Javier Suárez

This paper presents the philosophical implications of the position that we call mereological perspectivism. Mereological perspectivism asserts that determining whether a composite biological object constitutes a single unit (an individual) necessarily depends on the selection of privileged parts of the composite. First, we show how mereological perspectivism is used in several debates in philosophy of biology (biological individuality, ecosystem individuality). Second, we show the implications of mereological perspectivism for the general discussions on scientific perspectivism. Finally, we show how it can illuminate the debates about biological essentialism.

Potentialities are properties that can manifest. Many potentialities are instantiated in virtue of having causal bases that are possible causes for their manifestations. This chapter shows that a closer look at potentialities and their causal bases studied in the life sciences (from cell biology to psychiatry) gives us good reasons to revise our philosophical assumptions about causal bases. The investigation reveals that causal bases are often more dynamic and dependent on interactions with the environment than previously assumed. In particular, we argue that causal bases in the life sciences are regularly extrinsic and processual and that potentialities in the life sciences have causal bases at different levels.

We propose that top‐down coercive mechanisms have played a role in the origin and maintenance of the Major Transitions in Evolution (MTE). Top‐down coercion has potentially been underappreciated due to the lack of a conceptual framework. Therefore, we provide a formalized top‐down coercion framework for the MTE. Our conceptualization of top‐down biological coercion is a loss of potential due to a constraint from the top‐down. We also present three case studies of coercive top‐down mechanisms in the evolution of eukaryotic cells, multicellularity and eusocial insect colonies. The MTE project studies the origin and maintenance of new levels of individuality in the biological hierarchy. Previously, the MTE has been conceived as a bottom‐up process. Our coercion framework provides new empirical questions regarding the origin of transitions and helps reframe discussions of fitness in the MTE.

This article defends a selective eliminativist position with respect to the concept of “biological lineage” as used in certain areas of contemporary evolutionary biology. We argue that its primary epistemic roles in these contexts—explaining social evolution and cumulative selection—clash with empirical evidence, and that enforcing the concept of “lineage” even obstructs fruitful research avenues in several biological research fields, including phylogenetic research. Drawing on this, we suggest that, in many instances, it would be best to get rid of the concept of “lineage” and think in terms that are more closely aligned with the empirical complexity of the biological world. Specifically, this entails that, instead of looking for the lineage that may have given rise to the entities possessing certain biological characteristics, biologists should generally look for any potential process allowing such possession, with the formation of a lineage just one possibility among many.

The method of metaphysics of scientific practice consists in developing metaphysical claims on the basis of empirical information from and about scientific practice. This method stands in the tradition of naturalistic or scientific metaphysics on the one hand, and philosophy of science in practice on the other. In this chapter we draw on some of our own research to specify the method at work. We argue that the method is typically carried out in four steps: identifying the available empirical information that might be relevant to the metaphysical topic of interest; selecting the relevant empirical information; developing a preliminary metaphysical claim that the relevant empirical information supports; mutually adjusting empirical information and metaphysics. We reveal some major methodological challenges and present ways to overcome them. We conclude that when drawing metaphysical claims from scientific practicewe need to: first, make sure that our previous metaphysical views do not influence the empirical information or its selection in problematic ways; second, explain in what sense we are doing metaphysics, countering for instance the idea that metaphysical claims must be highly general and transdisciplinary. We end the chapter by pointing to the advantages of the method of metaphysics of scientific practice.

We argue that microbiome research should be more reflective on the methods that it relies on to build its datasets due to the danger of facing a methodological problem which we call “epistemic misalignment.” An epistemic misalignment occurs when the method used to answer specific scientific questions does not track justified answers, due to the material constraints imposed by the very method. For example, relying on 16S rRNA to answer questions about the function of the microbiome generates epistemic misalignments, due to the different temporal scales that 16S rRNA provides information about and the temporal scales that are required to know about the functionality of some microorganisms. We show how some of these exist in contemporary microbiome science and urge microbiome scientists to take some measures to avoid them, as they may question the credibility of the field as a whole.

This paper addresses the topic of determinism in contemporary microbiome research. I distinguish two types of deterministic claims about the microbiome, and I show evidence that both types of claims are present in the contemporary literature. First, the idea that the host genetics determines the composition of the microbiome which I call “host-microbiome determinism”. Second, the idea that the genetics of the holobiont (the individual unit composed by a host plus its microbiome) determines the expression of certain phenotypic traits, which I call “microbiome-phenotype determinism”. Drawing on the stability of traits conception of individuality (Suárez in Hist Philos Life Sci 42:11, 2020) I argue that none of these deterministic hypotheses is grounded on our current knowledge of how the holobiont is transgenerationally assembled, nor how it expresses its phenotypic traits.

This paper critically examines Mitchell’s integrative pluralism. Integrative pluralism is the view that scientific explanations should primarily aim to integrate descriptions from different ontological levels. We contend that, while integrative pluralism is a fundamental strategy in contemporary science, there are specific reasons why one should not expect integration in the sense developed by Mitchell to be the optimal strategy and the one that scientists should always aim for. Drawing on some examples from contemporary biology, we argue that integration is sometimes neither epistemically desirable, nor ontologically achievable. We conclude that integrative pluralism should thus be limited to a specific class of complex systems but cannot be generalised as the preferable research strategy without further information about the epistemic practices of the scientific community or the ontology of the system under investigation.

This paper presents an account of the nature of stem cells based on the philosophical concept of disposition. It is argued that stem cells can be conceived as dispositional objects, and adopting this attitude allows overcoming some of the controversies surrounding the nature of stemness (most notably, the state vs. entity debate) because it offers a framework that accommodates the lessons from different theories. Additionally, the account is simultaneously useful for interpreting stem cell experiments and guiding potential interventions. The account shows how different levels, both molecular and emergent network-level, play the primary causal role in explaining some empirical results, and hence they suggest that the explanations can be mechanistic or topological, respectively. The realization that any of these levels may play a more prominent causal role than another allows suggesting interventions at the genetic, molecular and population levels.

Despite the importance of network analysis in biological practice, dominant models of scientific explanation do not account satisfactorily for how this family of explanations gain their explanatory power in every specific application. This insufficiency is particularly salient in the study of the ecology of the microbiome. Drawing on Coyte et al. (2015) study of the ecology of the microbiome, Deulofeu et al. (2021) argue that these explanations are neither mechanistic, nor purely mathematical, yet they are substantially empirical. Building on their criticisms, in the present work we make a step further elucidating this kind of explanations with a general analytical framework according to which scientific explanations are ampliative, specialized embeddings (ASE), which has recently been successfully applied to other biological subfields. We use ASE to reconstruct in detail the Coyte et al.’s case study and on its basis, we claim that network explanations of the ecology of the microbiome, and other similar explanations in ecology, gain their epistemological force in virtue of their capacity to embed biological phenomena in non-accidental generalizations that are simultaneously ampliative and specialized.

A general view in philosophy of science says that the appropriateness of an object to act as a surrogate depends on the user’s decision to utilize it as such. This paper challenges this claim by examining the role of surrogative reasoning in high-throughput sequencing technologies as they are used in contemporary microbiome science. Drawing on this, we argue that, in technology-driven surrogates, knowledge about the type of inference practically permitted and epistemically justified by the surrogate constrains their use and thus puts a limit to the user’s intentions to use any object as a surrogate for what they please. Ignoring this leads to a serious epistemic misalignment, which ultimately prevents surrogative reasoning. Thus, we conclude that knowledge about the type of surrogate reasoning that the technologies being used allow is fundamental to avoid misinterpreting the consequences of the data obtained with them, the hypothesis this data supports, and what these technologies are surrogates of.

I investigate the intrinsic/extrinsic nature of stemness in muscle stem cells (MSC) by relying on recent research on quiescence, with the aim of shedding light on the nature of dispositions and deriving some consequences about stem cells. First, I argue why the study of quiescence is the best available way to establish any claim about the intrinsicness/extrinsicness of stemness at least is some stem cells. Drawing on that, I argue that MSC’s stem capacities result from the combination of intrinsic cues plus extrinsic factors from the stem cell niche, making stemness an extrinsic disposition in MSC. Importantly, it is shown that the niche allows the instantiation of stemness in MSC by acting as a masker of its manifestation. This shows that stemness is, at least in MSC, what I call an extrinsically structurally masked disposition (SMD); that is, a disposition whose instantiation requires, as a condition of possibility, the interaction between the bearer and a masker. Finally, I conclude by suggesting some potential consequences of this observation for the philosophical study of dispositions and for stem cell research.

This paper introduces the reconstitutor as a comprehensive unit of heredity within the context of evolutionary research. A reconstitutor is the structure resulting from a set of relationships between different elements or processes that are actively involved in the recreation of a specific phenotypic variant in each generation regardless of the biomolecular basis of the elements or whether they stand in a continuous line of ancestry. Firstly, we justify the necessity of introducing the reconstitutor by showing the limitations of other evolutionary conceptions of the unit of heredity, such as the replicator, the reproducer, and the Darwinian individual. We argue that these conceptions are based on the requirement of lineage formation, which we argue to be unnecessary for the existence of evolutionary heredity. In the second part, we introduce the reconstitutor, which we base on the concept of Stability of Traits, and illustrate how it covers cases of hereditary phenomena not covered by the previous accounts. Secondly, we illustrate how the reconstitutor could serve as a platform to rethink ecological inheritance and other forms of inheritance that have been recently introduced under the song/singer model of evolution.

COVID-19 has substantially affected our lives during 2020. Since its beginning, several epidemiological models have been developed to investigate the specific dynamics of the disease. Early COVID-19 epidemiological models were purely statistical, based on a curve-fitting approach, and did not include causal knowledge about the disease. Yet, these models had predictive capacity; thus they were used to ground important political decisions, in virtue of the understanding of the dynamics of the pandemic that they offered. This raises a philosophical question about how purely statistical models can yield understanding, and if so, what the relationship between prediction and understanding in these models is. Drawing on the model that was developed by the Institute of Health Metrics and Evaluation, we argue that early epidemiological models yielded a modality of understanding that we call descriptive understanding, which contrasts with the so-called explanatory understanding which is assumed to be the main form of scientific understanding. We spell out the exact details of how descriptive understanding works, and efficiently yields understanding of the phenomena. Finally, we vindicate the necessity of studying other modalities of understanding that go beyond the conventionally assumed explanatory understanding.

RESUMENLa teoría evolutiva de Margulis implica una auténtica revisión de algunos de los conceptos fundamentales de la biología tradicional. Uno de tales cambios tiene que ver con el controvertido debate acerca de las unidades de selección. Este trabajo considera la propuesta de Margulis como una nueva tradición de investigación (TI) y trata de evaluar sus consecuencias para la citada disputa. Se sugieren tres ideas: una, que su teoría supone la revisión de conceptos tradicionales de la biología; dos, que su estudio supone una reevaluación de debates tradicionales de la filosofía de la biología; tres, que estudiarlo desde los presupuestos de Lakatos y Laudan sobre las TI nos permite iluminar la utilidad de su teoría. AbstractMargulis’ evolutionary theory entails a revision of certain core concepts of traditional biology. One of these changes is related to the hot debate about units of selection. This paper considers Margulis’ proposal as a new research tradition (RT) and evaluates its consequences to the mentioned issue. Certain evidence strongly suggests: first, that her theory supposes the revision of many classical biological concepts; second, that her position yields a reappraisal of many traditional issues in philosophy of biology; third, that its study from Lakatos and Laudan’s ideas about RT allows us to light the richness of her theory.

An understanding of the factors behind the evolution of multicellularity is one of today’s frontiers in evolutionary biology. This is because multicellular organisms are made of one subset of cells with the capacity to transmit genes to the next generation and another subset responsible for maintaining the functionality of the organism, but incapable of transmitting genes to the next generation. The question arises: why do somatic cells sacrifice their lives for the sake of germline cells? How is germ/soma separation maintained? One conventional answer refers to inclusive fitness theory, according to which somatic cells sacrifice themselves altruistically, because in so doing they enhance the transmission of their genes by virtue of their genetic relatedness to germline cells. In the present article we will argue that this explanation ignores the key role of policing mechanisms in maintaining the germ/soma divide. Based on the pervasiveness of the latter, we argue that the role of altruistic mechanisms in the evolution of multicellularity is limited and that our understanding of this evolution must be enriched through the consideration of coercion mechanisms.

Recent research in philosophy of science has shown that scientists rely on a plurality of strategies to develop successful explanations of different types of phenomena. In the case of biology, most of these strategies go far beyond the traditional and reductionistic models of scientific explanation that have proven so successful in the fundamental sciences. Concretely, in the last two decades, philosophers of science have discovered the existence of at least two different types of scientific explanation at work in the biological sciences, namely: mechanistic and structural explanations. Despite the growing evidence about the radically different nature of these two types of explanation, no inquiry has been conducted to date to determine the ontological reasons that might underlie these differences, nor the way in which these types of explanations can be systematically related with each other. Here, we aim to cover this gap by connecting this plurality of research strategies with the existence of emergent levels of reality. We argue that the existence of these different—and apparently incompatible—explanatory strategies to account for biological phenomena derives from the existence of “ontological jumps” in nature, which generate different regimes of causation that in turn demand the development of different explanatory frameworks. We identify two of these strategies—mechanistic modelling and network modelling—and connect them to the existence of two ontological regimes of causation. Finally, we relate them with each other in a systematic way. In this vein, our paper provides an ontological justification for the plurality of explanatory strategies that we see in the life sciences.

The units or levels of selection debate concerns the question of what kind of biological systems are stable enough that part of their evolution is a result of the process of natural selection acting at their level. Traditionally, the debate has concerned at least two different, though related, questions: the question of the level at which interaction with the environment occurs, and the question of the level at which reproduction occurs. In recent years, biologists and philosophers have discussed a new aspect of this debate, namely the possibility that certain multi-species consortia formed by a host and its microbiome may act as a unit of selection. This thesis, however, has not been without criticism, as it is doubtful that such consortia could meet the conditions required to achieve the degree of stability that would allow them to experience natural selection. The purpose of this paper is to systematically examine such criticisms and to defend the thesis that the holobiont/hologenome can act as a genuine level of selection both in the form of an interactor and in the form of a reproducer. To do so, it will be argued that the microbiome should be characterized in functional rather than taxonomic terms.

Holobionts are symbiotic assemblages composed by a macrobe host plus its symbiotic microbiota. In recent years, the ontological status of holobionts has created a great amount of controversy among philosophers and biologists: are holobionts biological individuals or are they rather ecological communities of independent individuals that interact together? Chiu and Eberl have recently developed an eco-immunity account of the holobiont wherein holobionts are neither biological individuals nor ecological communities, but hybrids between a host and its microbiota. According to their account, the microbiota is not a proper part of the holobiont. Yet, it should be regarded as a set of scaffolds that support the individuality of the host. In this paper, we approach Chiu and Eberl’s account from a metaphysical perspective and argue that, contrary to what the authors claim, the eco-immunity account entails that the microorganisms that compose the host’s microbiota are proper parts of the holobiont. Second, we argue that by claiming that holobionts are hybrids, and therefore, not biological individuals, the authors seem to be assuming a controversial position about the ontology of hybrids, which are conventionally characterized as a type of biological individual. In doing so, our paper aligns with the contemporary tendency to incorporate metaphysical resources to shed light on current biological debates and builds on that to provide additional support to the consideration of holobionts as biological individuals from an eco-immunity perspective.

The appeal to mechanisms in scientific explanation is commonplace in contemporary philosophy of science. In short, mechanists argue that an explanation of a phenomenon consists of citing the mechanism that brings the phenomenon about. In this paper, we present an argument that challenges the universality of mechanistic explanation: in explanations of the contemporary features of the eukaryotic cell, biologists appeal to its symbiogenetic origin and therefore the notion of symbiogenesis plays the main explanatory role. We defend the notion that symbiogenesis is non-mechanistic in nature and that any attempt to explain some of the contemporary features of the eukaryotic cell mechanistically turns out to be at least insufficient and sometimes fails to address the question that is asked. Finally, we suggest that symbiogenesis is better understood as a pragmatic scientific law and present an alternative non-mechanistic model of scientific explanation. In the model we present, the use of scientific laws is supposed to be a minimal requirement of all scientific explanations, since the purpose of a scientific explanation is to make phenomena expectable. Therefore, this model would help to understand biologists’ appeal to the notion of symbiosis and thus is shown to be better, for the case under examination, than the mechanistic alternative.

Given one conception of biological individuality (evolutionary, physiological, etc.), can a holobiont – that is the host + its symbiotic (mutualistic, commensalist and parasitic) microbiome – be simultaneously a biological individual and an ecological community? Herein, we support this possibility by arguing that the notion of biological individuality is part‐dependent. In our account, the individuality of a biological ensemble should not only be determined by the conception of biological individuality in use, but also by the biological characteristics of the part of the ensemble under investigation. In the specific case of holobionts, evaluations of their individuality should be made either host‐relative or microbe‐relative. We support the claim that biological individuality is part‐dependent by drawing upon recent empirical evidence regarding the physiology of hosts and microbes, and the recent characterization of the ‘demibiont’. Our account shows that contemporary disagreements about the individuality of the holobiont derive from an incorrect understanding of the ontology of biological individuality. We show that collaboration between philosophers and biologists can be very fruitful in attempts to solve some contemporary biological debates.

Contemporary biological research has suggested that some host–microbiome multispecies systems (referred to as “holobionts”) can in certain circumstances evolve as unique biological individual, thus being a unit of selection in evolution. If this is so, then it is arguably the case that some biological adaptations have evolved at the level of the multispecies system, what we call hologenomic adaptations. However, no research has yet been devoted to investigating their nature, or how these adaptations can be distinguished from adaptations at the species-level (genomic adaptations). In this paper, we cover this gap by investigating the nature of hologenomic adaptations. By drawing on the case of the evolution of sanguivory diet in vampire bats, we argue that a trait constitutes a hologenomic adaptation when its evolution can only be explained if the holobiont is considered the biological individual that manifests this adaptation, while the bacterial taxa that bear the trait are only opportunistic beneficiaries of it. We then use the philosophical notions of emergence and inter-identity to explain the nature of this form of individuality and argue why it is special of holobionts. Overall, our paper illustrates how the use of philosophical concepts can illuminate scientific discussions, in the trend of what has recently been called metaphysics of biology.

Bourrat and Griffiths :33, 2018) have recently argued that most of the evidence presented by holobiont defenders to support the thesis that holobionts are evolutionary individuals is not to the point and is not even adequate to discriminate multispecies evolutionary individuals from other multispecies assemblages that would not be considered evolutionary individuals by most holobiont defenders. They further argue that an adequate criterion to distinguish the two categories is fitness alignment, presenting the notion of fitness boundedness as a criterion that allows divorcing true multispecies evolutionary individuals from other multispecies assemblages and provides an adequate criterion to single out genuine evolutionary multispecies assemblages. A consequence of their criterion is that holobionts, as conventionally defined by hologenome defenders, are not evolutionary individuals except in very rare cases, and for very specific host-symbiont associations. This paper is a critical response to Bourrat and Griffiths’ arguments and a defence of the arguments presented by holobiont defenders. Drawing upon the case of the hologenomic basis of the evolution of sanguivory in vampire bats, I argue that Bourrat and Griffiths overlook some aspects of the biological nature of the microbiome that justifies the thesis that holobionts are evolutionarily different to other multispecies assemblages. I argue that the hologenome theory of evolution should not define the hologenome as a collection of genomes, but as the sum of the host genome plus some traits of the microbiome which together constitute an evolutionary individual, a conception I refer to as the stability of traits conception of the hologenome. Based on that conception I argue that the evidence presented by holobiont defenders is to the point, and supports the thesis that holobionts are evolutionary individuals. In this sense, the paper offers an account of the holobiont that aims to foster a dialogue between hologenome advocates and hologenome critics.

Philippe Huneman has recently questioned the widespread application of mechanistic models of scientific explanation based on the existence of structural explanations, i.e. explanations that account for the phenomenon to be explained in virtue of the mathematical properties of the system where the phenomenon obtains, rather than in terms of the mechanisms that causally produce the phenomenon. Structural explanations are very diverse, including cases like explanations in terms of bowtie structures, in terms of the topological properties of the system, or in terms of equilibrium. The role of mathematics in bowtie structured systems and in topologically constrained systems has recently been examined in different papers. However, the specific role that mathematical properties play in equilibrium explanations requires further examination, as different authors defend different interpretations, some of them closer to the new-mechanistic approach than to the structural model advocated by Huneman. In this paper, we cover this gap by investigating the explanatory role that mathematics play in Blaser and Kirschner’s nested equilibrium model of the stability of persistent long-term human-microbe associations. We argue that their model is explanatory because: i) it provides a mathematical structure in the form of a set of differential equations that together satisfy an ESS; ii) that the nested nature of the ESSs makes the explanation of host-microbe persistent associations robust to any perturbation; iii) that this is so because the properties of the ESS directly mirror the properties of the biological system in a non-causal way. The combination of these three theses make equilibrium explanations look more similar to structural explanations than to causal-mechanistic explanation.

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages.

Margulis’ evolutionary theory entails a revision of certain core concepts of traditional biology. One of these changes is related to the hot debate about units of selection. This paper considers Margulis’ proposal as a new research tradition (RT) and evaluates its consequences to the mentioned issue. Three ideas are suggested here: firstly, that her theory represents the revision of many classical biological concepts; secondly, that her position implies a reappraisal of many traditional issues in philosophy of biology; and thirdly, that studying it from Lakatos and Laudan’s ideas about RT allows us to enlighten the richness of her theory.