Physical explanations and biological explanations,empirical laws and a priori laws

Philosophers intent upon characterizing the difference between physics and biology often seize upon the purported fact that physical explanations conform more closely to the covering law model than biological explanations. Central to this purported difference is the role of laws of nature in the explanations of these two sciences. However, I argue that, although certain important differences between physics and biology can be highlighted by differences between physical and biological explanations, these differences are not differences in the degree to which those explanations conform to the covering law model, which fits biology about as well as it does physics.     

Toward a conceptual framework for biology

Science progresses faster when researchers operate within an explicit framework of concepts and theories, but currently biology has no explicit, overarching conceptual framework and few general theories. The single general theory currently recognized is that of evolution, which was put forth by Charles Darwin 150 years ago. Recently, Scheiner and Willig (2008) explicated a similarly general theory of ecology. In this paper, using the theory of evolution as an exemplar, I discuss the nature of theory in biology and put forth an overarching theory, as well as new general theories for cells, organisms, and genetics. Along with the theories of evolution and ecology, these constitute a general conceptual framework for the biological sciences. This framework reveals linkages among the various parts of biology, makes explicit the assumptions behind more narrow theories and models, and provides new insights into the structures of biological theories. This framework can also be used as a teaching tool, moving the teaching of biology beyond the transference of a vast compendium of facts. My hope is that this essay will lead to a vigorous discussion and debate across all of biology about the nature and structure of its theories.     

Inferential explanations in biology

Among philosophers of science, there is now a widespread agreement that the DN model of explanation is poorly equipped to account for explanations in biology. Rather than identifying laws, so the consensus goes, researchers explain biological capacities by constructing a model of the underlying mechanism.We think that the dichotomy between DN explanations and mechanistic explanations is misleading. In this article, we argue that there are cases in which biological capacities are explained without constructing a model of the underlying mechanism. Although these explanations do not conform to Hempel’s DN model (they do not deduce the explanandum from laws of nature), they do invoke more or less stable generalisations. Because they invoke generalisations and have the form of an argument, we call them inferential explanations. We support this claim by considering two examples of explanations of biological capacities: pigeon navigation and photoperiodism. Next, we will argue that these non-mechanistic explanations are crucial to biology in three ways: (i) sometimes, they are the only thing we have (there is no alternative available), (ii) they are heuristically useful, and (iii) they provide genuine understanding and so are interesting in their own right.In the last sections we discuss the relation between types of explanations and types of experiments and situate our views within some relevant debates on explanatory power and explanatory virtues.     

HISTORICAL VERSUS SELECTIONIST EXPLANATIONS IN EVOLUTIONARY BIOLOGY

Abstract— Evolutionary changes require historical explanations, yet these are limited by the evolutionary processes we entertain and investigate. Using phylogenetic analysis, adaptation and natural selection can be tested as historical claims, but this is appropriate only in those special cases where change follows the scheme of one character-one function, singled out in new environmental circumstances. Systematic treatment of the evolutionary origin of characters (in particular, origin through ecological and developmental flexibility) lies outside the scope of selectionist explanations. Structural hypotheses about regularities in the directions of change, also analyzed phylogenetically, expand the scope of historical explanation to include the origin of characters, yet retain the view of organisms as passive and constrained objects of evolutionary change. Historical biology needs to encompass both the active responses of organisms and the construction by organisms of their own environments. For this to be realized will require changes in the concepts and practices of evolutionary biology, including a re-examination of the Lamarckian theme that the active responses of organisms have evolutionary significance—the rarity of individual-to-individual transmission of "acquired" characters does not disprove the possibility of their frequency increasing in a population.     

Dual Causality and the Autonomy of Biology

Ernst Mayr’s concept of dual causality in biology with the two forms of causes (proximate and ultimate) continues to provide an essential foundation for the philosophy of biology. They are equivalent to functional (=proximate) and evolutionary (=ultimate) causes with both required for full biological explanations. The natural sciences can be classified into nomological, historical nomological and historical dual causality, the last including only biology. Because evolutionary causality is unique to biology and must be included for all complete biological explanations, biology is autonomous from the physical sciences.     

Explanation in morphology

Summary In biology, and particularly in morphology, various types of explanation are found,e.g. causal, teleological, historical, etc.In this article an attempt has been made to analyse the relations between the various explanations to strive for an encompassing explanatory theory.The general structure of the explanatory theories appeared to be very similar, but the terms defining the phenomena and the types of the relations within the theories differ. To obtain a unifying theory it is necessary to develop methods to connect or transform the different elements.An important aspect is that the definition of selected phenomena (terms) depends on the explanation pursued.Lecture presented for the Netherlands Society for Theoretical Biology, December, 1971.     

Organic Codes: A Unifying Concept for Life

Although the knowledge about biological systems has advanced exponentially in recent decades, it is surprising to realize that the very definition of Life keeps presenting theoretical challenges. Even if several lines of reasoning seek to identify the essence of life phenomenon, most of these thoughts contain fundamental problem in their basic conceptual structure. Most concepts fail to identify either necessary or sufficient features to define life. Here, we analyzed the main conceptual frameworks regarding theoretical aspects that have been supporting the most accepted concepts of life, such as (i) the physical, (ii) the cellular and (iii) the molecular approaches. Based on an ontological analysis, we propose that Life should not be positioned under the ontological category of Matter. Yet, life should be better understood under the top-level ontology of “Process”. Exercising an epistemological approach, we propose that the essential characteristic that pervades each and every living being is the presence of organic codes. Therefore, we explore theories in biosemiotics and code biology in order to propose a clear concept of life as a macrocode composed by multiple inter-related coding layers. This way, as life is a sort of metaphysical process of encoding, the living beings became the molecular materialization of that process. From the proposed concept, we show that the evolutionary process is a fundamental characteristic for life’s maintenance but it is not necessary to define life, as many organisms are clearly alive but they do not participate in the evolutionary process (such as infertile hybrids). The current proposition opens a fertile field of debate in astrobiology, epistemology, biosemiotics, code biology and robotics.

     

Physical models of biological information and adaptation

The bio-informational equivalence asserts that biological processes reduce to processes of information transfer. In this paper, that equivalence is treated as a metaphor with deeply anthropomorphic content of a sort that resists constitutive-analytical definition, including formulation within mathematical theories of information. It is argued that continuance of the metaphor, as a quasi-theoretical perspective in biology, must entail a methodological dislocation between biological and physical science. It is proposed that a general class of functions, drawn from classical physics, can serve to eliminate the anthropomorphism. Further considerations indicate that the concept of biological adaptation is central to the general applicability of the informational idea in biology; a non-anthropomorphic treatment of adaptive phenomena is suggested in terms of variational principles.     

Emergent insights from the synthesis of conceptual frameworks for biological invasions

A general understanding of biological invasions will provide insights into fundamental ecological and evolutionary problems and contribute to more efficient and effective prediction, prevention and control of invasions. We review recent papers that have proposed conceptual frameworks for invasion biology. These papers offer important advances and signal a maturation of the field, but a broad synthesis is still lacking. Conceptual frameworks for invasion do not require invocation of unique concepts, but rather should reflect the unifying principles of ecology and evolutionary biology. A conceptual framework should incorporate multicausality, include interactions between causal factors and account for lags between various stages. We emphasize the centrality of demography in invasions, and distinguish between explaining three of the most important characteristics by which we recognize invasions: rapid local population increase, monocultures or community dominance, and range expansion. As a contribution towards developing a conceptual synthesis of invasions based on these criteria, we outline a framework that explicitly incorporates consideration of the fundamental ecological and evolutionary processes involved. The development of a more inclusive and mechanistic conceptual framework for invasion should facilitate quantitative and testable evaluation of causal factors, and can potentially lead to a better understanding of the biology of invasions.     

Living matter—nexus of physics and biology in the 21st century

Cells are made up of complex assemblies of cytoskeletal proteins that facilitate force transmission from the molecular to cellular scale to regulate cell shape and force generation. The “living matter” formed by the cytoskeleton facilitates versatile and robust behaviors of cells, including their migration, adhesion, division, and morphology, that ultimately determine tissue architecture and mechanics. Elucidating the underlying physical principles of such living matter provides great opportunities in both biology and physics. For physicists, the cytoskeleton provides an exceptional toolbox to study materials far from equilibrium. For biologists, these studies will provide new understanding of how molecular-scale processes determine cell morphological changes.The distinction between being “alive” or “not alive” has been a long-standing question for those interested in our natural world. In many ancient cultures, the difference between living organisms and inorganic matter was thought to be due to innate differences arising from a “vital force,” such that biology operated with different fundamental properties than the physical world. The ability to disprove such theories came about over the course of the 17th to the 19th centuries, as scientists developed theories of atoms and were able to synthesize organic matter from inorganic constituents. Over the past 100 years, developments in molecular biology and biochemistry have provided a wealth of information on the structure and function of biological molecules, much of which was acquired in collaborations between physical and biological scientists. Application of X-ray–scattering techniques first developed to study metals enabled discovery of the structure of complicated biological molecules ranging from DNA to ion channels. Use of laser trapping techniques first developed to trap and cool atoms enabled precise force spectroscopy measurements of single molecular motors. We now know that biological molecules, while more complicated than their inorganic counterparts, must obey the rules of physics and chemistry.This wealth of molecular-scale information does not directly inform the behaviors of living cells. The organelles within cells are made up of complex and dynamic assemblies of proteins, lipids, and nucleic acids, all immersed within an aqueous environment. These assemblies are somehow able to build materials that can robustly facilitate the plethora of morphological and physical behaviors of cells at the subcellular (intracellular transport), cellular (division, adhesion, migration), and multicellular (tissue morphogenesis, wound healing) length scales. The dynamic cytoskeleton transmits information and forces from the molecular to the cellular length scales. But what is it about the behaviors of biological molecules that endow cells with the ability to respirate, move, and replicate themselves robustly—all qualities we consider essential to “life”? For these questions, understanding of the physics and chemistry of systems of biological molecules is needed. Interactions that occur within ensembles of molecules lead to emergent properties and behaviors that cannot be predicted at the single-molecule level. These emergent chemical and physical properties of living matter are likely fundamentally different from inorganic or “dead” materials. Discovering the underlying principles of living matter provides fantastic opportunities to learn new physics and biology.The fields of condensed matter physics and materials science study the physical properties that emerge when objects (e.g., atoms, molecules, grains of sand, or soap bubbles) are placed in sufficiently close proximity, such that interactions between them cannot be ignored. Interatomic or intermolecular interactions give rise to emergent properties that are not seen in isolated species. Familiar examples involve electron transport across a material or a material''s response to externally applied magnetic fields or mechanical forces. These emergent properties, such as conductivity, elasticity, and viscosity, enable us to predict the behavior of a collection of objects in these condensed phases. In this paper, I will focus on my perspective of how approaches to understanding the mechanical properties of physical materials can inform understanding of the mechanical properties of living matter found within cells.In a crystal of metal, precisely organized atoms are located nanometers apart, and the energies of their interactions are on the scale of an electron volt (40-fold larger than thermal energy or twice the energy released on the hydrolysis of a single ATP molecule). These give rise to an energy density, or elastic modulus, on the order of gigapascals, which underlies the rigidity of metals. For small deformations, the restoring force between atoms means that this metal behaves like an elastic spring: after a force is applied, the metal returns to its original shape. Understanding force transmission through crystalline metals was facilitated by the development of elasticity theory in the 16th and 17th centuries. Fluids, such as water, lack crystalline order, but predictive understanding of fluid flows and forces was captured through development of theories of fluid dynamics. Now think of another material, Silly Putty, which behaves elastically at short timescales (it bounces like a rubber ball) but then oozes and flows at long timescales, acting like a viscous fluid. Silly Putty is made of long polymers that are trapped by one another at short timescales, but thermal energy is sufficient to allow them to diffuse and translocate at long timescales. Silly Putty is also a “soft material,” in that the polymer''s interaction energies are at the thermal energy level, and its length scale is at the micrometer level. Materials like Silly Putty were thought to be too complicated for analytical theory. It was only in the middle of the 20th century that the theoretical framework to understand these “messy” and “disorganized” polymer-based materials was developed.The most powerful theories for understanding these vastly different forms of physical matter were developed in the absence of even the simplest of computers. The theories relied on developing physical properties or parameters to describe the material with a “mean field,” a type of coarse-graining that identifies the essential properties of individual constituents and interactions but ignores many other details. These mean fields give us new intuitions concerning the origin of material properties and give rise to definitions of physical parameters, such as elasticity and viscosity. However, these theories also require materials that do not jostle around a lot and remain close to equilibrium. In fact, understanding materials “far from equilibrium” has been identified as a major challenge in physics for the next century (National Research Council, 2007) .Materials formed by dynamic protein assemblies in the cytoskeleton are disorganized, heterogeneous, and driven far from equilibrium. Motor proteins generate local stresses, and their activity is spatially modulated. The polymerization and depolymerization of cytoskeletal polymers is controlled by a myriad of regulatory proteins. All these dynamic molecular processes endow the cytoskeletal assemblies with unique behaviors that enable them to support complex physiological tasks. It is likely these dynamics also provide underlying robustness of the cells in response to fluctuating and changing environments. These properties make living cells exquisite materials that cannot be captured by existing frameworks of physical matter. I suspect that we have not yet identified the important parameters needed to characterize their properties. The rich dynamics created by active biological matter present a formidable challenge in the area of materials science.How do we hope to understand the properties of these complex cytoskeletal assemblies and materials? It may seem as though understanding cytoskeletal machinery is an insurmountable feat, the approaches that have been successful for physical materials will not work, and we must rely on complex simulations that require modeling of all individual components. This may be true. However, I think that this is a pessimistic view. Just consider how complicated physical materials would be if we did not have the appropriate parameters to describe the macroscopic responses and had instead became obsessed about knowing the details of all the interactions between underlying atoms and molecules? In the same vein, I believe that predictive insights into biological matter will emerge through development of new physical theories that use mean-field approaches to understanding materials that contain active components and are driven far from equilibrium. The burgeoning field of active-matter physics is currently considering these questions (Ramaswamy, 2010) . However, these theoretical approaches require physical measurements of cells and cellular proteins that may not be clearly linked to a physiological process or have a clear biological context. Materials built from cytoskeletal proteins in vitro should also provide an excellent source of experimental measurements, but closer collaboration with theorists working in this field and collaboration between biochemists and experimental physical scientists is needed to develop control over such materials. Developing predictive physical theories of the cytoskeleton will elucidate principles of why “the whole is more than the sum of its parts” that will provide greater control and design over living matter, in the same way that engineering has provided great advances in applications of materials from the physical world.What do biologists gain from theories of living matter? These theories will provide a crucial link between molecular and cellular length scale behaviors and will provide insight into the mechanisms of why specific molecular perturbations alter cell behavior. Moreover, they should provide us with general design principles of living matter. What are the basic aspects of a machine needed to separate chromosomes, establish polarity, or generate contractile forces that is utilized across different cell types? Can knowing these aspects provide insight into the evolution of cellular machines and the robustness of cell behavior? Thus, study of cellular materials both provides new opportunities for physicists and will provide crucial predictive understanding of cell physiology.Open in a separate windowMargaret L. Gardel     

Emergence,therefore antireductionism? A critique of emergent antireductionism

Emergent antireductionism in biological sciences states that even though all living cells and organisms are composed of molecules, molecular wholes are characterized by emergent properties that can only be understood from the perspective of cellular and organismal levels of composition. Thus, an emergence claim (molecular wholes are characterized by emergent properties) is thought to support a form of antireductionism (properties of higher-level molecular wholes can only be understood by taking into account concepts, theories and explanations dealing with higher-level entities). I argue that this argument is flawed: even if molecular wholes are characterized by emergent properties and even if many successful explanations in biology are not molecular, there is no entailment between the two claims.     

The complexity of living bodies and the structure of biological theories

It has been suggested that biological theories differ from physical theories because the subject matter of biology differs from the subject matter of physics especially in the fact that living bodies are more complex than nonliving bodies. It is shown that the interactional complexity of living bodies can only be expressed by invoking biological theories. The claim that living bodies are complex is, therefore, ultimately a claim about the nature of scientific theories rather than a claim about the nature of the subject matter of biology resting upon a presystematic judgement.     

The female orgasm and the homology concept in evolutionary biology

The definition of homology and its application to reproductive structures, external genitalia, and the physiology of sexual pleasure has a tortuous history. While nowadays there is a consensus on the developmental homology of genital and reproductive systems, there is no agreement on the physiological translation, or the evolutionary origination and roles, of these structural correspondences and their divergent histories. This paper analyzes the impact of evolutionary perspectives on the homology concept as applied to the female orgasm, and their consequences for the biological and social understanding of female sexuality and reproduction. After a survey of the history of pre-evolutionary biomedical views on sexual difference and sexual pleasure, we examine how the concept of sexual homology was shaped in the new phylogenetic framework of the late 19th century. We then analyse the debates on the anatomical locus of female pleasure at the crossroads of theories of sexual evolution and new scientific discourses in psychoanalysis and sex studies. Moving back to evolutionary biology, we explore the consequences of neglecting homology in adaptive explanations of the female orgasm. The last two sections investigate the role played by different articulations of the homology concept in evolutionary developmental explanations of the origin and evolution of the female orgasm. These include the role of sexual, developmental homology in the byproduct hypothesis, and a more recent hypothesis where a phylogenetic, physiological concept of homology is used to account for the origination of the female orgasm. We conclude with a brief discussion on the social implications for the understanding of female pleasure derived from these different homology frameworks.     

Computational approaches for the prediction of protein function in the mitochondrion

Understanding a complex biological system, such as the mitochondrion, requires the identification of the complete repertoire of proteins targeted to the organelle, the characterization of these, and finally, the elucidation of the functional and physical interactions that occur within the mitochondrion. In the last decade, significant developments have contributed to increase our understanding of the mitochondrion, and among these, computational research has played a significant role. Not only general bioinformatics tools have been applied in the context of the mitochondrion, but also some computational techniques have been specifically developed to address problems that arose from within the mitochondrial research field. In this review the contribution of bioinformatics to mitochondrial biology is addressed through a survey of current computational methods that can be applied to predict which proteins will be localized to the mitochondrion and to unravel their functional interactions. genomic context; proteome     

Non-conflict theories for the evolution of genomic imprinting

Theories focused on kinship and the genetic conflict it induces are widely considered to be the primary explanations for the evolution of genomic imprinting. However, there have appeared many competing ideas that do not involve kinship/conflict. These ideas are often overlooked because kinship/conflict is entrenched in the literature, especially outside evolutionary biology. Here we provide a critical overview of these non-conflict theories, providing an accessible perspective into this literature. We suggest that some of these alternative hypotheses may, in fact, provide tenable explanations of the evolution of imprinting for at least some loci.     

Are there adequate data to assess how well theories of rarity apply to marine invertebrates?

Understanding the biology of rare species is a very important part of conservation biology. Most of our current understanding of rarity has, however, come from studies of terrestrial plants, birds, mammals and some insects. Freshwater and marine habitats are underrepresented in published studies of rare species or conservation biology. We therefore have little knowledge about how well our understanding of what makes particular species rare and how rare species persist applies to marine invertebrates which form a major component of coastal biodiversity. In this review, I examine some theories about rarity with respect to intertidal and shallow subtidal invertebrates to identify whether there are adequate data to apply these theories to marine invertebrates and how well such theories apply. The general conclusions are that the lack of quantitative data on abundances, ranges, habitat-requirements, dispersal and connectedness among populations for marine invertebrates means that their status as rare species cannot really be assessed appropriately. It is also unlikely that, without extensive sampling programmes and considerable expense, adequate data could be obtained for these small, cryptic animals, which typically have very patchy, variable and unpredictable patterns of distribution and abundance. Intertidal and subtidal assemblages are diverse, including species with many different life-histories from many phyla, occupying the same suite of habitats. It is therefore suggested that future research on rare organisms in marine habitats should build upon the long and successful history of experimental marine studies to test specific hypotheses about processes influencing rarity in the field. Such studies would not only add a new dimension to our current understanding of rarity, but would also provide badly-needed data on the status of rare marine invertebrates. abundances, invertebrates, marine, range, rarity     

A role for the mismatch repair system during incipient speciation in Saccharomyces

The cause of reproductive isolation between biological species is a major issue in the field of biology. Most explanations of hybrid sterility require either genetic incompatibilities between nascent species or gross physical imbalances between their chromosomes, such as rearrangements or ploidy changes. An alternative possibility is that genomes become incompatible at a molecular level, dependent on interactions between primary DNA sequences. The mismatch repair system has previously been shown to contribute to sterility in a hybrid between established yeast species by preventing successful meiotic crossing-over leading to aneuploidy. This system could also promote or reinforce the formation of new species in a similar manner, by making diverging genomes incompatible in meiosis. To test this possibility we crossed yeast strains of the same species but from diverse historical or geographic sources. We show that these crosses are partially sterile and present evidence that the mismatch repair system is largely responsible for this sterility.     

The Neutral—Niche Debate: A Philosophical Perspective

Ecological communities around the world are under threat while a consensus theory of community structure remains elusive. In the last decade ecologists have struggled with two seemingly opposing theories: niche-based theory that explains diversity with species’ differences and the neutral theory of biodiversity that claims that much of the diversity we observe can be explained without explicitly invoking species’ differences. Although ecologists are increasingly attempting to reconcile these two theories, there is still much resistance against the neutral theory of biodiversity. Here we argue that the dispute between the two theories is a classic example of the dichotomy between philosophical perspectives, realism and instrumentalism. Realism is associated with specific, small-scale and detailed explanations, whereas instrumentalism is linked to general, large-scale, but less precise accounts. Recognizing this will help ecologists get both niche-based and neutral theories in perspective as useful tools for understanding biodiversity patterns.     

Darwin's legacy: why biology is not physics, or why evolution has not become a common sense

Cosmology and evolution together have enabled us to look deep into the past and comprehend evolution-from the big bang to the cosmos, from molecules to humans. Here, I compare the nature of theories in biology and physics and ask why physical theories get accepted by the public without necessarily comprehending them but biological theories do not. Darwin's theory of natural selection, utterly simple in its premises but profound in its consequences, is not accepted widely. Organized religions, and creationists in particularly, have been the major critic of evolution, but not all opposition to evolution comes from organized religions. A great many people, between evolutionary biologists on one hand and creationists on the other, many academics included, who may not be logically opposed to evolution nevertheless do not accept it. This is because the process of and the evidence for evolution are invisible to a nonspecialist, or the theory may look too simple to explain complex traits to some, or because people compare evolution against God and find evolutionary explanations threatening to their beliefs. Considering how evolution affects our lives, including health and the environment to give just two examples, a basic course in evolution should become a required component of all our college and university educational systems.