The relationship between development and evolution has recently becomea lively debated topic among philosophers and biologists. Thisinterest has been increasingly stirred through at least sixdevelopments since the 1990s: First, new findings of the moleculargenetic mechanisms underlying the development and evolution havetriggered new ideas about evolutionary change. These discoveries andconceptual innovations eventually led to the foundation of the newfield of evolutionary developmental biology (evo-devo). Second, theability to rapidly sequence genes and genomes allowed geneticcomparisons to be made between species. Prior to this, evolutionarygenetics was confined to allelic differences within a species. Third,new discoveries of environmentally sensitive channels of extra-genetictransmission of information between organisms (e.g., epigeneticinheritance and transmission of microbiota) led to attempts to moreclosely connect development and inheritance, and, as a consequence,evolution. Fourth, developmental plasticity, the ability of theenvironment to elicit different normal phenotypes from the samegenotype, was found to be universal among multicellular organisms, andthe molecular mechanisms by which environmental agents generate thesealternative phenotypes have been elucidated in many cases. Fifth,eukaryotic organisms now appear to be holobionts, consortia ofdifferent species working in mutualistic symbioses. Moreover, almostall multicellular organisms develop as holobionts, where symbiontsprovide signals for development. This has opened new questionsconcerning whether evolution can occur by altering symbiontrelationships during development. Sixth, the behavioral patternsdeveloped by organisms are increasingly discussed not only as effectsof adaptive processes, but as starting point of evolutionarytrajectories. This includes hypotheses conceptualizing organisms asagents that co-modify the selective pressures effecting them (andother species).

These recent trends towards intertwining, rather than separating, theconcepts of development and evolution is in contrast to authoritativevoices in evolutionary biology (and to a substantial part inphilosophy of biology) during most of the twentieth century (see Sapp1983; Burian 2004; Amundson 2005). As a consequence of this tradition,this new integration faces a number of conceptual and methodologicalchallenges. In this entry, the focus will be on debates associatedwith the relationship between development and evolution. Otherdiscussions about what each of the two components, development andevolution, are or how they are (or should be) conceptualized orstudied individually is only addressed when relevant to this maintopic.

• 1. Evolution and development in historical context
• 2. Conceptual partings and unifications of evolution and development
  • 2.1 The proximate-ultimate distinction
  • 2.2 The integration of proximate and ultimate causes
• 3. Developmental change as the mechanism for evolution
• 4. Ontological challenges of developmental evolution
• 5. Explanations of developmental evolution
  • 5.1 Mechanistic explanations of developmental evolution
  • 5.2 The explanatory power of developmentalist explanations of evolution
• 6. Organismal agency in development and evolution
• 7. Anthropological and ethical dimensions of developmental evolution
• 8. Concluding Remarks
• Bibliography
• Academic Tools
• Other Internet Resources
• Related Entries

1. Evolution and development in historical context

Originally, the concepts of evolution and development were closelyconnected. In fact, since the end of the 17^th century theconcept of ‘evolution’ was widely used to describeindividual developmental processes, and ‘developmentalhypotheses’ often referred to what is now called evolution. Inaddition, development (‘Entwicklung’) was often consideredto not only describe ontogenetic changes in organisms (Goethe 1790;Debraw 1777) but also (what we consider today) phylogenetic changes.For example, Friedrich W.J. Schelling (1798) states that the sequenceof stages of all organic beings took shape through the gradualdevelopment (‘Entwicklung’) of the same organization. Thissituation changed in the beginning of the 19^th century whenevolution was used by some authors for labeling transgenerationaltransformations of organisms, among others by Charles Lyell and GeorgW.F. Hegel. However, in his first edition of hisOrigin ofSpecies (1859) Darwin did not use the term evolution, likelybecause he wanted to distance his theory from earlier developmentalunderstandings of the word. Instead, he spoke of ‘descent withmodification.’ But especially due to the work of HerbertSpencer, evolution became established as a concept that concerns the“transformation of species”, and Darwin’s theory oftransformation through selection was increasingly seen as the exemplarof a “theory of evolution” (Spencer 1852 [1891], 1).According to this theory, evolution is driven through organisms“struggle for existence” (Darwin) and the “survivalof the fittest” (Spencer).

Besides this increasing conceptual parting, however, evolution anddevelopment were considered to be closely interrelated processes bymany scholars throughout the 19^th century. By building onearlier comparative studies of embryonic processes across various taxa(by, among others, Karl E. von Baer (1827)), Ernst Haeckel (1872)argued that all metazoans have the same early stages of development(see Gould 1977). He understood these developmental patterns asmodifications of a basic type. According to this biogenetic law, thedevelopment of individuals is a recapitulation of species’evolutionary changes:

During the rapid and short course of its individual development, theorganic individual […] repeats the most important changes ofform, which its ancestors passed through during their long and slowcourse of paleontological development, following the laws of heredityand adaptation. (Haeckel 1866, II 300)

This idea had already been anticipated by Johann F. Meckel (1812), andby Richard Owen (1837) who parallelized ‘transmutations’of embryonic forms and transmutations of species. Also Darwin (1837)describes the genesis of individuals as a ‘shortenedrepetition’ of morphological changes during phylogeny. In 1864,Fritz Müller’sFür Darwin showed thathomologous embryonic and larval structures showed phylogeneticrelationships between animal groups and that larvae were also subjectto selection and that, therefore, the larvae of different speciesshould be expected to diverge from each other. From the end of the19^th century the view of development as a recapitulation ofevolution became problematic (see Rasmussen 1991). First, during thistime the view of a linear evolution from invertebrates oververtebrates to humans was increasingly questioned. Second, based oncomparative studies of cell lineages across different taxa in the1890s, scholars criticized that not only adult organisms but theirdevelopmental pathways evolve, too. For example, in 1898, embryologistEdmund B. Wilson (1898) was using homologous patterns of embryoniccell division to unite taxa, while embryologist Frank R. Lillie (1898)studied differences in these patterns to show the origins ofevolutionary adaptations. These studies were (at least to some part)in conflict with Haeckel’s recapitulationism, as they sawontogeny not as sufficiently explained by historical causes, butrather in need of independent mechanistic laws operating on physicalproperties of the ontogenetic material (see Berrill & Liu 1948;Guralnick 2002). In addition, this cell lineage research suggestedthat Haeckel’s law on the relationship between development andevolution should be inverted, as cell lineages evolve overevolutionary time. In more general terms, this means that developmentdoes not merely mimic evolution. Rather, it is a source of variationand thus a cause of evolution and adaptation: “ontogeny does notrecapitulate phylogeny, it creates it” (Garstang 1922: 81). Inshort, evolution proceeds through modifications of development.

Another characteristic of 19^th century (and early20^th century) biology was the lack of a clear conceptualparting between developmental and reproductive processes. Especiallythe idea of the inheritance of acquired characteristics was part ofthe mainstream of evolutionary thought (Bowler 1983, 2017; Gissis& Jablonka 2011). According to Jean Baptiste Lamarck (1809),changes in the structure or function of traits that occurred during anorganism’s life, depending on their use or disuse, could beinherited. Darwin adopted this idea in his inheritance theory ofpangenesis (see Holterhoff 2014). He suggested that all body parts, ateach developmental stage, contain small particles, so-called‘gemmules,’ that were sensitive to environmental changes.These particles accumulate in the reproductive organs and aretransmitted to the next generation. Thus, gemmules modified duringdevelopment lead to modifications in the next generation. Darwinstated that due to this close connection of development andreproduction, “inheritance must be looked at as merely a form ofgrowth” (Darwin 1868: II, 404). However, this connection betweendevelopment and inheritance was increasingly criticized in the firsthalf of the 20^th century. Following August Weismann (1892),the inheritance of environmentally induced variation in somatic cells(the body) of organisms was increasingly called into question. Insexually reproducing organisms, only the germ plasm used for the spermand eggs were recognized as carrying information that are passed on tothe next generation. According to this view, the germ line was immunefrom variation occurring in the somatic cells of the body, and thusthe inheritance of acquired characteristics or other theories ofplasmatic inheritance were rejected (but see Sapp 1983; Harwood 1993;Jablonka & Lamb 2014; Gilbert & Epel 2015).

This parting between development, on the one side, and inheritance andevolution, on the other, through Weismann’s theory ofinheritance, had long lasting effects on how biologists reasoned aboutevolution in the following years (see McCord 2024). Despite continuingefforts of neo-Lamarckians well into the 1930s (Bowler 1983),development increasingly vanished from the landscape of evolutionarytheory. By uniting Mendel’s laws and new findings in geneticswithin the statistical framework of population genetics, the modernsynthesis came to understand evolution as a change in the frequenciesof different alleles in a population (Fisher 1930; Dobzhansky 1951).Variation relevant to evolution was produced only in the genes of thegerm line. These random changes, mutations, were screened off from thedevelopmental history of the individual. In the 1940s and 50s thisview was increasingly manifested, not least through the so-called‘central dogma’ of molecular biology (Crick 1958). Itstates (similar to Weismann’s theory) that information flowsalways from the DNA to proteins, never vice versa. Thus, phenotypicchanges during development could not affect the genes. The trend tolink genes with populations in evolutionary explanations, rather thandevelopment with evolution reached its climax with the gene’seye view of evolution (Williams 1966; Dawkins 1976). By focusing onthe question how certain traits like altruistic behaviors could bebeneficial, it identified genes, rather than organisms, as the soleunits of selection. In addition, it described development as nothingbut the readout of a genetic program (Maienschein 2003; Pigliucci2010). Genes were seen to control the development of traits andorganisms’ behaviors, and they replicate to secure their ownfurther propagation in groups and populations.

During the twentieth century these partings of development andevolution led to statements such as: “Problems concerned withthe orderly development of the individual are unrelated to those ofthe evolution of organisms through time” (Wallace 1986: 149).However, such statements were also targeted by a number of critics.For example, the exclusion of developmental biology from the modernsynthesis (Harrison 1937; De Beer 1954; Waddington 1957; Hamburger1980), the explanatory autocracy of the adaptationist program inevolutionary biology (Gould & Lewontin 1979), and the omission ofa satisfying theory of evolutionary novelty, in which developmentwould play a role (Goldschmidt 1940), have been criticized. Not leastdue to this rejection of developmental perspectives, evolutionarytheory has been blamed to constrain the direction of research inevolutionary biology (Provine 1989; Amundson 2005). In fact,throughout the twentieth century a number of theories have been putforward, which argue that due to the partial independence of geneticand phenotypic variation, evolutionary research should put moreemphasis on how the changes in developmental and behavioral patternsmight drive or bias evolutionary change. This includes cases of‘canalized’ developmental pathway which are maintained,even though the genotype or the environment might have changed to somedegree (Waddington 1957; see also Nijhout 2002; Gilbert & Epel2015; Sultan 2015). In addition, studies of phenotypic plasticityshowed that a trait of an organism can react to an environmental inputin various ways, and that the genome codes for a wide range ofpotential phenotypes (Waddington 1942; Nijhout 1990; Pigliucci 2001).The evolutionary relevance of these findings was emphasized by studiesthat investigated how the range of variation changes from a plastictrait to a fixed or canalized one over the course of generations(Suzuki and Nijhout 2008; West-Eberhard 2003; B. Baker et al. 2019).What is more, according to the so-called ‘Baldwin effect,’learned behavioral patterns (e.g., the acclimatization to a newstressor) that were initially rather plastic, can affect thereproductive success of individuals and thus, across generations andthrough natural selection, be gradually incorporated into the geneticand epigenetic makeup of a species (Baldwin 1896; Simpson 1953; Piaget1976 [1978]; Newman 2002). While some of these developmentalperspectives on evolution were consistent with the population geneticframework of evolutionary biology and were, in fact, graduallyincorporated into evolutionary thought, others posed more seriouschallenges for theoretical integration.

By building on the latter, more problematic set of approaches(especially in recent years), evolutionary theory has been facingcalls from developmentally oriented biologists and philosophers ofbiology to widen the standard explanatory and methodologicalapproaches on evolution (Bonner 1958; Alberch 1982; Bonner 1982; Raff& Kaufman 1983; Gilbert, Opitz, & Raff 1996; Schlichting &Pigliucci 1998; West-Eberhard 2003; G. Müller 2007; Pigliucci& Müller 2010; Bateson & Gluckman 2011; Jablonka &Lamb 2014; Laland et al. 2014, 2015; Gilbert & Epel 2015; G.Müller 2017; for discussion, see Futuyma 2017; Huneman &Walsh 2017; Fábregas-Tejeda & Vergara-Silva 2018; Baedke etal. 2020b; Edelaar et al. 2023, Lala et al. 2024). In line with olderaccounts, they argue that evolutionary change should not only orprimarily be investigated and explained as a change in genotypefrequencies in populations but (also) on the level of the developingand acting individual – the organism (see Baedke andFábregas-Tejeda 2023). The underlying idea is that phenotypicvariation and the flexibility of organisms’ responses toenvironmental cues may introduce non-random variation and thus maybias and/or direct morphological evolution to some degree. Thisincludes not only environmentally induced changes in regulatoryprocesses but also the physical constraints of the developing embryo.By focusing on these phenomena, Kevin Lala and colleagues (Laland etal. 2014: 161) state: “An alternative vision of evolution isbeginning to crystallize, in which the processes by which organismsgrow and develop are recognized as causes of evolution”. Othershave argued that genes are probably more often followers in evolutionthan leaders (West-Eberhard 2003, 2005). In other words, theenvironmentally responsive, developing and acting organism takes thelead. It introduces in a non-random way new phenotypes intopopulations that are subsequently stabilized by genes.

This ‘development first view’ (or sometimes called‘plasticity first view’) of evolution is currently adoptedespecially by researchers in three fields of research: evolutionarydevelopmental biology (evo-devo), epigenetics, and niche constructiontheory. Evo-devo studies how developmental pathways evolve and, moreimportant for the above view, how developmental constraints and biasescan affect evolutionary trajectories (Raff 1996, 2000; Gerhart &Kirschner 1997; Love 2003; Amundson 2005; Laubichler & Maienschein2007; Sansom & Brandon 2007; G. Müller 2007; Minelli 2022;Laubichler 2010; Pigliucci & Müller 2010; Gilbert & Epel2015; Moczek et al. 2015; Levis & Pfennig 2020; Nuño de laRosa & Müller 2021; Lala et al. 2024). In other words,because of the existing developmental mechanisms, some trajectories ofdevelopment are more readily available than others. This opens upareas of questions that include asking how changes in gene regulatorynetworks cause modifications of developmental processes (associatedwith major shifts in morphological ‘body plans’) and thusproduce evolutionary novelties, or what capacities organisms have togenerate heritable, adaptive phenotypic variation and thus to evolvein evolution (i.e. their evolvability; see Hendrikse et al. 2007).This focus on how phenotypic variation is produced, rather than how itis selected, is supported by studies in epigenetics on inter- andtransgenerational epigenetic inheritance (see the entryinheritance systems), like through the transmission of regulatory factors of gene activity(Jablonka & Lamb 1989, 2014, Jablonka & Raz 2009, Perez &Lehner 2019). Other forms of extra-genetic inheritance include thetransmission of microbiota (Gilbert et al. 2012, 2014, Browne et al.2017) and behaviorally mediated parental effects (Kappeler &Meaney 2010; Rilling & Young 2014). This set of processes containsenvironmentally sensitive non-genetic sources of variation fororganisms that can be transmitted across generations, in many casesdecoupled from the transfer of genetic information. Finally, nicheconstruction theory highlights the self-perpetuating and reciprocaleffects of organisms, as agents, that construct their own niche(and/or that of other species) during development (Lewontin 1982;Sterelny 2001; Day et al. 2003; Odling-Smee et al. 2003; Laland et al.2008, 2009, 2011; Odling-Smee 2010; Chiu & Gilbert 2015, 2020;Aaby & Ramsey 2022). These recent studies on developmentalevolution will be discussed in detail in section 3.

The various developments in the above fields have led to an increasingphilosophical interest in conceptual and explanatory issues arisingfrom the new junction of development and evolution. These includeclarifying what causal relationships and boundaries exist between thetwo realms, which different roles developmental processes play inrecent evolutionary research, and how these new roles affectontological frameworks of the organism. In addition, thesedevelopments have been accompanied by debates about the structure ofexplanations in developmental evolution. Finally, they have informeddiscussions on anthropological understandings and ethical questionsconcerning humans. We will now discuss the debates on these issues indetail.

2. Conceptual partings and unifications of evolution and development

One of the most central topics of early philosophy of biology in the1960s and 1970s was the attempt to develop a suitable conceptualframework that would support the parting between development andevolution in line with the central assumption of the Modern Synthesisthat evolution is a change in the genetic composition of populationsonly (Dobzhansky 1951: 16; see also Charlesworth et al. 2017). Thismeans, as a consequence, that development does not (or not insignificant ways) causally effect evolution. Over the decades thisassumption has been supported by the historically influentialconceptual distinction between proximate causes and ultimate causes(Mayr 1961).

2.1 The proximate-ultimate distinction

The dual framework ‘proximate vs. ultimate’ provides aqualitative distinction of biological causality (for relateddistinctions, see J. Baker 1938; Tinbergen 1951, 1963). It holds thatbiologists who study proximate causes ask how questions aboutindividual developmental processes. Thus, functional biologistsinterested in such proximate causes study how systems work. Instead,evolutionary biologists that study ultimate causes ask why questions,like why phylogenesis has produced particular evolutionary functions.According to this distinction, at least on the surface, proximatecauses resemble Aristotelian efficient causes while ultimate causesresemble Aristotelian final causes. To illustrate this distinction,Mayr (1961) draws on an example of avian migration. Migration can bestudied by asking how birds migrate (i.e., how they develop skillslike navigation) or why they migrate (i.e., due to what selectiveadvantage). These two investigations are understood to be bothimportant and complementary. However, they should be treated asdistinct from one another.

The proximate-ultimate distinction can be given an epistemic orontological reading. First, authors have interpreted it asdistinguishing different kinds of explanations (Amundson 2005; Calcott2013; Scholl & Pigliucci 2015). This epistemic reading includesthat how questions cannot be addressed by explanations citing ultimatecauses (i.e., telling a story of adaptation) and that why questionscannot be addressed by explanations citing proximate causes (i.e.,telling a story of trait development). Second, authors haveinterpreted this distinction as one between different ontologicalclasses of causes working in ontogenetic and phylogenetic processes(Laland et al. 2013a). This ontological reading is backed uptheoretically by Weismann’s concept of the separation of germline and soma, which provides a demarcation line between two distinctclasses of causes. To this day, biologists and philosophers have notreached a consensus on how exactly the division‘proximate-ultimate’ or ‘how-why’ should beunderstood, epistemically or ontologically (Francis 1990; Dewsbury1992, 1999; Sterelny 1992; Beatty 1994; Ariew 2003). Despite this lackof agreement this framework has been applied in various fields, fromevolutionary biology (E. O. Wilson 1975 [2000: 23]), evolutionarypsychology (Daly & Wilson 1978; Crawford 1998) and behavioralecology (Morse 1980: 92–95) to human sciences, like in humancooperation (Marchionni & Vromen 2009) and developmentalpsychology (Lickliter & Berry 1990). Especially in evolutionarybiology it has contributed to mainstream causal reasoning for a longtime, even among evolutionary biologists interested in developmentalprocesses (see, e.g., Maynard Smith 1982: 6).

2.2 The integration of proximate and ultimate causes

There has been constant criticism of the proximate-ultimatedistinction (since even before Mayr 1961), and against its underlyingidea to downgrade the explanatory or causal relevance of developmentto evolution. More recently, the discussion of this issue gained pacethrough new findings in fields such as epigenetics, evo-devo and nicheconstruction theory (Thierry 2005; Laland et al. 2011, 2013a, 2013b;Haig 2011, 2013; Scott-Phillips et al. 2011; Dickins & Rahman2012; Guerrero-Bosagna 2012; Calcott 2013; Dickins & Barton 2013;Gardner 2013; Mesoudi et al. 2013; Martínez & Esposito2014; Scholl & Pigliucci 2015; Baedke 2018; Uller & Laland2019; Brown 2021). In this context, some scholars argue that theproximate-ultimate distinction stands “at the center of some ofcontemporary biology’s fiercest debates” (Laland et al.2011: 1512) about the role of developmental plasticity, nicheconstruction and inclusive inheritance for evolutionary trajectories.Participants in this debate have argued that we should, due todifferent epistemic or heuristic reasons, keep Mayr’sproximate-ultimate distinction (Scott-Phillips et al. 2011; Dickins& Barton 2013) or a revised or reinterpreted form of it (Scholl& Pigliucci 2015; Otsuka 2015), expand it by a third intermediateform of explanations (Haig 2013), or replace it with a concept of‘reciprocal causation’ (Laland et al. 2011, 2013a, 2013b,2015; Mesoudi et al. 2013). In line with earlier philosophical work(Oyama 1985; Keller 2010; Griffiths & Stotz 2013), the latter ideaof reciprocal causation should allow describing the feedback processesbetween causal factors in evolving systems. This includesorganisms’ capacity of phenotypic plasticity or, morespecifically, their activities to alter selection pressures.Paradigmatic feedback cases are niche construction behaviors oforganisms that modify their environments and thus shape naturalselection pressures working on them. In other words, reciprocalcausation holds that organisms are not only effects of adaptiveprocesses, but also causal starting points of evolutionarytrajectories. In this sense this framework argues against the causaland/or explanatory asymmetry claim of the proximate-ultimatedistinction. It highlights the important role of development forevolution.

Against this new approach, scholars have argued that reciprocalcausation does, in fact, not pose any conceptual challenges forevolutionary biology, as it has been included since quite some timeago in the field (Svensson 2018). A true challenge, however, is todevelop this idea into a methodologically sound framework that allowsstudying and modeling complex non-linear relations between organismsand environments without merging the two into one inextricable unit(Baedke et al. 2021). Other have cast doubt on the central causal rolethe unit of the organism is supposed to play in this reciprocityframework (Baedke 2019a), or questioned whether this conceptualizationcan, in fact, capture all causal dependency relations of interest forevolutionary biology (Martínez & Esposito 2014; Scholl& Pigliucci 2015). Moreover, some argued that also this frameworkrelies on the dichotomy between development and evolution (Dickins& Barton 2013; Martínez & Esposito 2014) and that it isnot conducive to successful biological science, as it does not lead tofalsifiable questions (Dickins & Rahman 2012) and bleeds proximateand ultimate explanations into each other so that their distinctionbecomes meaningless (Gardner 2013; one should mention, however, thatthis might be the very aim of this approach). More generally, it hasbeen requested that advocates of this approach should provide moreconceptual clarifications on what reciprocal causation actually issupposed to mean (Buskell 2019). On an empirical level, a surveysuggested that biologists hold ambiguous positions about thedescriptive and empirical accuracy, explanatory merits and practicalutility of the conceptual framework of reciprocal causation (Hazelwood2023).

Besides distinguishing development and evolution in a qualitativemanner as proximate and ultimate causal processes, a less commonattempt is to quantitatively distinguish (or relate) the two. Here,especially distinctions based on the rates or time scales on whichdifferent developmental and biological processes occur have been made(see the entrylevels of organization in biology). For example, Conrad H. Waddington (1957) developed a hierarchicalmodel of time scales that includes biochemical processes on lowermolecular levels of organization with a faster rate, medium pacedprocesses of development on a medium level, and evolutionary processeson higher levels with a slower rate. According to such a view,evolutionary processes are simply processes occurring with a differentrate and thus at a different level than developmental ones. Thus, theydiffer gradually rather than in kind. Rate-based distinctions havebeen described to be consistent with the ultimate-proximate framework(when interpreting it as one that distinguishes different timescalesof phenotypic change; see Francis 1990; Haig 2013) or as differentfrom proximate-ultimate distinctions (Baedke & Mc Manus 2018). Inaddition, time-scale (or size-scale) conceptualizations have beenapplied for developing methodologies and multi-scale modeling thatintegrate, among others, developmental and evolutionary processes (S.Levin 1992; Green & Batterman 2017; Duckworth 2019).

Holobionts and developmental plasticity add new layers of complexityto the challenge of integrating proximate and ultimate causes acrossreciprocal organism-environment relationships. In mammalian holobionts(Gilbert et al. 2015, 2024), for instance, the microbes are part ofthe host’s environment, while the host is the environment forthe microbes (Formosinho et al. 2022). Moreover, in mammalianholobionts, the microbial portion of the organism can change andevolve quicker than the mammalian portion. In addition, as ErnestEverett Just (1933) and Richard Lewontin (1983) have argued,developmental plasticity means that evolution is not merely about theorganism but concerns changes in the organism-environment system.

3. Developmental change as the mechanism for evolution

The idea that evolutionary and developmental changes are closelylinked is central for the field of evolutionary developmental biology(evo-devo). The field describes itself as the science that studies howalterations in development create the variations that nature canselect (Raff & Kaufman 1983; Gilbert, Opitz, & Raff 1996). Inother words, natural selection did not create variation; developmentcreates variation. Development is the artist; natural selection is thecurator (Gilbert 2006, 2019). Both have creative agency; but they areworking at different levels. Although this view had been expressed byscientists such as Thomas Huxley, Julian Huxley, Conrad H. Waddington,and Richard B. Goldschmidt, it gained credence through more recentdiscoveries that explained how normal development could occur. Chiefamong these discoveries was the explication of developmental pathwaysthat connected embryonic induction with gene expression. Here,paracrine factors (proteins that influence the behaviors or geneexpression patterns of neighboring cells) secreted by one set of cellswere received by receptors on the membranes of other cells. Thesereceptors then activated proteins within the cytoplasm, whicheventually activated or repressed proteins that entered the nucleus toregulate transcription of particular genes.

The second major discovery that promoted evo-devo was the discovery ofmodular enhancers. The above-mentioned transcription factors wouldbind to specific regions of DNA, called enhancers. Most genes havemultiple enhancers. Thus, a gene might have a ‘limb’enhancer that activates the expression of the gene in the limb, and an‘eye’ enhancer that enables the expression of the gene inthe eye. Moreover, each enhancer usually binds several transcriptionfactors and could be activated in Boolean <and> or <or>fashion. There are also enhancers whose bound transcription factorsinhibit gene expression. Evolution could occur by creating or deletingenhancers, thereby enabling genes to be expressed differently indifferent species. King and Wilson (1975) and Jacob (1977) hadspeculated that evolution occurred by changes in gene regulation. Thisprovided a model for such regulation, and some of the best examplesare seen in the divergence of humans from other apes (Geschwind &Konopka 2012; Pollard et al. 2006). As Haraway (2008) noted,“relationships are the smallest possible pattern foranalysis”, and the relation between enhancer and transcriptionfactor may indicate the ‘natural kinds’ of the biologicalworld (Gilbert & Bard 2014). Moreover, the entities defined byenhancer/transcription factor interactions during development areoften unexpected and do not mirror intuitive boundaries betweenentities in adults. Activation of genes to form the distal rib, forinstance, is controlled by a different enhancer than that whichactivates genes in the proximal rib (Guenther et al. 2008).

These differences in gene expression could be categorized into fourcategories (Arthur 2004). One of these categories involves theplace of gene expression, where different populations ofcells express a particular gene in different species. For instance,thegremlin gene in the duck hindlimb webbing protects thesecells from cell death, enabling webbed feet (Laufer et al, 1997;Merino et al. 1999). A second category involves changes in thetiming of gene expression, as in the continued expression ofthefgf8 gene at the tip of the dolphin forelimbs, thusenabling the extension of its flippers (Richardson &Oelschläger 2002). A third category of change involvesalterations in themagnitude of gene expression, as in thedifferences inBmp4 gene expression that determine the widthof finch beaks (Abzhanov et al. 2004). A fourth category focuses onthe alterations of the actualprotein sequence of regulatoryproteins, as in the changes of theAntennapedia gene ininsects, which restrict insects from forming more than six limbs(Galant and Carroll 2002).

The third discovery was the elucidation of gene regulatory networks(GRNs). A GRN is based on paracrine factors, signal transductioncascades, and transcription factors. While ignoringpost-transcriptional gene regulation, this concept attempts to explainhow initial conditions (RNAs and proteins within the oocyte, positionof the embryo within the uterus, etc.) could create the conditionswhereby cell types differ, even though their genomes are identical.Spearheaded by Eric Davidson (2001, 2006), this approach uses systemstheory concepts such as modularity and dissociability to explain howthe genes interact in a hierarchical manner to produce different celltypes (Levine & Davidson 2005) and how the cell types in relatedspecies could differ by the recruitment (co-option) of a particularGRN by altering transcription factor binding. More generally, thediscovery of GRNs has enabled the integration of developmental biologywith paleontology (Jablonski 2017; Hinman et al. 2003; Hinman &Cheatle Jarvela 2014) and may also be extended into areas ofsymbiosis, niche construction, and the evolution of eusociality(Laubichler & Renn 2015; Haana & Abouheif 2021).

A fourth discovery of evolutionary developmental biology was theimportance of developmental plasticity for evolution (Nijhout 1990;Hall 1992; Gilbert 2001; Pigliucci 2001). By the end of the 20thcentury, the roles of temperature, sunlight, diet, crowding, maternalbehaviors, and predation were seen to have major roles in effectingphenotypes in plants and animals. Thus, the environment not onlyselected variations, it helped produce them. Since evo-devo postulatedthat changes in development cause evolution, and since developmentalplasticity played a role in development, then it became necessary tolook at changes in plasticity as being part of evolution. In the earlyyears of the 21st century, developmental plasticity was seen to playroles in evolutionary change (West-Eberhard 2003; Abouheif et al.2014; Suzuki & Nijhout 2006; Rajakumar et al. 2018; Levis &Pfennig 2020), and the notion of ‘plasticity-firstevolution’ (or ‘development-first evolution’)integrated data from numerous sources into a program where thephysiological ability to alter one’s phenotype due toenvironmental agents could become canalized and genetically fixed byselection. The mechanisms by which such plasticity-first evolution waseffected (unmasking and selection of cryptic genetic variants,stress-related inability of molecular chaperones to allow properfolding of mutant proteins, etc.) became a new research program.

A fifth discovery was the realization that one of the majorenvironmental agents effecting development were symbiotic microbes(McFall-Ngai 2002; Gilbert et al. 2012, 2015). The notion of theholobiont (i.e. an integrated composite organism composed of microbialand host eukaryotic species) organized much of the data to look at theroles of microbes in causing both the normal development of theorganism and variations of normal development (in disease andevolution) (Rosenberg & Zilber-Rosenberg 2016; see entrybiological individuals). For instance, in the mouse, the normal development of the immunesystem and the gut capillary network depends upon specific bacteriaobtained during birth. These bacteria induce the expression ofparticular genes in the eukaryotic cells, and the proteins made bythese genes influence cell fate (Hooper et al. 2001; McFall-Ngai etal. 2013). Signals for normal mammalian development, for instance, canbe provided by metabolites derived from food digested by themother’s microbiome. Such intergeneration developmentalsymbiosis appears to be needed for the maturation of the auditoryneurons in fetal mouse brains as well as for pancreatic celldevelopment (Kimura et al. 2020; Vuong et al. 2020). Symbioticmicrobes can also be provided by the starter set of microbes acquiredduring the movement through the birth canal. These microbes arecrucial in maturing the capillary network of the intestine, the immunesystem of the gut, and the neurons involved in coordinatingperistalsis (see Gilbert 2024). In other words, the bacteria can actas an embryonic cell, regulating gene expression in neighboring cells.Here, the eukaryotic organism needs and expects these bacteria to bepresent for normal development.

As in the other cases of developmental plasticity, the next step wasto see if changes in developmental symbionts could produce changes inevolution (see O’Malley 2015). It was shown that changes insymbionts could provide selectable variants for evolution (Zhang etal. 2019), and it could provide the basis for reproductive isolationthrough cytoplasmic incompatibility or mating preference (Brucker& Bordenstein 2013; Sharon et al. 2010). One of the mostinteresting possibilities, though, comes from the view that most, ifnot all, eukaryotic organisms are holobionts, and that symbionts opennew evolutionary trajectories. Symbiotic microbes, for instance, havelong been known to be responsible for the plant-digesting enzymes inthe stomachs of ruminants. Without cellulose-digesting bacteria, cowscannot digest grass or grain. Moreover, the microbes help create therumen after they colonize the digestive system at birth. Thus, thesemicrobes also induce the formation of the organ that houses them, andallows them to function (Gilbert 2020; Chiu & Gilbert 2020).Developmental symbiosis (sympoiesis) thus has opened evolutionarytrajectories for certain mammals. This is an example of bothdevelopmental symbiosis and niche construction. Niche constructiondepends on developmental plasticity (Laland et al. 2008).

These five new (or renewed) aspects of developmental biology haveseveral philosophical implications. Among others, they concernontological questions of what developing organisms are and how theyshould be explained from an integrated perspective of developmentalevolution.

4. Ontological challenges of developmental evolution

Many ontological debates on the relation between development andevolution focus on the organism as the central unit, which bothdevelops and evolves, in contrast to, for example, genes orpopulations. Several theories have tried to clarify the nature of thisorganismic unit: According to one influential view, the organism is aseries of integrated processes during a life cycle (Bonner 1965;Nicholson & Dupré 2018; Fusco 2019a; DiFrisco 2019), withcomplex and reciprocal relationships between the whole organism andits parts (Gilbert & Sarkar 2000; Esposito 2016; Peterson 2017).Central elements of this view are based on an organicist frameworkdeveloped by Kant, which states that in organisms “the parts,with respect to both form and being, are only possible through theirrelationship to the whole” and “that the parts bindthemselves mutually into the unity of a whole in such a way that theyare mutually cause and effect of one another” (Kant 1790/1793[1902: 373]; see also Lenoir 1982). Haraway (2008) highlights thelatter idea of reciprocal interaction between organisms’ partsby saying: “Reciprocal induction is the name of the game”(2008: 228) and it is “reciprocating complexity all the waydown” (2008: 42).

These ideas of life cycle integration and ubiquitous reciprocitysuggest a more processual and organism-centered ontologicalperspective on the organism. This view is becoming important instudies of evolution. For example, in order to understand how nervoussystems evolve one need to consider that the nervous system of thedeveloping organism has different functions than the adult nervoussystem and may be used to coordinate body construction as it develops(M. Levin 2019; Fields et al. 2020). That developing systems showexaptation and competition and that evolving systems show cooperationhas allowed Fields and Levin (2020) to suggest that developmental andevolutionary processes can be integrated on a scale-free level throughthe language of information processing and communication.

A second ontological consideration states that developing and evolvingorganisms are integrated collective individuals, so-called holobiontsor ‘meta-organisms’ (Zilber-Rosenberg & Rosenberg2008; Bosch & McFall-Ngai 2011; Gilbert, Sapp, & Tauber 2012;see also O’Malley 2017; Baedke et al. 2020a). Therefore, ourdiscussions about evolution must take into consideration that eachorganism is a consortium having numerous genomes, not just one, astraditionally assumed. Mathematical modeling of the evolution ofholobionts that take this diversity into consideration is justbeginning (Roughgarden et al. 2018; Osmanovic et al. 2018; Roughgarden2020). This new ontological framework states that symbiosis is thenorm; it is not peripheral. These symbionts can act at differentstages of the life cycle and are seen to scaffold development(Griesemer 2014; Chiu & Gilbert 2015; Minelli 2016). In‘scaffolding,’ each developmental stage is made possibleby entities and processes that catalyze these activities, which allowsnovel and evolutionary relevant processes to occur at lowerdifficulties and costs.

A third ontological point concerns the exact nature of the linkbetween development and evolution. Two approaches have been putforward: One draws on the idea that the biological entity that iscausally efficacious in both realms can only be found on the level ofintegrated collectives of symbiotic interactions. Following in thetradition of Leibnitz’ notion of compossibility as well asMargulis’ (1999) claim that we live on a ‘symbioticplanet,’ this view argues that symbiotic collectives are notonly essential units of development but also of evolution. This leadsto a view of evolution that is not centered on interspecies conflictand competition between individuals (Huxley 1888; Williams 1966;Dawkins 1976). Instead the entities that evolve are cooperativeco-developing collectives (Rosenberg & Zilber-Rosenberg 2016;Roughgarden et al. 2018; for discussion, see Peacock 2011;O’Malley 2014; Doolittle 2016; Suárez 2018).

A different ontological framework links development and evolutionthrough the entity of the acting organism (Nicholson 2014, 2018; Walsh2015; for discussion, see Pradeu 2016; Baedke 2019a; Baedke &Fábregas-Tejeda 2023). These approaches usually are lessrelated to symbiosis research than to studies on niche construction ormaternal effects. Here the organism (e.g., a beaver that builds a damor an earthworm that processes the soil) is constructed as aself-determined agent that through its behavior modulates theselection pressures acting on it (Odling-Smee et al. 2003; Uller &Helanterä 2019; see also section 6). Thus, so the argument goes,it can bias and direct population dynamics. The holobiont perspectiveand the niche construction perspective join together when oneappreciates that the microbes and the host form each other’senvironment. Here, the symbionts (as in the cattle rumen) are involvedin constructing their own niche within the host (i.e., the largesymbiont), and the host and microbes scaffold each other’sdevelopment and evolution (Laland et al. 2008; Chiu & Gilbert2015, 2020).

Whether or not we consider collectives or organismic individual agentsas the core entities partaking both in development and evolution,attempts to integrate the two realms have to show in each case that,in fact, it is the same unit that developsand evolves. Inother words, if we want to unify development and evolution through theunit of the biological individual (being the one entity that partakesin both) this unit needs to meet criteria of both physiological (e.g.,metabolic) and evolutionary individuality (see the entry onbiological individuals). Evolutionary individuals have been traditionally conceptualized asreproductive units with differential fitness and shared lineages(so-called ‘Darwinian individuals’; see Godfrey-Smith2009) or as units of selection (‘interactors’; see Hull1980). Unfortunately, both of these units do not always coincide(Godfrey-Smith 2013; Pradeu 2016). For example, some organisms(holobionts) form developing but no reproductive units, as theyinclude a multitude of lineages (e.g., microbial ones). Other possibleunits of selection (like genes or populations) are not identical withphysiological individuals. Thus, a physiological individual may notnecessarily be an evolutionary unit or vice versa.

This brings us to a fourth ontological point: developmentalplasticity, which is considered to bias or even guide evolutionarytrajectories (West-Eberhard 2003; Radersma et al. 2020). The conceptof plasticity states that development can be regulated in importantways by the environment. This rules out genetic determinism (but notnecessarily environmental determinism; see Waggoner and Uller 2015).In the original conception of phenotype production (i.e.,development), Wilhelm Johannsen (1909) had pointed out that thephenotype is the product of both the genotype and environmentalcircumstance, and Woltereck (1909; see also Sarkar 1999) argued thatwhat was inherited is the “Reaktionsnorm”, apotential to generate phenotypic variations in response toenvironmental agents. In line with this view, many embryology texts inthe late 1800s (e.g., Hertwig 1894) had promulgated the perspectivethat development demanded both the interactions between embryoniccells and the further interactions of those cells with the environment(see Nyhart 1995).

Despite this history, developmental plasticity was marginalized asgenetic explanations came to the fore in the mid-20th century (Sarkar1998; Keller 2002). Against this background, embryologists such asLewis Wolpert (1994) could ask whether an organism’s phenotypecould be computed if we had the total description of the egg. Due tothe above findings on organism-environment interaction (see section3), a different view emerged that more seriously considers theenvironmental-responsiveness and plasticity of the developingphenotype. This view includes a shift from externalist to internalistor constructionist understandings of the organism-environmentrelationship (Godfrey-Smith 1996). While the externalist view –the orthodox view in evolutionary theory – conceptualizesproperties of organisms as a result of their environments (i.e.natural selection targeting genetic programs), the internalist viewsees “one set of organic properties in terms of other internalor intrinsic properties of the organic system” (Godfrey-Smith1996: 30). According to these two accounts, organisms occupy anenvironment that covaries with them or that is largely independent oftheir variation. The above research in developmental evolutionsuggests a switch from an externalist to a constructionistperspective, in which the organism actively molds its internal statesand responds to and alters its external environment (see Laland et al.2014, 2015). In addition, in this framework the causal role of theenvironment also becomes more complex. It now includes the idea thatthe environment has active agency that can determine the phenotype.Rather than conceptualizing the environment as nothing but a passivefilter for evolution, in this view the environment plays a role inactuating the phenotype in addition to selecting it (Moczek 2015;Gilbert & Epel 2015).

5. Explanations of developmental evolution

Besides these discussions about the ontology of developing andevolving organisms, other central philosophical debates on theinterface between development and evolution have targeted the topic ofscientific explanation. This refers to the questions of what studiesof developmental evolution (should) explain and how they explain.

5.1 Mechanistic explanations of developmental evolution

Philosophers of science have long argued for the explanatory autonomyof biological explanations. Especially, they have criticizedunderstanding biological explanation as similar to law-based accountsof explanations in physics (see Lange 2007). In contrast, scholarshave argued that explanation in the biosciences often includesdescribing a mechanism that brings about a certain biologicalphenomenon (Bechtel & Richardson 1993; Craver 2007; Bechtel &Abrahamsen 2010; see also entrymechanisms in science). Especially evo-devo has been described as a paradigmatic mechanisticscience, which – against the ultimate-proximate distinction– seeks to identify developmental mechanisms that can explainevolutionary change in phenotypes (Gilbert 2003; Hall 2012). Thismechanistic approach is often flanked by mathematical models ofvarious developmental patterns, from changes in gene regulatorynetworks to growth patterns of organisms, and by historical narrativeson how organisms and species evolve (Jaeger & Sharpe 2014; Winther2015). However, besides the accepted centrality of mechanisticexplanation for developmental evolution, a much-debated topic concernswhat exactly a developmental mechanism is and how it functions inevolutionary explanation compared to standard explanations citingnatural selection.

Philosophers of biology (in the so-called new mechanistic philosophy)have conceptualized mechanistic explanations in biology as theconstruction of models of mechanisms that connect parts of systems,located on one level of organization, with behaviors of the wholesystem, usually located on a higher level of organization (Machamer etal. 2000; Craver 2007; Illari & Williamson 2012). In thisframework, mechanistic models relate different compositional levels oforganization, like genes and phenotypes or cells and tissues. Theseinter-level relations exist between causal capacities of parts of asystem and their organization and the capacities of a system as awhole. Such relations are established following a procedure ofdecomposition and localization (Bechtel & Richardson 1993; Craver2007; Menzies 2012). This conceptual framework to describe biologicalmechanisms and mechanistic explanation has been developed based oncase studies in molecular and cell biology. However, scholars havecast doubt on whether it is also useful to describe mechanisticexplanations in studies of development and developmentalevolution.

With respect to development, it has been argued, first, thatorganization plays a different role in mechanistic developmentalexplanations (Mc Manus 2012). In contrast to the above framework,which usually presupposes that levels of organization are simplythere, and thus it does not have to clarify how levels of organizationactually originate, the origin of levels and other forms oforganization (e.g., spatial axes) are specifically addressed inmechanistic developmental explanations. Second, philosophers haveargued that the relations between levels traced by developmentalmechanisms are not exhausted by the synchronic, constitutive relationsbetween parts and wholes, as some new mechanists suggest (Craver &Bechtel 2007). In contrast, developmental explanations trace changingdiachronic relationships between causal capacities of a system atdifferent levels of organization at different time intervals (Ylikoski2013; Baedke & Mc Manus 2018; Baedke 2020; see also Love &Hüttemann 2011). Third, it has been argued that explanations inevo-devo using developmental mechanisms face a challenge due to theheterogeneity of these mechanisms (Love 2018). When trying tointegrate two types of explanations of developmental mechanisms– explanations of highly conserved molecular genetic mechanisms,like gene regulatory networks, and explanations of cellular-physicalmechanisms, like cell migration – sometimes a tradeoff emerges.Rather than allowing a more complete explanation, integrating the twomechanisms may lead to a less general explanation, sincenon-phylogenetically conserved cellular-physical mechanisms yield lessgenerality in explanations. This tradeoff can introduce an explanatorybias to projects that seek to integrate development and evolution. Itcould lead researchers to favor the generality of explanations, whichcite highly conserved molecular genetic mechanisms and nocellular-physical mechanisms, over integrated explanations citing bothkinds of mechanisms.

With respect to the concept of mechanism in developmental evolution,Brigandt (2015) highlights that some mechanistic explanations inevo-devo – like those on how development biases evolution(Radersma et al. 2020), how novel variation arises throughdevelopmental plasticity (Pigliucci 2001, Gilbert 2006), and howorganisms generate heritable, adaptive phenotypic variation(evolvability; see Brown 2014) – significantly expand thestandard analysis of decomposition and localization by dynamicalmodels (see also Bechtel & Abrahamsen 2010; Brigandt 2013; Baedke2020). These models allow predicting the dynamics of developingsystems. After decomposing a system and identifying causalcontributions of parts or sub-systems, dynamical models help todemonstrate how these contributions operate together to bring about awhole system’s behavior. In this way, they answer howdevelopmental mechanisms create evolutionary relevant qualitativechanges in phenotypic properties, like robustness, phenotypicplasticity, and modularity, through underlying quantitative changes intheir component parts and activities. One classical example of thisare mathematical models on the robustness of spatial patterning andsegmentation inDrosophila. They provide quantitativeinformation about the interaction of underlying gene networkcomponents, including, for example, gene transcription rates and decayrates of gene products (von Dassow et al. 2000).

Other discussions on the interface between development and evolutionfocus on how to understand the commonly used notion of‘conserved mechanism’ (Love 2024) or the structure ofexplanations that address how developmental mechanisms evolve. Forexample, Calcott (2009, 2013) has argued that Mayr’s distinctioncharacterizes two kinds of explanation: developmental explanation thatanswers ‘How doindividuals workat atime?’ and evolutionary explanation that answers ‘Howdopopulations changeover time?’ However,there is a third kind, called ‘lineage explanation’.Lineage explanation differs from the above by answering ‘How doindividuals changeover time?’ As Rudy Raff,one of the pioneers of evo-devo, summarized this idea when comparinghis work to that of standard evolutionary biologists (Amundson 2005,p. 253), “they’re interested in species; we’reinterested in bodies.” Therefore, lineage explanation offer aseries of mechanistic models, which trace differences between thedevelopmental mechanisms of individuals that produce the relevantmorphological structures at different times. Over evolutionary time,these relations undergo small modifications, which ultimately bringabout novelties, like eyes, teeth, and feathers, in the whole system(Jernnall et al. 2000; Salazar-Ciudad & Jernvall 2010; Machado etal 2023). Thus, lineage explanation expands the standardsphilosophical framework from a single description of a mechanism intoa series of mechanistic models. Despite this expansion, however, thereremains the general challenge to combine and integrate mechanisticexplanations operating on the level of individuals with more classicpopulation-level approaches to evolution (Villegas 2024).

5.2 The explanatory power of developmentalist explanations of evolution

Besides these debates about the structure of biological mechanisms andtheir role in explaining developmental evolution, philosophers ofbiology and biologists have discussed, more generally, which virtuedevelopmental explanations (could) have for addressing evolutionaryphenomena. Related to this issue is the question how much – inthe sense of what kind of facts – natural selection alone canexplain (see the entryadaptationism). It is widely accepted that such explanations can address the generaldynamics of trait frequencies and survival (see Sober 1984). However,whether this also holds for addressing in more detail the developmentof particular traits of individuals is an unsettled issue. While someauthors claim that evolutionary explanations by natural selection canexplain why a particular individual has a certain trait rather thananother trait (Neander 1995; Forber 2005) others deny this (Sober1984; Stegmann 2010). What is more, it has been argued thatintegrating explanations of developmental phenomena, likedevelopmental bias, phenotypic plasticity, niche construction, andinclusive inheritance, to the explanatory framework of evolutionarytheory would lead to a “significantly expanded explanatorycapacity” of this theory (Pigliucci & Müller 2010: 12;Lala et al. 2024). However, while there is often agreement inevolutionary biology over the existence of these developmentalphenomena (Laland et al. 2014; Wray et al. 2014), their explanatoryrelevance is questioned. Against this background, scholars have begunanalyzing based on which criteria of explanatory power, likeprecision, proportionality, sensitivity, and idealization,developmentalist evolutionary explanations are better thanselectionist explanations. This includes identifying, which tradeoffsbetween explanatory standards (e.g., between precision and sensitivityor idealization) those accounts face that seek to integratedevelopmental and evolutionary explanations (Baedke et al. 2020b;Uller et al. 2020).

6. Organismal agency in development and evolution

Another long-standing debate that is especially reemerging in recentyears starts from the observations that organisms actively react toenvironmental factors, self-establish and -maintain their organizationdespite various changes, co-direct their plastic development, and thusbias and mediate their ecological interactions and evolutionarytrajectories. This discussion needs to be placed against the broaderhistory of concepts like agency, goal-directedness, purposiveness, andteleology in biology. In the 20^th century, debates onteleology (see the entryteleological notions in biology) drew on the relation between evolution and development especially todistinguish the teleological dimension of development fromnon-teleological evolutionary processes (Mayr 1961). Others introducednew conceptual frameworks, like teleonomy, which refer to onlyapparently purposeful systems (Pittendrigh 1958; see Dresow & Love2023). Such frameworks attempted to understand the phenomenon oforganismal agency as resulting from so-called ‘externalteleology’ (i.e. as a product of external selective forces andadaptation), not from organisms’ ‘internalteleology’ or ‘intrinsic purposiveness’, which wasdiscredited as a view that inevitably leads to vitalist speculations(Baedke & Fábregas-Tejeda 2023). Due to this, one findsrather few discussions on the purposeful organismic agent as astarting point to understand the teleological nature of developmentand evolution during this time (but see, e.g., Russell 1950; Piaget1976 [1978]).

In recent years, this situation has changed drastically through manyworks discussing organismal agency (e.g., Moreno & Mossio 2015;Walsh 2015, 2021; Riskin 2016; Okasha 2018; Corning et al. 2023;Mitchell 2023; Moczek & Sultan 2023; Fábregas-Tejeda et al.2024; Rupik 2024). This debate has particularly been stimulated by newdevelopmentalist accounts of evolution, especially in nicheconstruction theory, studies of plasticity-led evolution, and(eco-)evo-devo. These views, first, induced a shift away from pastattempts to explain away organismal agency as a mere consequence ofadaptation. Second, they reopened the door to again explore differentframeworks that allow conceptualizing organisms’ intrinsicpurposiveness (e.g., Aaby & Desmond 2021; Walsh 2021; Sultan etal. 2022; Patten et al. 2023; Nuño de la Rosa 2023; Jaeger2024). These accounts address (again) questions about the ontologicalstatus of organisms and the epistemic role agency and teleology can(or should) play in development and evolution: For example, how wouldadopting a view of organisms’ internal teleology enhance ourunderstanding and explanations of developmental and evolutionaryprocesses? How should biologists conceptualize the apparentpurposiveness of organisms’ activities? Through concepts likeorganization, autonomy, and control or through goal-directedness or bydrawing on ecological ‘affordance’ frameworks (Moreno& Mossio 2015; Walsh 2015; Babcock & McShea 2024; for otherolder framework, see Fábregas-Tejeda 2024). Further issuesconcern whether the scope of agency is restricted to goal-directedbehaviors or whether it also can be ascribed to all or only specific(e.g. plastic) developmental processes (Sultan et al. 2022; Nahas2024; Walsh & Sultan 2024). And: What evolutionary consequences,if any, result from organisms’ agential activities? Addressingthese questions will determine which role organismal agency will playin future studies at the interface between development andevolution.

This path forward will also depend on whether ontological or epistemicviews of agency will be adopted in biology. In the current livelydebate we still see a wide spectrum of positions. On the one side ofthis spectrum, we find classical Kantian views according to whichagency is merely a heuristic tool (or epistemic framework) forbiologists to temporarily deal with the intricacies of developmentaland evolutionary phenomena until mechanistic research catches up (Kant1790/1793 [1902]; see Desmond & Huneman 2020). On the other sidewe find ontological or naturalizing views arguing that agency andpurposiveness in developmental evolution can only be understood as acapacity that fundamentally belongs to organisms (Walsh 2015).

7. Anthropological and ethical dimensions of developmental evolution

Besides the above debates about the relationship between evolution anddevelopment and about the role agency could play in linking the two,there are other, more general debates about anthropological andethical issues that concern developmental evolution. They emerge fromtwo developments: one the one hand, research on developmentalevolution has given the organism concept (and organismal agency) a newrelevance in biology; and this concept has often been used as abiological counterpart to concepts of societal relevance, like person,individual, and body. On the other hand, many of the empiricaldevelopments described in Section 3 went along with biomedicaladvances and debates in postgenomic fields like epigenetics,proteomics, exposomics, and microbiome research. These two trends ledto at least four anthropological and ethical discussions about how weconceptualize developing and evolving humans, their life, body andhealth, as well as how we assign responsibilities for healthcareinterventions:

First, new findings in the plasticity of developing organism, theirinterconnectedness, and modes of transgenerational transmission ofinformation have affected scientific and public understandings of whathumans are. For example, if humans are conceptualized as holobionts– as collectives of co-developing and co-evolving organisms– this also means that development is a matter ofco-construction, of interactions between species. It means, as Haraway(2016) has phrased it, that we – as humans – veryliterally ‘become with others.’ In this context,sympoiesis (developmental symbiosis) means that development isco-development. Against this background, John Dupré and MaureenO’Malley (2009) see living entities as interactive collections:“Life, we claim, is typically found at the collaborativeintersections of many lineages, and we even suggest that collaborationshould be seen as a central characteristic of living matter”.Due to this interrelatedness, Bapteste et al. (2021) and Gilbert(2021) suggest that microbiome research induces ade-anthropocentrification of humans’ perception of the world.These views are in line with Hans Jonas’ (1966, 1984) olderargument that humans maintain their humanity through a deep connectionwith nature, particularly through their symbiotic interconnectednesswith other living beings. As human expansion threatens the naturalworld and existence of other beings, Jonas emphasized the importanceof caring for all of nature’s future. He viewed this as a‘solidarity of interest’ with the organic world.

Second, biological and biophilosophical debates on developmentalevolution and organisms’ plasticity and environmentalresponsiveness have informed debates on what the human body is. Forexample, Jörg Niewöhner (2011) states that a new concept ofthe human body is currently emerging in modern biology, the so-called‘embedded body’. According to this view, the human body isno longer a machine-like unit, which is genetically programmed,neurally controlled and bounded by the skin, but an open, dynamic, and‘attentive’ unit which cannot be grasped in isolation fromits material and social environment (see also Baedke 2017; Frost2020). Additionally, the body is embedded into different time scalesranging from its evolutionary and transgenerational to ontogeneticpast, which permanently constitute its present. Others have argued,against the standard human birth narrative and Aristotle, who definedthe temporal boundaries of individuals at birth and death, that birthin humans is not the creation of a new individual. Instead, birthshould be understood as the origin of a new multi-species collective(Gilbert 2014; Chiu & Gilbert 2015).

Another anthropological issue arising from recent research in evo-devoand epigenetics is, third, the question how to define normality andhealth in humans (see Baedke 2019b). There is evidence that bacteriaare needed for our normal cognitive and social development. Forexample, germ-free mice are asocial and have autistic-like behavior(Desbonnet et al. 2014) and this behavior can be replicated byimplanting the microbiome from autistic patients (but not controlpatients) into germ-free mice (Sharon et al. 2019). Such cases suggestthat biological normality is not an intrinsic property to organismsbut emerges through interconnections with other organisms and theenvironment. In addition, our understanding of health is increasinglychallenged. This especially refers to the view that describes humanhealth as freedom and autonomy from external interference. It usuallysees bacteria as deviations from the norm and parasitism aspathological, because it threatens and contaminates the purity of theindividual’s energy pattern. Instead, in a (more processual)holobiontic framework, microbes are needed for normal development andare thought to prevent the development of certain diseases (Blaser2014; Bello et al. 2018; Kirjavainen et al. 2019). In addition,certain entities, bacteria or viruses, previously thought to beharmful, are now increasingly considered to be ‘good’ or‘healthy’ collaborators, not ‘bad intruders’(Tauber 2008; Dupré & Guttinger 2016; Sariola & Gilbert2020). In more general term, this means that since microbiota areincreasingly recognized as important components that stabilize normaldevelopment and co-evolve with humans, they therefore carry traitscrucial for humans’ fitness, i.e., health. This new perspectivecould lead to radical changes in personalized surveillance andtreatment of disease, and, more generally, to new strategies in policymaking, which replace the idea of preserving the autonomous individualfrom contamination by the idea of maintaining (equilibrium states of)collectives of co-developing and co-evolving individuals. This newperspective also highlights the question of how organisms evolve suchthat they can distinguish those microbes most likely to be pathogenicfrom those that are expected to become mutualistic symbionts (Tauber1994; Pradeu and Cooper 2012).

Finally, on a more ethical dimension, these findings abouthumans’ openness to their environments and to one another hasled, first, to discussions about who takes responsibilities forhumans’ health states and interventions (e.g., on epigenomes andmicrobiomes) on intra- and transgenerational timescales (Gluckman etal. 2009; Dupras & Ravitsky 2016). If plastic development canshape evolution, who is responsible for developmental outcomes andevolutionary trajectories in humans? Should the individual being asthe central heath care agent, which is ‘freed from the chains ofits genes’ (Pickersgill et al. 2013), take this responsibility?Or should collectives, such as national states or internationalbodies, be responsible for levels of toxins in the environment as wellas for the food individuals eat and stress they are exposed to(Hedlund 2012)? The latter account of responsibility aims to preventoveremphasizing the causal role of mothers as the most central publichealth care agents who should be held accountable (and guilty) iftheir children or later generations become sick (Richardson et al.2014).

8. Concluding Remarks

The relationship between evolution and development has been a longdebated topic in the history of biology and philosophy of biology.This entry has sampled a small portion of work relevant to theconceptual, ontological, epistemological, anthropological and ethicalreflections on this relationship. Besides the issues discussed here,philosophers of biology and biologists have discussed howdevelopmental and phylogenetic approaches to homology can beintegrated (Amundson 2005; Wagner 2014; DiFrisco 2019, 2023; DiFrisco& Jaeger 2021; McKenna et al. 2021; and the entry ondevelopmental biology), what challenges interdisciplinary collaboration faces when studyingcomplex phenomena of developmental evolution (Love 2024), what modelorganisms (Love 2009; Lloyd et al. 2012; Minelli & Baedke 2014;Zuk et al. 2014; the entry ondevelopmental biology) and representational tools scientists (should) use for studyingrelationships between evolution and development (e.g., normal plates,cell fate maps, epigenetic landscapes), and what the epistemic andheuristic roles of these tools in scientific practice are (see Haraway1976; Gilbert 1991; Hopwood 2007; Love 2010; Baedke 2013; Baedke &Schöttler 2017; Nicoglou 2018).

At the moment, philosophical debates about the appropriate conceptualand explanatory approach to combine or integrate developmental andevolutionary processes have not reached a consensus. Indeed, onerecent attempt at integration recognizes a large diversity ofapproaches. Under that banner of “welcome pluralism,” Lalaand colleagues (2024) admit that the various theoretical andconceptual frameworks are highly heterogeneous, and they find this amark of a “healthy science.” At the same time, anypluralism of such kind needs to be able to draw boundaries andestablish clear criteria when and why to adopt which exact epistemicstandards, models, and explanatory or conceptual frameworks (ratherthan others). Thus, an important aim for future philosophical researchis to understand the obstacles for the stabilization andsolidification of these frameworks, to identify their explanatoryvirtues and limitations, as well as to call attention to their effectson how we understand humans and human health.

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Other Internet Resources

• Evolutionary Developmental Biology, a reference guide for biological, historical, and philosophicaltopics on evo-devo, edited by Laura Nuño de la Rosa and Gerd B.Müller
• The Embryo Project Encyclopedia, with entries on the history of research on developmentalevolution
• Evo-devo, a general introduction to evolutionary developmental biology, at theUnderstanding Evolution website (U. California/Berkeley)
• Evo-devo resources, at the Understanding Evolution website (U. California/Berkeley)
• New York Times video with Sean Carroll on evo-devo
• Evo-devo song, by Tim Blais

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