Development and mechanistic explanation

Within modern philosophy of biology the topic of mechanistic explanation has become a central theme for critical discussion. The neo-mechanical philosophers have developed accounts that emphasize intervention and manipulation as the central epistemic tools that allow gaining epistemic access upon the mechanisms and have argued that the processes of inter-field integration across disciplines can be understood through the analysis of mechanisms spanning multiple levels. In this paper I revisit current proposals on mechanistic explanation in order to show some of their limitations when dealing with developmental mechanisms. I basically argue that (i) developmental mechanisms cannot be accommodated within a framework centered upon the mutual manipulation principle, (ii) the distinction between causal relations vs. constitutive relations cannot be easily demarcated within developmental biology and (iii) the notion of "part" underlying the neo-mechanical accounts on explanation is not suitable for developmental biology.     

Introduction—development and phylogeny of the arthropods: Darwin’s legacy

In the present essay, I first recall the genealogical concept of classification settled by Charles Darwin in the Origin of Species. Darwin tightly linked what we now call phylogeny and development. He often insisted to take into account embryonic and larval characters, most often using as examples his favourite animals, the cirripedes. Then I discuss remaining problems, and also perspectives, to address the link between phylogeny and development in the modern terms of molecular and cladistic phylogenetics and of molecular and genetic developmental biology.     

Evolutionary developmental biology its roots and characteristics

The rise of evolutionary developmental biology was not the progressive isolation and characterization of developmental genes and gene networks. Many obstacles had to be overcome: the idea that all genes were more or less involved in development; the evidence that developmental processes in insects had nothing in common with those of vertebrates.Different lines of research converged toward the creation of evolutionary developmental biology, giving this field of research its present heterogeneity. This does not prevent all those working in the field from sharing the conviction that a precise characterization of evolutionary variations is required to fully understand the evolutionary process.Some evolutionary developmental biologists directly challenge the Modern Synthesis. I propose some ways to reconcile these apparently opposed visions of evolution. The turbulence seen in evolutionary developmental biology reflects the present entry of history into biology.     

Discussion: Turing's theory of morphogenesis—Its influence on modelling biological pattern and form

The evolution of spatial pattern is a central issue in developmental biology. Turing's (Phil. Trans. R. Soc. Lond. B237, 37–72, 1952) chemical theory of morphogenesis is a seminal contribution. In this talk I give a personal and necessarily limited view of its impact on mathematical and developmental biology. I briefly describe some of the interesting mathematical aspects of Turing's reaction-diffusion mechanism and discuss some of the different models which Turing's vision inspired. The emphasis throughout is on the practical biological applications of the various theories.     

Plant fibers: Initiation,growth, model plants,and open questions

Fibers, which are used as a major raw material in the paper industry, as structural components in timber for building, and in the manufacture of wooden items, are among the most important renewable resources. Billions use wood as a major energy source, and fibers are an energy-rich component of wood. They are used for various textiles and as raw material for composites. In this review, I describe the basic characters of fibers, their structure, development, uses, and some of the current major model plants for fiber formation. I discuss open developmental questions and various aspects of further research. Most of the recent progress in the biology of fiber formation, especially in their cell-wall chemistry, emerged from studies of several model plants: Arabidopsis thaliana, Populus sp., Eucalyptus sp., flax, and hemp. I stress the critical need to combine the use of modern methods of research with classical botany. Approaching the issue of fiber formation only by molecular or only by classical methods will not only limit the progress, but may result in critical mistakes. Considering the importance of fibers to humanity, it is surprising how little we know about the biology of fiber formation and how little it is studied as compared, for instance, to the effort to study the genetics and cell biology of flower organ identity.     

Evolution and development: Past, present, and future

The paper tries to set right certain ideas about the history of evolutionary developmental biology. The main point is, that we had to enface the dominance of a comparative approach towards evolutionary developmental biology before 1900, which even later on was effective in Russia, for example, till the 1930s. The problem of the experimentalist approach set against this tradition was and is that there is no concept of gestalt that may allow to integrate the former comparative views and the modern mechanistic interpretations. We argue, that it would be wrong just to describe the comparative tradition as being outdated, as it may allow to get the framework for a dynamical concept of Gestalt that may integrate the ideas of morphogenesis and pattern formation worked out in evo-devo recently.     

What is a species? Essences and generation

Arguments against essentialism in biology rely strongly on a claim that modern biology abandoned Aristotle’s notion of a species as a class of necessary and sufficient properties. However, neither his theory of essentialism, nor his logical definition of species and genus (eidos and genos) play much of a role in biological research and taxonomy, including his own. The objections to natural kinds thinking by early twentieth century biologists wrestling with the new genetics overlooked the fact that species have typical developmental cycles and most have a large shared genetic component. These are the “what-it-is-to-be” members of that species. An intrinsic biological essentialism does not commit us to Aristotelian notions, nor even modern notions, of essence. There is a long-standing definition of “species” and its precursor notions that goes back to the Greeks, and which Darwin and pretty well all biologists since him share, that I call the Generative Conception of Species. It relies on there being a shared generative power that makes progeny resemble parents. The “what-it-is-to-be” a member of that species is that developmental type, mistakes in development notwithstanding. Moreover, such “essences” have always been understood to include deviations from the type. Finally, I shall examine some implications of the collapse of the narrative about essences in biology.     

Using evolutionary genomics,transcriptomics, and systems biology to reveal gene networks underlying fungal development

Fungal model species have contributed to many aspects of modern biology, from biochemistry and cell biology to molecular genetics. Nevertheless, only a few genes associated with morphological development in fungi have been functionally characterized in terms of their genetic or molecular interactions. Evolutionary developmental biology in fungi faces challenges from a lack of fossil records and unresolved species phylogeny, to homoplasy associated with simple morphology. Traditionally, reductive approaches use genetic screens to reveal phenotypes from a large number of mutants; the efficiency of these approaches relies on profound prior knowledge of the genetics and biology of the designated development trait—knowledge which is often not available for even well-studied fungal model species. Reductive approaches become less efficient for the study of developmental traits that are regulated quantitatively by more than one gene via networks. Recent advances in genome-wide analysis performed in representative multicellular fungal models and non-models have greatly improved upon the traditional reductive approaches in fungal evo-devo research by providing clues for focused knockout strategies. In particular, genome-wide gene expression data across developmental processes of interest in multiple species can expedite the advancement of integrative synthetic and systems biology strategies to reveal regulatory networks underlying fungal development.     

Origins of multicellular complexity: Volvox and the volvocine algae

The collection of evolutionary transformations known as the ‘major transitions’ or ‘transitions in individuality’ resulted in changes in the units of evolution and in the hierarchical structure of cellular life. Volvox and related algae have become an important model system for the major transition from unicellular to multicellular life, which touches on several fundamental questions in evolutionary biology. The Third International Volvox Conference was held at the University of Cambridge in August 2015 to discuss recent advances in the biology and evolution of this group of algae. Here, I highlight the benefits of integrating phylogenetic comparative methods and experimental evolution with detailed studies of developmental genetics in a model system with substantial genetic and genomic resources. I summarize recent research on Volvox and its relatives and comment on its implications for the genomic changes underlying major evolutionary transitions, evolution and development of complex traits, evolution of sex and sexes, evolution of cellular differentiation and the biophysics of motility. Finally, I outline challenges and suggest future directions for research into the biology and evolution of the volvocine algae.     

Moss systems biology en route: phytohormones in Physcomitrella development

The moss Physcomitrella patens has become a powerful model system in modern plant biology. Highly standardized cell culture techniques, as well as the necessary tools for computational biology, functional genomics and proteomics have been established. Large EST collections are available and the complete moss genome will be released soon. A simple body plan and the small number of different cell types in Physcomitrella facilitate the study of developmental processes. In the filamentous juvenile moss tissue, developmental decisions rely on the differentiation of single cells. Developmental steps are controlled by distinct phytohormones and integration of environmental signals. Especially the phytohormones auxin, cytokinin, and abscisic acid have distinct effects on early moss development. In this article, we review current knowledge about phytohormone influences on early moss development in an attempt to fully unravel the complex regulatory signal transduction networks underlying the developmental decisions of single plant cells in a holistic systems biology approach.     

E. E. Just's broad,yet hidden,influence on modern cell and developmental biology

This year, 2019, marks the centennial of embryologist E. E. Just's discovery of what is known as the fast block to polyspermy. Just's observation of the subtle changes that occur at the egg's surface during fertilization (and in experimental parthenogenesis) led him to postulate that the egg, and indeed every cell, possesses a property he called independent irritability, which represents the cell's ability to respond in a physiologically‐relevant way to a variety of signals or triggers. In this paper, I argue that Just's concept of independent irritability informed his contemporary Johannes Holtfreter as Holtfreter attempted to explain the phenomena of embryonic induction and competence and that Holtfreter, in turn, influenced Marc Kirschner and John Gerhart in their formulation of the theory of facilitated variation. Just's influence is especially evident in Gerhart and Kirschner's presentations of what they call weak linkage—a property of living systems that allows core processes and components to be mixed and matched in different ways to generate novel traits. Unfortunately, the connection between Holtfreter's work and Just's has remained hidden. This paper gives examples of phenomena that exhibit weak linkage, and it lays out the case that Just's concept of independent irritability, through Holtfreter, Gerhart, and Kirschner, has broadly infiltrated modern cell and developmental biology.     

Stages in the development of a model organism as a platform for mechanistic models in developmental biology: Zebrafish, 1970-2000

Model organisms became an indispensable part of experimental systems in molecular developmental and cell biology, constructed to investigate physiological and pathological processes. They are thought to play a crucial role for the elucidation of gene function, complementing the sequencing of the genomes of humans and other organisms. Accordingly, historians and philosophers paid considerable attention to various issues concerning this aspect of experimental biology. With respect to the representational features of model organisms, that is, their status as models, the main focus was on generalization of phenomena investigated in such experimental systems. Model organisms have been said to be models for other organisms or a higher taxon. This, however, presupposes a representation of the phenomenon in question. I will argue that prior to generalization, model organisms allow researchers to built generative material models of phenomena - structures, processes or the mechanisms that explain them - through their integration in experimental set-ups that carve out the phenomena from the whole organism and thus represent them. I will use the history of zebrafish biology to show how model organism systems, from around 1970 on, were developed to construct material models of molecular mechanisms explaining developmental or physiological processes.     

Post‐translational modifications as key regulators of apicomplexan biology: insights from proteome‐wide studies

Parasites of the Apicomplexa phylum, such as Plasmodium spp. and Toxoplasma gondii, undergo complex life cycles involving multiple stages with distinct biology and morphologies. Post‐translational modifications (PTMs), such as phosphorylation, acetylation and glycosylation, regulate numerous cellular processes, playing a role in every aspect of cell biology. PTMs can occur on proteins at any time in their lifespan and through alterations of target protein activity, localization, protein–protein interactions, among other functions, dramatically increase proteome diversity and complexity. In addition, PTMs can be induced or removed on changes in cellular environment and state. Thus, PTMs are likely to be key regulators of developmental transitions, biology and pathogenesis of apicomplexan parasites. In this review we examine the roles of PTMs in both parasite‐specific and conserved eukaryotic processes, and the potential crosstalk between PTMs, that together regulate the intricate lives of these protozoa.     

Generating phenotypic variation: prospects from "evo-devo" research on Bicyclus anynana wing patterns

Understanding the generation of phenotypic variation is an important challenge for modern evolutionary biology, and butterfly wing patterns are an exciting system that can shed some light on this issue. Here, we report on recent advances in the genetics of Bicyclus anynana butterflies. This system provides the potential for a fully integrated study of the evolutionary and developmental processes underlying diversity in morphology.     

Cytological examination of reduction bodies of Corvomeyenia carolinensis harrison (Porifera: Spongillidae)

Freshwater sponges, Corvomeyenia carolinensis Harrison, were placed into tap water to induce degenerative reduction body formation. Reduction bodies were examined using light and electron microscopy in order to define their histochemical and ultrastructural characteristics. The reduction body of freshwater sponges is an extremely simple developmental system consisting primarily of an archeocyte reserve delimited by a simple squamous pinacoderm. The freshwater sponge reduction body displays many similarities to overwintering phases of marine sponges. The system presents an unusually straightforward vehicle for investigations of degeneration and regeneration as processes in developmental biology and may represent a reasonable vehicle in which to examine the process of the genesis of lysosomes.     

Homology and heterochrony: the evolutionary embryologist Gavin Rylands de Beer (1899-1972)

The evolutionary embryologist Gavin Rylands de Beer can be viewed as one of the forerunners of modern evolutionary developmental biology in that he posed crucial questions and proposed relevant answers about the causal relationship between ontogeny and phylogeny. In his developmental approach to the phylogenetic phenomenon of homology, he emphasized that homology of morphological structures is to be identified neither with the sameness of the underlying developmental processes nor with the homology of the genes that are involved in the development of the structures. De Beer's work on developmental evolution focused on the notion of heterochrony, arguing that paedomorphosis increases morphological evolvability and is thereby an important mode of evolution that accounts for the origin of many taxa, including higher taxa.     

Four decades of teaching developmental biology in Germany

I have taught developmental biology in Essen for 30 years. Since my department is named Zoophysiologie (Zoophysiology), besides Developmental Biology, I also have to teach General Animal Physiology. This explains why the time for teaching developmental biology is restricted to a lecture course, a laboratory course and several seminar courses. However, I also try to demonstrate in the lecture courses on General Physiology the close relationship between developmental biology, physiology, morphology, anatomy, teratology, carcinogenesis, evolution and ecology (importance of environmental factors on embryogenesis). Students are informed that developmental biology is a core discipline of biology. In the last decade, knowledge about molecular mechanisms in different organisms has exponentially increased. The students are trained to understand the close relationship between conserved gene structure, gene function and signaling pathways, in addition to or as an extension of, classical concepts. Public reports about the human genome project and stem cell research (especially therapeutic and reproductive cloning) have shown that developmental biology, both in traditional view and at the molecular level, is essential for the understanding of these complex topics and for serious and non-emotional debate.