Tiago Rama

This article characterizes Oyama’s concept of ontogenesis of information formally. I apply the mathematical notion of synergistic information to the framework proposed by Griffiths et al. (2015) for developmental information and specificity. This allows us to examine the specificity revealed by the interaction of variables as a result of interventions on interactions. I define Developmental Synergistic Information as the specificity of interacting variables obtained by measuring how much mutual information interventions on interactions carry about the effect variable. To formalise this concept, I use partial information decomposition, one of the most robust frameworks for analysing synergistic information. Some examples of developmental synergistic information are presented. Finally, I consider the philosophical implications of developmental synergistic information, arguing that it supports important tenets of organism-centered biology: (i) synergistic information has ontogeny—order is generated in epigenesis; (ii) such information is non-transmissible through channels of inheritance—the specificity of outcomes must be reconstructed anew in each generation; (iii) the developmental organism (or any synergistic system under consideration) is itself a cause of development—causation resides in the coaction of developmental variables; and (iv) the developmental context of information must be taken into account—that developmental causation is always embedded in and constrained by a developmental matrix of other specifiers.

Abstract: Inheritance theory is broadening by emphasizing two kinds of diversity, both justified by recent empirical advances: the material substrates of inheritance beyond genes, and the directions of inheritance relationships beyond vertical, parent-to-offspring transmission. There are different inheritance materials—genetic, epigenetic, parental, cultural—and inheritance directions—vertical, oblique, horizontal. This shift is accompanied by theoretical reconceptualizations associated with developmental and organismal views of inheritance, which understand inheritance as the transgenerational reconstruction of traits that explain trait persistence across a lineage. Within this framework, this article draws attention to inverted inheritance: inheritance relationships that run from a younger generation to an older one. Although this idea may initially seem at odds with how inheritance is generally understood and studied, this article argues that inverted inheritance is compatible with—and indeed illuminated by—the current expansion of inheritance theory. While it highlights relationships that invite empirical investigation, it also sheds light on our conceptualization of what inheritance is.

Wakefield’s “hybrid” theory of mental illness defines the (dys)function of mental traits in terms of the evolutionary account of biological functions known as the Selected-Effect Theory. This evolutionary framework, however, has been subject to serious challenges in recent decades. Accordingly, the article sets out—programmatically and through a detailed literature review—an appraisal of hybrid accounts of mental health in light of debates around the philosophy of biological functions. Our analysis proceeds in two stages. First, it extends and systematizes three central critiques to the Selected-Effect Theory to Wakefield’s account: (i) the explanatory inadequacy of historical and population-based accounts of function for defining mental illness, (ii) the possibility that the origins and persistence of mental traits may resist functional explanation, and (iii) the claim that functional explanations of the mind’s evolution extend beyond selection-based accounts. Second, we explore alternative frameworks through which biological theories of function can shape the definition of mental illness—particularly the causal-role theory and the organizational account.

The career of the developmental psychobiologist Gilbert Gottlieb revolved around the construction of a probabilistic view of epigenetics. In drastic contrast to instinct and nativist theories which portray development as a determinate sequence of changes controlled by a genetic master plan, Gottlieb’s experiments and theory picture (behavioral) development as a highly multicausal complex process, with bidirectional interactions across different levels of organization and profound influences from ecological causes. Notwithstanding, the rejection of deterministic epigenetics was accompanied by the neglect of the concept of information in developmental biology, which various developmental systems theorists advocated. Information, particularly its interpretation in the gene-centered framework, was seen as an obstacle to probabilistic epigenetics. In recent years, however, Paul Griffiths and colleagues have developed an informational view of development that encompasses the theoretical and experimental background of developmental systems theory. There is thus an opportunity to build a bridge between probabilistic epigenetics and information theory, and this paper explores this avenue. Here I explore probabilistic epigenetics by adopting an informational view that allows us to represent and model specific aspects related to the probabilistic nature of development: dynamic epigenetic landscapes, probabilistic phenotypic repertoires, and the multicausality, non-linearity, and bidirectionality of developmental processes. This paper intends to enrich probabilistic epigenetics with informational tools while extending the applicability and utility of Griffiths et al.’s theory to previously unexplored developmental phenomena.

An agential and organismic view of development has brought back certain concepts in biology, such as teleology and normativity. What are the goals and norms that a developmental system pursues? Where do phenotypes in ontogenesis come from? The common answer is that genes contain the recipe to build organisms. However, the agential and organismic view of development seeks to offer a different answer beyond any reductionist explanations. For this reason, teleological development has become a central explanans in developmental and evolutionary biology. However, the motivation for this article is that teleological development as an explanandum has not been treated with the same intensity and rigor despite its ubiquity as an explanans. In particular, a common and now widely adopted strategy for explaining teleological development is to appeal to a closely related, yet fundamentally distinct, conception of teleology: the theory of autonomous systems, a theory designed to deal with physiological-level processes, not developmental ones. This article explores the (generally unanalyzed) differences between teleological development and teleological physiology by arguing that autonomous systems theory falls short of explaining the specificity of teleological development. The main conclusion is that the agential and organismic views in biology ought to be broadened to include different goals beyond self-maintenance; that to be alive is not only about the maintenance of an organization. The distinctiveness of teleological development entails specific philosophical puzzles—what I call Bernard and Baer’s desiderata—that are neither present in nor solved by autonomous systems theory.

Teleosemantics explains meaning by appealing to the biological norms that make error possible, but most work in the field still anchors those norms in evolutionary “selected‐effect” functions. We develop an organismic alternative grounded in the self-maintenance of autonomous systems. Building on sensorimotor theory and enactivism, we reconceptualise goals as second-order constraints—attractors in a dynamic sensorimotor field—and show how they are nested into a heterarchy of means–end relations connecting global self‑maintenance of the sensorimotor organization and identity of system with the most basic sensorimotor coordinations. Drawing on this framework, we identify the minimal necessary requirements for genuine teleological behaviour: 1. Initiation that individuates an action in relation to a goal, 2. Modulatory execution that adaptively compensates perturbations or deploys alternative strategies in relation to the goal, and 3. Meaningful termination through successful acknowledgment and transition, goal-sensitive persistence, or coherent abandonment. We locate normativity and error in the dynamic presuppositions among sensorimotor schemes and goals: failure manifests as incoordination within, or across, goal branches. We finally illustrate our account with a detailed analysis of the well known A-not-B error in psychology, showing how infants meet our teleological criteria while revealing incoordination as tension breakdowns inside a heterarchy of goals and sensorimotor schemes. Teleological behaviour, therefore, is the management of tensions and dynamic presuppositions within the goal heterarchy, not a chain of mechanical causes, thus delivering a naturalised notion of purposiveness that opens up an alternative route to teleosemantics beyond evolutionary selection.

Musical metaphors represent a means by which biologists illustrate a particularly complex phenomenon: how organisms regulate their life processes according to their living conditions. This explanatory role can be traced back to the early embryologists and organicism. Also, nowadays, musical metaphors are used by those who resist viewing organic processes as reading (performing) a musical score written in a genetic script. In contrast to this reductionist view, organic processes are presented under a Jazz band view in which organic activities are the result of live improvisations based on various sources of information available. This article explores the philosophical and historical foundations of the Jazz band view in the context of plant phytoacoustics, the field concerned with understanding the regulatory role of sound in plants. The incredible ability of plants to perceive, produce, and communicate sounds is central to various regulatory processes in development, maintenance, and reproduction. Musical metaphors and the place of sound in the life cycle of organisms converge in the same explanatory target, regulation, regardless of whether it is explained metaphorically or literally. This allows us to pose some programmatic questions for phytoacoustics that are motivated by current research topics in the philosophy of biology.

Abstract: Ernst Mayr famously distinguished between proximate and ultimate causal explanations. His view was decisive in the fact that development was not included in the theory of evolution. Nevertheless, the explanatory role of developmental processes in evolution is a central theme of current theoretical biology, which has led to several revisions of Mayr’s distinction. One of the reasons for this is that the interactions between organisms and the environment influence evolution. This is a cornerstone of the Extended Evolutionary Synthesis. How are these reciprocal interactions integrated into evolutionary theory? I will argue that this question can sometimes be answered by adopting an interactivist view based on a reinterpretation of reciprocal causation: developmental processes (such as niche construction or developmental plasticity) interact reciprocally with natural selection to produce evolved traits. In the following, I will critically analyze Mayr’s critics to determine whether the integration of development and evolution based on the reinterpretation of reciprocal causation is an appropriate alternative. I will argue that this is not the case, since this interactivist framework raises several explanatory problems. Instead, I should reconsider Mayr’s distinction by adopting an alternative view of evolutionary causation, known as the statisticalist view of natural selection, which posits that the only level of causation is the individual level and that ultimate explanations are statistical in nature. I argue that this framework avoids the explanatory problems identified earlier. To explore the relationship between organisms–environment reciprocal causation and ultimate explanations, I introduce the concept of ‘statistical reciprocity’ to measure the statistical effects of reciprocal causation in population changes. I situate this proposal within a general framework I call ‘population ontogenetics’, which is seen as an attempt to unify development and evolution beyond interactivist positions.

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, a similar situation can be found in contemporary evolutionary theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis refrain from any substantial change by reinterpreting and explaining individual-level phenomena from a population perspective. Finally, I will approach these historical and contemporary debates by arguing for the statistical reading of natural selection, which holds that explanations by natural selection are statistical. My main conclusion is that the historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. Adopting the statistical point of view has three advantages for the issues discussed in this paper. First, understanding historical conceptual change as part of an explanatory shift fits with the emergence of population biology as a discipline that employs statistical methods. Second, concerning current debates in evolutionary biology, the statisticalist reading can validate the goal of both sides of the dispute. It ascribes an invaluable role to the population statistical explanation of the MS and also commends the study of developmental and organismal causes of adaptive evolution. Finally, the division of explanatory roles in evolutionary biology, embarrassed by statisticalism, can be related to the different interpretations that important biological concepts have undergone throughout history and contemporary biology, i.e., that the division of explanatory roles allows for a division of conceptual interpretations.

This paper argues that a central explanatory role for the concept of _Umwelt_ in theoretical biology is to be found in developmental biology, in particular in the effort to understand development as a goal-directed and adaptive process that is controlled by the organism itself. I will reach this conclusion in two (interrelated) ways. The first is purely theoretical and relates to the current scenario in the philosophy of biology. Challenging neo-Darwinism requires a new understanding of the various components involved in natural selection processes. An important prerequisite is to understand developmental change in a teleological way. Here, the concept of _Umwelt_ plays a crucial role: if organisms are responsible for generating adaptive variation in specific environments, we need a theory that explains the context-dependent nature of adaptively oriented processes. The _Umwelt_ is thus a central element in determining the goal that an adaptive process pursues. The second path in my analysis also has a historical dimension. I will present Karl Ernst von Baer’s reflections on teleological development and his influence on Jacob von Uexküll’s thinking. I will present various ideas developed by Baer, such as the distinction between _Ziel_ and _Zweck_ and the use of musical metaphors, which can help to understand development teleologically and give Uexküll’s theory a central place in this framework.

A common explanatory error in science concerns the conflation of the epistemological roles between two domains. Here we will address a specific case: when explanations of development replace evolutionary explanations or vice versa. Ernst Mayr famously distinguished between proximate and ultimate causal explanations in biology. His view was central to the Modern Synthesis’ exclusion of development from evolutionary theory. Nonetheless, the explanatory role of developmental processes in evolution is a central theme in current theoretical biology which has prompted several revisions of Mayr’s distinction. Here we will review these reviewers to determine whether the integration of development and evolution is based on an appropriate reinterpretation of Mayr’s distinction. In many cases, revisionists suggest an interactionist alternative, in which proximate and ultimate causes interact to produce evolved traits. The most frequent interactionist account relies on the idea of reciprocal causation. We will argue that this perspective is still problematic and that the boundaries between explanatory domains are crossed. Instead, we should rethink Mayr’s distinction by adopting an alternative view of evolutionary causation, the so-called Statisticalist view, which maintains that the only level of causation is the individual level. By ruling out two different levels of causation, this framework is appropriate to avoid fallacious explanations and reconsider reciprocal causation entirely as a proximate phenomenon. We introduce the concept of “statistical reciprocity” to explain the statistical effects of reciprocal causality in populations and outline some ideas of “population ontogenetics” as a prominent framework for unifying development and evolution beyond interactionist positions.

The use of evolutionary explanations to explain phenomena at the individual level has been described by various authors as an explanatory error, the so-called Phylogeny Fallacy. In this paper, this fallacy will be analyzed in the context of teleosemantics, a central project of the philosophy of mind whose main aim is to naturalize intentional systems by appealing to their biological teleofunctions. I will argue that those teleosemantics projects that invoke evolutionary functions generally commit the fallacy. First, I will point to various arguments in the literature that point to this fallacy. However, a more general argument will also be made. To illustrate this puzzling scenario, I will present two desiderata that any teleosemantic project must fulfill. I will argue that naturalizing intentionality based on natural selection creates an explanatory gap between types and tokens. This gap prohibits an adequate explanation of the desiderata. To close the gap, teleosemantics invokes replicator biology, a view of inheritance that has already been identified to commit the fallacy. This leads teleosemantics into a complex situation: to close the gap, it must commit the fallacy.

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, I argue that a similar situation can be found in contemporary biological theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis avoid any substantial change by reinterpreting and explaining individual-level phenomena from a population perspective. Finally, I will approach this debate by advocating the statistical reading of natural selection, which holds that explanations by natural selection are statistical. I will argue that the above historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. The reinterpretation of concepts at the individual level is part of the explanatory framework of the Modern Synthesis and the empty role of development within this framework. Moreover, the statistical perspective adopted here allows for the integration of two explanatory models: population-statistical and individual-causal. Finally, I will argue that this pluralistic framework can help to define the explanatory scope of the different biological approaches in order to achieve a coherent integration of development into evolution without rejecting population thinking.

In evolutionary theory, Explanatory Externalism—one of the pillars of the Modern Evolutionary Synthesis—holds that natural selection is the sole adaptive force driving evolution. This paper highlights several challenges to Explanatory Externalism, primarily advanced by developmental biology and its various subfields and theories. Based on this debate, I examine the implications for one of the most established accounts of biological function: the selected-effect theory. While externalist readings of selected-effect theory are common, I argue a conditional claim: if and when SE is interpreted within an externalist framework, two central desiderata remain unresolved—the explanatory and the discriminatory desiderata. Internalist explanations are indispensable in evolutionary biology for two reasons. First, in connection with the explanatory desideratum, internalist explanations are necessary to account for the existence of traits—not only at their origin but also in their transgenerational persistence. Second, concerning the discriminatory desideratum, an internalist framework shows that traits may exist in nature without being either selected or accidental, arising instead from structural properties and constraints that render them necessary conditions for the organism’s existence. Due to the limits of an externalist SE, I propose a possible alternative in which selected functions are reconceptualized from an internalist perspective. Although this framework remains underdeveloped, key ideas can be outlined through the Agential Paradigm in biology. In this view, an organism’s capacity to regulate its environmental interactions generates specific trait functions, which in turn determine selection conditions across generations. Rather than treating selection as an external force imposing order on randomness, trait function itself drives selection—no selection without function. Selected functions are thus linked to the adaptive origins of traits and their persistence across phylogeny. From this agential perspective, developmental processes generate specific functions, which in turn cause the selection of traits. This article, consequently, offers an appraisal of selected-effect functions grounded in a reconceptualization of selection within the Agential Paradigm.

Abstract: This paper argues that a central explanatory role for the concept of Umwelt in theoretical biology is to be found in developmental biology, in particular in the effort to understand development as a goal-directed and adaptive process that is controlled by the organism itself. I will reach this conclusion in two (interrelated) ways. The first is purely theoretical and relates to the current scenario in the philosophy of biology. Challenging neo-Darwinism requires a new understanding of the various components involved in natural selection processes. An important prerequisite for the explanation is the ability to understand development in a teleological way. Here, the concept of Umwelt plays a crucial role: if organisms are responsible for generating adaptive variation in specific environments, we need a theory that explains the context-dependent nature of adaptively oriented processes. The Umwelt is thus a central element in determining the goal that an adaptive process pursues. The second path in my analysis also has a historical dimension. I will present Karl Ernst von Baer’s reflections on teleological development and his influence on von Uexküll's thinking. I will present various ideas developed by von Baer, such as the distinction between Ziel and Zweck and the use of musical metaphors, which can help to understand development teleologically and give von Uexküll’s theory a central place in this framework.

Abstract: The use of evolutionary explanations to account for proximate phenomena has been labeled by various authors as an explanatory error, the so-called phylogeny fallacy. In this paper, this fallacy will be analyzed in the context of teleosemantics. I will discuss whether teleosemantics projects that rely on the Selected-Effect Theory of Functions (i.e., mainstream teleosemantics) generally commit the fallacy. To frame the discussion, I will present two desiderata that, as argued here, every teleosemantic project must fulfill. The actuality desideratum, motivated by Brentano’s problem, demands for a proximate analysis of the properties that make a system intentional. The historical desideratum, on the other hand, says that a teleological view of intentionality requires a historical dimension in order to understand the causal basis of teleological explanations. While I give some reasons why mainstream teleosemantics might be implicated in the phylogeny fallacy, my main argument aims to highlight an explanatory gap in mainstream teleosemantics: the population perspective on teleofunctions is inadequate to explain the proximate properties of intentional systems. In other words, the account of the historical desideratum in population-evolutionary terms prevents an adequate solution to the actuality desideratum. This is related to two central goals of teleosemantics: The goal of explaining content determination by recourse to natural selection processes makes it impossible to solve another goal of teleosemantics, the so-called problem of representational status. I analyze this situation and argue that inherited information (sometimes interpreted in a teleosemantic sense) plays the role of closing the gap. This leads teleosemantics into a complex situation: in order to close the gap, it must commit the fallacy. At the end of the article, I propose an alternative to fulfill both desiderata, motivated by an evo-devo perspective. I state that lineage explanations have the property of providing historical explanations based on proximate causes.

The foundations of biology have been a topic of debate for the past few decades. The traditional perspective of the Modern Synthesis, which portrays organisms as passive entities with limited role in evolutionary theory, is giving way to a new paradigm where organisms are recognized as active agents, actively shaping their own phenotypic traits for adaptive purposes. Within this context, this article raises the question of whether contemporary biological theory is undergoing a cognitive revolution. This inquiry can be approached in two ways: from a theoretical standpoint, exploring the centrality of the cognitive sciences in current theoretical biology; and from a historical perspective, examining the resemblance between the current state of theoretical biology and the Cognitive Revolution of the mid-20th century. Both inquiries yield affirmative answers, though important nuances will be emphasized. The cognitive sciences' explanatory framework is employed to elucidate the agentic characteristics of organisms, establishing a clear parallelism between the Cognitive Revolution and the present state of theoretical biology

Disputes about the causal structure of natural selection have implications for teleosemantics. Etiological, mainstream teleosemantics is based on a causalist view of natural selection. The core of its solution to Brentano’s Problem lies in the solution to Kant’s Puzzle provided by the Modern Synthesis concerning populational causation. In this paper, I suggest that if we adopt an alternative, statisticalist view on natural selection, the door is open for two reflections. First, it allows for setting different challenges to etiological teleosemantics that arise if a statisticalist reading of natural selection is right. Second, by providing a different solution to Kant’s Puzzle based on individual causes of evolution, statisticalism promotes a different answer to Brentano’s Problem, what I label as Agential Teleosemantics.

El objetivo de este ensayo es analizar cómo confluyen en una misma base teórica tres discursos de diferentes áreas. Uno de ellos es la ciencia, mediante el genocentrismo defendido en la síntesis moderna en Biología. Otro es el de la religión, mediante la perspectiva finalista del alma y del diseño divino, que comparten las religiones abrahámicas. Y finalmente, en la esfera político-social, donde diferentes movimientos conservadores se oponen a replantear la agenda de derechos. La confluencia teórica que nuclea a las tres áreas mencionadas es un preformismo sobre los organismos. Por lo tanto, primero, veremos cómo el preformismo se encuentra presente en biología. Segundo, señalaremos dos ideas relacionadas al preformismo que subyacen tanto en la biología como en la religión. Finalmente, analizaremos cómo ambos discursos, el científico y el religioso, se encuentran presentes y empleando un rol argumentativo en debates actuales sobre derechos humanos, específicamente, relacionados al aborto ya los derechos de la población lgtbiq+. La principal conclusión es que la síntesis moderna promueve erróneamente una defensa de un preformismo genético, con connotaciones religiosas, que es usado por movimientos conservadores en el ámbito político-social.

The field of the philosophy of biology is flourishing in its aim to evaluate and rethink the view inherited from the previous century ---the Modern Synthesis. Different research areas and theories have come to the fore in the last decades in order to account for different biological phenomena that, in the first instance, fall beyond the explanatory scope of the Modern Synthesis. This thesis is anchored and motivated by this revolt in the philosophy of biology. The central target in this context is the possibility of naturalizing teleology, a classical nightmare for the history of biology itself. This requires, principally, understanding the causes of teleological explanations without assuming an unfashioned backward causation of sorts. As the riddles of teleological explanations are about their temporal dimension, I analyze different temporal scales of biological processes: evolutionary, developmental, and physiological. The first one is the one defended in the context of the Modern Synthesis. As expected, one of the aims of this thesis is to evaluate the adequacy of an evolutionary account of teleological explanation. The scrutiny is negative. Evolutionary explanations in the context of the Modern Synthesis lack the necessary causal roots to naturalize teleology. Concerning the physiological scale, a long tradition pushed up by Kant and the organicist movement in the 20th century allows us to better understand how teleological explanations can be naturalized in physiological process. The key notions in this temporal scale are self-organization and the recursive, looped character of physiological process. While the physiological scale may be suitably accounted by contemporaries views, such as Autonomous Systems Theory, different central teleological phenomena remain unexplained from a purely physiological perspective. In particular, different issues concerning the (adaptive) construction of organism ---such as plasticity, robustness, variation, novelty, inheritance--- deserve an ontogenetic analysis. The principal aim of this thesis is to provide a theory of teleological development that falls beyond the Modern Synthesis' framework and is prompted by different insights from the history of biology. I call it Agential Teleosemantics. It rests on two central pillars. First, that developmental processes, beyond any gene-centered stance, can be understood in informational terms; i.e. developmental processes are about the interaction of developmental resources conveying biological information. The second ingredient is agentivity, namely the idea that development is regulated by an agentive system according to the adaptiveness of the phenotypic outcomes produced. The role of agency in Agential Teleosemantic is equivalent to the role of genes in the Modern Synthesis: it is responsible for explaining the order and the adaptive complexity in the living realm. The second target of this thesis regards the possibility of naturalizing intentional explanations in cognitive science. The central project involved in such an aim is known as teleosemantics. Classical teleosemantics however is etiological: it explains the teleofunctions of representational systems in terms of evolutionary processes. The different disputes in the contemporary philosophy of biology provide two insights to analyze teleosemantics in cognitive science. First, the challenges against the Modern Synthesis must be extended to the evolutionary approach of etiological teleosemantics. Second, as Agential Teleosemantics suggests an alternative source for teleofunctions ---ontogeny, I offer an attempt to integrate Agential Teleosemantics into cognitive science in order to provide an alternative teleosemantic project to understand intentional explanations in cognitive science.