biology which focuses on questions in the intersection of evolution and development and has been seen by many as a potential synthesis of these two fields. This synthesis is the topic of the books reviewed here. Integrating Evolution and Development (edited by Roger Sansom and Robert Brandon), is a collection of papers on conceptual issues in Evo-Devo, while From Embryology to Evo-Devo (edited by Manfred Laubichler (...) and Jane Maienschein) is a history of the problem of the relations between ontogeny and phylogeny. (shrink)
In a now classic paper published in 1991, Alberch introduced the concept of genotype–phenotype (G!P) mapping to provide a framework for a more sophisticated discussion of the integration between genetics and developmental biology that was then available. The advent of evo-devo first and of the genomic era later would seem to have superseded talk of transitions in phenotypic space and the like, central to Alberch’s approach. On the contrary, this paper shows that recent empirical and theoretical advances have only (...) sharpened the need for a different conceptual treat- ment of how phenotypes are produced. Old-fashioned metaphors like genetic blueprint and genetic programme are not only woefully inadequate but positively misleading about the nature of G!P, and are being replaced by an algorithmic approach emerging from the study of a variety of actual G!P maps. These include RNA folding, protein function and the study of evolvable soft- ware. Some generalities are emerging from these disparate fields of analysis, and I suggest that the concept of ‘developmental encoding’ (as opposed to the classical one of genetic encoding) provides a promising computational–theoretical underpinning to coherently integrate ideas on evolvability, modularity and robustness and foster a fruitful framing of the G!P mapping problem. (shrink)
The concept of developmental constraint was at the heart of developmental approaches to evolution of the 1980s. While this idea was widely used to criticize neo-Darwinian evolutionary theory, critique does not yield an alternative framework that offers evolutionary explanations. In current Evo-devo the concept of constraint is of minor importance, whereas notions as evolvability are at the center of attention. The latter clearly defines an explanatory agenda for evolutionary research, so that one could view the historical shift from ‘developmental (...) constraint’ towards ‘evolvability’ as the move from a concept that is a mere tool of criticism to a concept that establishes a positive explanatory project. However, by taking a look at how the concept of constraint was employed in the 1980s, I argue that developmental constraint was not just seen as restricting possibilities (‘constraining’), but also as facilitating morphological change in several ways. Accounting for macroevolutionary transformation and the origin of novel form was an aim of these developmental approaches to evolution. Thus, the concept of developmental constraint was part of a positive explanatory agenda long before the advent of Evo-devo as a genuine scientific discipline. In the 1980s, despite the lack of a clear disciplinary identity, this concept coordinated research among paleontologists, morphologists, and developmentally inclined evolutionary biologists. I discuss the different functions that scientific concepts can have, highlighting that instead of classifying or explaining natural phenomena, concepts such as ‘developmental constraint’ and ‘evolvability’ are more important in setting explanatory agendas so as to provide intellectual coherence to scientific approaches. The essay concludes with a puzzle about how to conceptually distinguish evolvability and selection. (shrink)
This volume joins a growing list of books, monographs, and proceedings from scientific meetings that attempt to consolidate the wide spectrum of approaches emphasizing the role of development in evolution into a coherent and productive synthesis, often called evo-devo. Evo-devo is seen as a replacement or amendment of the modern synthesis that has dominated the field of evolution since the 1940s and which, as even its architects confessed, was fundamentally incomplete because development remained outside its theoretical framework (Mayr (...) and Provine 1980).As the volume attests, there is now a strong feeling that the time is ripe for the onsolidation of evo-devo, and that the field is mature enough so that mapping the theoretical terrain and experimental approaches is both feasible and scientifically productive. Now is an appropriate time to try to weave the strands of reasoning leading to the developmental perspective and offer a synthesis. (shrink)
In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability (...) offers a robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology3.1 The explanatory targets of evolutionary biology3.2 Selection-based explanations3.3 Lineage explanations3.4 Evolvability-based explanations3.5 What properties must evolvability have?4 What Evolvability Really Is4.1 Making sense of ft4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)
Evolutionary developmental biology (evo-devo) offers both an account of developmental processes and also new integrative frameworks for analyzing interactions between development and evolution. Biologists and philosophers are keen on evo-devo in part because it appears to offer a comfort zone between, on the one hand, what some take to be the relative inability of mainstream evolutionary biology to integrate a developmental perspective; and, on the other hand, what some take to be more intractable syntheses of development and evolution. (...) In this article, I outline core concerns of evo-devo, distinguish theoretical and practical variants, and counter Sterelny's recent argument that evo-devo's attention to development, while important, offers no significant challenge to evolutionary theory as we know it. (shrink)
John Reiss is a practicing evolutionary biologist (herpetology) who by his own account happened to be in the right place (Harvard’s Museum of Comparative Zoology) at the right time (the 1980s) to hear echoes of the debate about sociobiology that had been raging there between E. O. Wilson and, on the other side, Stephen Jay Gould and Richard Lewontin (xiv). Reiss is not concerned with sociobiology, at least in this book, but with the adaptationism that Gould and Lewontin saw in (...) the sociobiologists’ approach to cooperative behavior. At Harvard, Reiss was guided by Pere Alberch, in whose laboratory Gould’s stress on developmental constraints was being transformed into a now influential version of the Evo-devo movement (xiv, 327). On Alberch’s view, which Reiss accepts, variation in the rate, timing, placement, and intensity of gene products during the ontogenetic process, rather than mutation in structural genes, constitutes the proximate source of the phenotypic variation on which natural election works (327-29). Reiss does not think that Evo-devo, at least as he construes it, does away with natural selection. Rather, he seeks to identify the role played by selection in retaining or eliminating the variation generated in the developmental process. Selection, he argues, enables organisms, populations, species, and other lineages to maintain the presumptively adapted conditions of existence to which their very persistence already testifies. “Adaptedness,” Reiss writes, “is not a product of evolution; it is a condition for evolution” (22). He thinks that this fact, as he takes it to be, belies the adaptationist assumption that organisms are collections of independently optimal adaptations that arise by way of concerted spurts of directional selection. “It is a mistake,” he writes, “to atomize organisms and to explain each part as the solution of a problem raised by the environment” (295). (shrink)
Jackendoff's “mentalistic” semantics looks more radical than it is. It can best be understood as a necessary corrective to the traditional oversimplification that holds that psychological variation “cancels out” on the path from word to world. This reform parallels the “evo-devo” reform in evolutionary biology.
Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the (...) need for a broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (shrink)