Interdisciplinary Lessons For The Teaching of Biology From The Practice of Evo-Devo

Sci & Educ (2013) 22:255-278 OI 10.1007/11191-011-9351-6 Interdisciplinary Lessons for the Teaching of Biology from the Practice of Evo-Devo Alan C. Love Published online: § April 2011 © Springer Science+Business Media B.V. 2011 Abstract Evolutionary developmental biology (Evo-devo) is a vibrant area of contemporary life science that should be (and is) increasingly incorporated into teaching curricula. Although the inclusion of this content is important for biological pedagogy at multiple levels of instruction, there are also philosophical lessons that can be drawn from the scientific practices found in Evo-devo. One feature of particular significance is the interdisciplinary nature of Evo-devo investigations and their resulting explanations. Instead of a single disciplinary approach being the most explanatory or fundamental, different methodologies from biological disciplines must be synthesized to generate empirically adequate explanations. Thus, Evo-devo points toward a non-reductionist epistemology in biology. I review three areas where these synthetic efforts become manifest as a result of Evo- devo's practices (form versus function reasoning styles; problem-structured investigations; idealizations related to studying model organisms), and then sketch some possible applications to teaching biology. These philosophical considerations provide resources for life science educators to address (and challenge) key aspects of the National Science Education Standards and Benchmarks for Scientific Literacy. 1 From Content to Practice, to Pedagogy Geophilomorph centipedes always have an odd number of leg bearing body segments. The total mumber can range from 27 to 191 though the predominant range is between 39 and 85 in the ~1,000 species (Minelli 2009, Chap. 3; Arthur 1999, 2011, Chap. 13). Is it selectively advantageous to have an odd rather than even number of body segments (or vice versa)? Maybe there is a relevant fitness difference, but it seems unlikely. A decreased number of segments might favor enhanced locomotory capacities, but then lower numbers of segments (rather than odd or even counts) would be favored. Evidence from developmental genetics suggests that specific developmental mechanisms establish the pattern. A.C, Love (3) Department of Philosophy, Minnesota Center for Philosophy of Science, University of Minnesota, 831 Heller Hall, 271 19th Ave. S., Minneapolis, MN 55455-0310, USA e-mail: aclave@umn.eda © Springer256 A.C. Love Gene expression occurring during the formation of segments in ontogeny begins with a periodicity of two segments at a time; subsequently, a new periodicity of one segment at a time emerges, leading to an odd final count for the leg-bearing body segments (Chipman et al. 2004), But there is nothing special about being odd. Other segmented invertebrates only have an even number of body segments—leeches always have 32. It appears that natural selection does not explain body segment number in geophilomorph centipedes, or other segmented animals (Davis and Patel 1999; Minelli and Fusco 2004). Developmental Processes that generate this phenotypic bias must be invoked to comprehend the evolutionary pattem of variation. ‘The centipede example is one of many available from the content of evolutionary developmental biology (Evo-devo) that can be deployed in the classroom. For serially repeated elements, other options include the consistent pattern of seven cervical vertebrae across diverse mammals, except for sloths and manatees (Van Sittertet al. 2010; Narita and Kuratani 2005; Hautier et al. 2010), or angiosperm phyllotaxis (leaf-arrangement) and the Fibonacci number series (Cooke 2006). These and other examples abound in the abundant and accessible presentations of Evo-devo (e.g., Shubin 2008; Arthur 2006, 2011; Carroll 2005; Kirschner and Gerhart 2005; Minelli 2009), including documentary television spe- cials (http://www. pbs.orghwgbh/nova/evolution/darwin-never-knew.html). From Hox genes to the origin of turtle shells, Evo-devo is an exciting area of research that should be incorporated into biological pedagogy. It is not just that development evolves—develop- ‘ment also structures the evolution of organismal traits. Therefore, understanding development is essential to understanding evolution. Much of the framing around Evo-devo research is the purported inadequacies of standard neo-Darwinian evolutionary theory (or the “Modern Synthesis”). ‘The Modem Synthesis established much of the foundation for how evolutionary biology has been discussed and taught for the past sixty years. However, despite the monikers of ‘Modern’ and “Synthesis,” it was incomplete (Carroll 2005, 7). ‘The Modern Synthesis of 1940 was not so much wrong as it was incomplete. ...On the question of how the altered genotype caused an altered phenotype, the Modem Synthesis was silent (Kirschner and Gerhart 2005, 29). ‘This orientation is evident in my claim that natural selection is not the explanation for leg- bearing segment numbers, and it is observable more generally (Miiller 2007; Pigliucei 2007: Pigliueci and Miller 2010; Laubichler 2010; Carroll 2008). This is also related to claims that teaching evolution from an Evo-devo perspective is more compelling than the traditional emphasis on population genetics and microevolution Gilbert 2003): Millions of biology students have been taught the view (rom population genetics) that ‘evolution is change in gene frequencies’ ...This view forces the explanation toward mathematics and abstract, descriptions of genes, and away from butterflies and zebras ...The evolution of form is the main drama of lifes story, both as found inthe fossil record and in the diversity of living species. So, let's teach that story. Instead of ‘change in gene frequencies,’ let's uy ‘evolution of form is change in development’ (Carll 2005, 294) ‘Thus, itis not just that standard neo-Darwinian theory was empirically inadequate but also that it was an obstacle to communicating the core phenomenon of evolution to students. Educators confronted with the myriad of empirical content found in Evo-devo face a dilemma. On the one hand, the framing of Evo-devo as a rival or successor to neo- Danwinian evolutionary theory sets up an antagonism across different areas of biological research. This is especially dangerous given the combustible atmosphere surrounding the teaching of evolution, and problematic interpretations of the nature and significance of D SpringerInterdisciplinary Lessons for the Teaching of Biology 257 scientific controversy (Sarkar 2007; Scott and Branch 2003). On the other hand, incor- porating recent scientific developments into the biological curriculum is a herculean task, especially because the life sciences are composed of diverse and heterogeneous disciplines. While students need to be exposed to new and significant discoveries across the spectrum of biological inquiry,’ Evo-devo is only one of many areas that must (seemingly) be stuffed into the crowded curriculum. It appears analogous to the case of Alice in Through the Looking Glass, when the Red Queen says, “it takes all the running you can do, to keep in the same place.” If so, the situation is not just difficult but disheartening because most agree that science education needs to move beyond the status quo.” And the Red Queen also declared that, “if you want to get somewhere else, you must run at least twice as Fast.” One way to escape from (or at least circumvent) this dilemma is to shift our attention from the content of Evo-devo to its practices. Instead of chasing the newest theoretical or empirical content, and evaluating whether it is or isn’t a challenge to evolutionary theory, wwe can ask if there are lessons to be learned from how Evo-devo biologists go about their scientific tasks—how they practice science, Roughly. a scientific practice picks out a way of acting or proceeding in the empirical investigation of the natural world. Some of these practices are material and involve the physical arrangement of laboratory or museum space, experimental apparatuses, and animal husbandry. Material practices have implica- tions for what kinds of information can be gathered about natural systems and to what degree findings can be extrapolated (Burian 1993; Love 2009b). Other practices are conceptual and involve the use of reasoning tools (e.g., statistical) or specific concepts (eg., heterochrony). Concepts are often viewed in terms of their content (what they represent or refer to); ‘heterochrony’ represents changes in the initiation, rate, or completion of events during development. But concepts also can be considered from the vantage point of practice. What is the role of the concept ‘heterochrony’ in Evo-devo research? Is it used to categorize phenomena or explain phenomena, or does it need explaining? Does it guide research or influence the choice of methods? These are questions about the practices involved in using a specific concept, which have an impact on theoretical commitments in biology such as how empirical content is interpreted or changes over time (Brigandt 2010b, this issue).” One distinctive feature of the scientific practices found in Evo-devo is their interdisciplinarity (Love 2003, 2005, 20082, 20103). Interdisciplinary research involves the coordination or synthesis of methods, concepts, and explanatory resources (and standards) from different scientific approaches.’ Although Evo-devo is often characterized as an integration of embryology with evolutionary theory, it involves a larger synthesis of diverse approaches such as genomics, morphology, paleontology, and systematics (Hall ‘ence content increases and changes, and a teacher's understanding... must keep pace" (National Sclence Education Standards 996 (NSES}, 57). > “Reforming science education requires substantive changes in how science is taught. ...The vision of science education described by the standards requires changes throughout the entire system” (NSES, 5,28). ® The distinction between conceptual and material practices is congruent withthe learning of science as an active process: “the tom “active process” implies physical and mental ativity. Hands-on activities are not tenough—students also must have “minds-on” experiences” (NSES, 20) “ Inteedisciplinary and mulidiseiplinary can be distinguished so chat the former describes research involving the formation of a new discipline whereas the latter only requires existing disciplines to work together transiently. The present use of “interisciplinary” does not invoke this distinetion because the saliont feature is that multiple approaches must be synthesized in order to produce an adequate explanation (whether transiently oF not) © Springer258 A.C. Love 1999). Interdisciplinarity is of philosophical concen because it points toward the possibility of a non-reductionist epistemology in biology (Brigandt and Love 2008; Brigandt 2010a, this issue; Love 20082), especially as a counterweight to predominantly reductionist. methods (Robert 2004). The “fundamentality” of one particular perspective producing the most empirically adequate explanations is rejected, often with an explicit dismissal of explanations from one approach—“to state that a genetic mutation led to a favored character, which, in tum, was selected is utterly uninformative in explaining innovation” (Wagner et al. 2000, 822-823)—and a concomitant embrace of the kinds of approaches. that must be synthesized—“paleontologists. comparative anatomists, developmental biologists, and molecular geneticists are all contributing data aimed at clarifying the genetic basis for novel structures like heads, tails, and limbs” (Freeman 2002, 475-476). How do biologists know what is needed to explain particular phenomena? How do scientists know when they have the appropriate mixture of biological disciplines? Increasing collaboration across different sciences makes these philosophical questions more urgent and germane to educating the next generation of biologists, as well as the wider public. In this paper I review three areas where these synthetic, interdisciplinary efforts are manifest in Evo-devos practices: form versus function reasoning styles; problem-structured investigations; and, idealizations derived from studying model organisms. T draw a broad ‘lesson’ from each area: (I) reasoning in biological science is not homogeneous; (II) problems structure and guide biological research rather than hypothesis testing or theory confirmation; and, (III) different disciplinary approaches use reasoning strategies that simultaneously reveal some features and systematically overlook others. Then I sketch some possible applications for teaching biology, with a special emphasis on the National Science Education Standards (1996) and Benchmarks for Scientific Literacy (2009). Although this doesn’t exhaust the philosophical possibilities offered by Evo-devo, it highlights several relevant elements for pedagogy, such as the need for natural phenomena to be explored with different methods. An emphasis on practice is in accord with teaching science as inquiry.’ It is not just what Evo-devo says about living systems that needs to be communicated but how biologists go about investigating and explaining. Although re- contextualization is required to move these lessons into the classroom, the philosophical considerations distilled from these interdisciplinary practices provide novel resources for life science edueators—and challenge some cherished assumptions. By implication, they offer the prospect of going beyond running in place to maintain the status quo and reject the assumption that one must run twice as fast to get anywhere new. 2 Reasoning Styles: Synthesizing Form and Function Scientists exhibit different characteristic ways of investigating natural phenomena or “styles of reasoning” (Crombie 1994). One marker of divergent reasoning styles in biology is the distinction between form and function, Researchers whose style emphasizes “form” © Bvo-devo research is labeled diversely (eg, evolution of development, developmental evolution, oF rmicroevolution of development) but I will use "Evo-devo” for expository simplicity. (For further discussion, see Brigandt and Love 2010.) © 4 multifaceted activity that involves making observations; posing questions; examining books and other sources of information to see what is already known; planning investigations; reviewing what is already {noun in light of the student's experimental evidence: using tools to gather, analyze, and interpret daa: proposing answers, explanations, and predictions; and communicating the results. Inquiry requires use of assumptions, use ofentical and logieal thinking, and consideration of alternative explanations” (NSES, 23). D SpringerInterdisciplinary Lessons for the Teaching of Biology 259 scrutinize the composition and arrangement, shape or appearance of organic materials, whereas researchers whose style stresses “function” examine activities at various levels of organization performed or displayed by organisms (Bock and von Wahlert 1965). No biologist thinks form or function can be ignored, but stylistic preferences can shape the relations between them: “Is function the mechanical result of form, or is form merely the manifestation of function or activity?” (Russell 1982/1916, ix), Darwin's primary reasoning style was functional (Love, in press). The activities performed or displayed by organisms, such as reproduction or foraging, undergird his ideas about a struggle for existence and differential fitness effects of variation in populations. Adaptations are traits suited for doing something (functioning). Thus, a kind of functional reasoning motivates the centrality of natural selection as an explanation for the origin and maintenance of adaptive traits in organisms (Depew and Weber 1996). It meshes with a version of naturalized teleology that accounts for the presence of an organismal trait by appeal to its purposes (Lennox 1993; Reiss 2009; Walsh 2008): “modifications are accumulated by natural selection for the good of the being” (Darwin 1964/1859, 85-86). Many of Darwin's contemporaries, both supporters and detractors of his theory of evolution, utilized a form reasoning style. Richard Owen attempted to blend function and. structure (form), but came to see the latter as preeminent (Amundson 2005, Chap. 4, 2007; Rupke 2009, Chap. 4), Functional reasoning was unable to account for the unity of structural composition across vertebrate skeletons adapted to wildly different functions (Owen 2007/1849), ‘T. H. Huxley shared Owen’s form-orientation, in part due to the common influence of transcendental morphology from continental Europe: “adaptation ‘was no big deal... Huxley was ...still pushing homologies, and form over function” (Ruse 2003, 140), His aim in the early 1850s was to show that different morphological features of mollusks were modifications of a common structural plan: “that the squid’s arm was a homologue of the snail’s foot” (Desmond 1997, 174). “Adaptation was, if anything, a hindrance to the morphologist’s aim of discerning relationships. Huxley accepted adaptation, but he was fundamentally indifferent to it” (Ruse 2003, 140). These reasoning style differences between form and function remain salient today, especially in and around Evo-devo, They help explain why researchers adopt material and conceptual practices. For example, comparative morphology and embryology exemplify form-oriented reasoning (‘compositional’ biology) with practices that tack the organization, structure, and transformation of organic parts on developmental and evolutionary time scales (Winther 2006). These reasoning style differences also illuminate why continuing. difficulties remain in synthesizing disciplinary approaches in biology (Laubichler and Maienschein 2009). Many of the communities within Evo-devo’s interdisciplinary nexus (eg., embryology and developmental biology, morphology, and systematics) favor a form- oriented reasoning style, in part because of a comparative methodology. Proponents of twentith-century Evo Devo emphasized the need to study form as a corrective to what they perceived to be an over-eliance on functional (fitness) or optimality (adaptation) type explanations in evolutionary biology ...The emphasis within these kinds of developmental explanations hhas thus been on form ~ or rather the (conserved) structure of the developmental system (Laub ichler 2009, 11, 37), Arguably, this is an unresolved fault line between neo-Darwinism and Evo-devo (Amundson 2005, Chap. 11). Those who foreground organismal performance relative to environmental conditions (function) are at odds with ‘structuralists’ (form) who keep development at the center of evolutionary studies. The former explain evolutionary change from one adult phenotype to another via population processes such as natural selection, whereas the latter explain evolutionary change from one ontogeny to another via © Springer260 A.C. Love developmental processes such as morphogenesis. There may be no way to reconcile the two styles of reasoning into a single perspective. Thus, one of the manifestations of interdisciplinarity in and around Evo-devo is the intersection (and sometimes clash) of form and function reasoning styles. A particularly poignant tension between form and function in Evo-devo research is found in the contested concept of functional homology (Love 2007). In molecular developmental genetics, a core approach in Evo-devo, the conserved role of homeobox. gene expression during embryogenesis that establishes the axial patterns of animals is sometimes referred to as functionally homologous over and above the relation of structural homology that obtains between DNA sequences (Manak and Scott 1994). Other Evo-devo researchers complain that, from the perspective of comparative morphology and systematics, “functional homology’ is a contradiction (Abouheif et al. 1997). Homologous structures are the same by virtue of descent from a common ancestor, regardless of what functions these structures are involved in, whereas analogous structures are similar by virtue of selection favoring comparable functional outcomes, regardless of common descent. But etholo; 's also describe behaviors (functions) as homologous (Wenzel 1992). ‘The standpoint of scientific practice and form versus function reasoning styles suggest that the tension in the meanings for “function” and “homology” is related to the diverse explanatory ends in different areas of research (Amundson and Lauder 1994; Brigandt 2003. this issue). One of the most contentious areas of debate in philosophy of biology is function ascription (Allen et al. 1997; Ariew et al. 2002; Lewens 2004), Through attention to the heterogeneity of concept use in actual scientific practice, four separate meanings of ‘function’ in biology can be distinguished (Wouters 2003, 2005): activity (what something does), causal role (contribution to a capacity), fitness advantage or viability (value of having something), and selected effect or etiology (origination and maintenance via natural selection). Philosophical debate has raged about which sense (if any) is most appropriate for biological reasoning or most general in scope. But this kind of debate is not new and pre-Darwinian investigators of physiology drew a distinction between ‘use’ and ‘activity’ (Furley and Wilkie 1984, 58-69). The functional use of a feature represents ‘what itis for,’ whereas the functional activity of a feature picks out ‘what it does’ (or how it works). Multiple a multiple uses; the terms do not have the same extensions. They also are not equivalent epistemologically because we may know what something does but not what it is for and, alternatively, what something is for but not how it accomplishes this. This physiological distinction sheds light on the modern philosophical debate about function (Table 1). Use-function can correspond to causal role, viability, or selected effect, functions, but activity-function closely maps onto ‘mere’ activity and does not invoke a specific variety of functional contribution. As a result, a non-contradictory combination of function and homology can be isolated: homology of function refers to the same activity- function in different animals under every variety of form and use-function. Homologies between functions—construed as activities—obtain even as structures (form) related to an activity change, and despite different uses the activity is involved in. In the context of Evo- devo, we can observe this distinction in careful discussions of regulatory gene function. tivities might underlie a particular use, and one activity may be in the service of When studying the molecular evolution of regulatory genes, their biochemical and developmental function must be considered separately. The biochemical function of PAX-6 and eyeless are as ‘general transcription Tactors (which bind and uetivate downstream genes), but their developmental function is their specific involvement in eye morphogenesis (Abouheif 1997, 407). D SpringerInterdisciplinary Lessons for the Teaching of Biology 261 ‘Table 1A classification of four different interpretations of function in contemporary biology Senses of function Use (what itis for) Activity (what it is) according to the use versus Selected effect x activity distinetion Viability x Causal role x “Mere? activity x Modified from Love (2007) The biochemical function is the activity-function and the developmental function is the use-function, which differ in their evolutionary trajectories. Biochemical activity-fuunctions of regulatory genes are often conserved (1., homologous), while being available for co- option to make use-function contributions to distinct developmental processes. The same genes are evolutionarily stable in terms of activity-function and evolutionarily Tabile in terms of use-function.” This account of activity-function homology demonstrates how conflicts between form and function reasoning styles that emerge in interdisciplinary contexts might be resolvable by attending to the conceptual practices of different biological sub-disciplines. If we shochor these practices into a unified account, we risk misunderstanding how biology works. Biologists classify entities in terms of different senses of function and form. Homology is a form-oriented concept that describes a relation of sameness between structures independent of function (e.g., tetrapod forelimbs). Analogy is a function-oriented concept that describes a relation of similarity between structures due to performance constraints (e.g., body shape in sharks and dolphins). Biologists prioritize one reasoning, style over another differently depending on their explanatory goals. Many times these differences only become manifest in interdisciplinary contexts where they are set side by side, sometimes revealing that traits individuated using structural and functional criteria are not translationally equivalent (Wagner 2001); e-g., the traits might not exhibit coin- cident spatial boundaries and therefore cannot be utilized interchangeably (cf. Wimsatt 2007, Chap. 9). An interdisciplinary lesson for pedagogy can be recognized: reasoning in biological science should not be treated homogeneously. There are differences between form and function reasoning styles, and there are differences in how to understand the meaning of function itself. The heterogeneity results from two distinct epistemic situations. First, because there are form-oriented or structure aspects to evolutionary phenomena (e. homology), adaptation and natural selection (functional reasoning style) do not exhaust evolutionary explanation. In this case it is possible that the heterogeneity can be reconciled since the different styles of explanation concern different aspects of biological phenomena. Second, heterogeneity can be the consequence of different scientific aims; form-oriented and function-oriented reasoning styles can approach the same phenomena differently. The example of functional homology displays this vividly since function-oriented reasoning in terms of use-function cannot be conjoined to homology without contradiction, If we demand a single, homogeneous perspective, then we do violence to biology’s practices. Therefore we should teach the heterogeneity of reasoning in biology. Some have argued that assessments of homology ultimately rely on prior ascriptions of adaptive (use) 7" situation is even more complicated because its the parts of genes that correspond to different activity and use functions In homeobox genes, an open reading frame contains at least two “component hom Jogues": the DNA binding mf and a sequence involved in the relevant protein-protein interaction fora development process. There is no single activty-functin forthe entire open reading fame, which makes ‘se-function equivalence (homology) claims for homeobox genes ambiguous (Lynch und Wagner 2010). © Springer262 A.C. Love function (ie., form reasoning is subordinated to functional reasoning): “All significant biological kinds—including homologies—are (implicitly or explicitly) to be characterized in terms of selected-effect (SE) functions” (Rosenberg and Neander 2009, 307). This viewpoint does not correspond to biological practice, both with respect to homology and. function (Griffiths 2006; Amundson and Lauder 1994; Love 2007, in press; Wouters 2005). In addition to the different senses of function noted above, homology assessments are often made without any knowledge of a trait's SE function, both in the paleontological record, where this information is often absent, and in the genome, where the inference is made on the basis of sequence (form) homology. Biologists avoid using SE functions to individuate traits in order to test hypotheses about the operation of natural selection on those traits (thereby avoiding circularity of reasoning). And further individuation of homologues is accomplished by appeal to form features. For example, the second cervical vertebra in mammals is distinguished by its odontoid process (a shape feature), not any adaptive effects. Rosenberg and Neander (2009) claim that “homology and function ...are notions shot through with Darwinian presuppositions about ancestry and adaptation,” (309) but this assumes that reasoning in biology is homogeneous. The utilization of homology and function in pre-Darwinian frameworks, the stability of trait individuation across the theoretical transition to evolution (Griffiths 2007), the recognition by ancient physiologists, that senses of function must be differentiated, and the existence of four meanings of function in contemporary biology (Wouters 2003, 2005) all speak against this claim. Biology is composed of multiple perspectives that correspond to diverse aspects of natural phenomena and distinct explanatory aims. Sometimes function is preeminent (when explaining adaptation); sometimes form is preeminent (when explaining homology). Instead of a general asymmetry between form and function, these divergent reasoning styles manifest local asymmetries depending on the perspectives from which trait cate- gorization operates in different areas of biology. Local asymmetries become salient in interdisciplinary research where their potential incompatibility (due to distinct investigative aims) is palpable. The first interdisciplinary lesson from the reasoning practices of Evo-devo is that we must teach the heterogeneity of reasoning in biology (see also Bri- gandt, this issue). 3 Interdisciplinary Explanation: Answering Many, Different Questions Interdisciplinary explanations are sought for complex scientific problems that cannot be adequately addressed by one discipline (Mitchell 2003): “many features of nature are directly influenced by multiple mechanisms that fall into disparate scientific disciplines” (McPeek 2006, $1). Evolutionary theory is constituted by these kinds of problems, including speciation, evolvability, and adaptation (Love 2010a). One central problem for Evo-devo is how new features arise in the history of life—the problem of evolutionary novelty (Love 2006, 2008a; Brigandt and Love 2010). For example, how do bird feathers evolve from scale-like precursors that are dramatically different? Evo-devo researchers claim that adequate explanations for the origin of novelties are lacking, and only a synthesis that offers interdisciplinary explanations can provide them. The subtitle for a symposium on the origin of feathers makes this explicit: “A Problem Demanding Inter- disciplinary Communication” (Maderson and Homberger 2000). And researchers regularly use the vocabulary of problems: “The origination of morphological structures, body plans, and forms should be regarded as a problem distinct from that of the variation and D SpringerInterdisciplinary Lessons for the Teaching of Biology 263, diversification of such entities (the central theme of current neo-Darwinian theory)” (Miller and Newman 2003, preface). ‘The crucial point is that these complex problems actually concern more than one question. These problems, or problem agendas, are ‘lists’ of interrelated questions (both empirical and conceptual), united by a connection to natural phenomena (Love 2005, 20083). As a consequence, problem agendas are indicative of long-term investigative Programs and require contributions from more than one disciplinary approach. The interrelations among the questions are not haphazard but constitute an anatomy or problem. structure, which provides an organizational framework for coordinating interdisciplinary inquiry and governing what must be included to produce an acceptable explanation. This coordination derives from the criteria of explanatory adequacy associated with problem agendas. Assessing answers to questions in a problem agenda requires criteria of adequacy that determine the acceptability of proposed answers. Although often only implicit in research, these criteria can be isolated by a process of abstraction and reconstruction from. scientific discourse (Love 2008¢). ‘The problem agenda of evolutionary innovation and novelty is a signature item in Evo- devo (Miller and Newman 2005; Love 2003a; Miiller and Wagner 2003). Evolutionary innovations and novelties are distinctive features, such as avian flight or the vertebrate head, whose origination marked a significant departure ftom the variation available in ancestral lineages. These structures and activities are claimed to be outside the explanatory capabilities of contemporary evolutionary theory, understood as selection operating on phenotypes and sorting genotypes, thereby altering allele frequencies (Miiller and Newman 2005; Kirschner and Gerhart 2005; Wagner et al. 2000). This judgment derives from the fact that development is not factored into most of the explanatory models from evolutionary genetics. The problem agenda contains empirical questions such as “how did vertebrate jaws originate?” or ‘how did avian flight originate?” and conceptual questions about the relative contributions of different variation generating mechanisms, such as the regulation of gene expression or phenotypic plasticity due to environmental variables. We can characterize the problem agenda as trying to explain how qualitatively new phenotypic variation originates at particular phylogenetic junctures. This is an inquiry about development from the viewpoint of phylogeny. comparing discrepancies between past ranges of variation and those presently available. Innovation can be equated with the origin of (activity) function features and novelty with the origin of form features (Love 2006). This introduces our first criterion of adequacy: any adequate explanatory framework for the origin of new features must treat both form and function, ideally in terms of form- function complexes or characters (Wagner 2001). Explaining the origin of vertebrate jaws (@ novelty) requires examining how jaws function because the articulation of bones, muscles, and nerves that support this activity shaped the patterns of variation at the phylogenetic juncture. Different reasoning styles will emphasize form more than function (and vice versa), but the criterion of adequacy serves as a reminder to address (ultimately) the two in combination. Second, any adequate framework must confront the origination of innovations and novelties at all levels of biological organization and express relations between these levels relevant to the production of variation. These relations can be understood as spatial compositional’) or temporal (‘procedural’) hierarchies, in both developmental processes. and evolutionarily across generations; some entities are parts of larger entities, and some events are caused by or occur after earlier events. When combined with the first criterion, a possibility space begins to emerge and directs us to inquire more specifically into these Kinds of relations to comprehend the origination of phenotypic variation. A compositional © Springer264 A.C. Love ‘Table 2. Logical possibilities involved in assessing the adequacy of explanations for evelutionsry innovations and novelties with respect tothe level of biological hierarchy in conjunction with the form-function Hierarchy Form Function Time Developmental Compositional me Evolutionary Developmental Procedural “r Evolutionary Hierarchies can be either compositional (part-whole) or procedural (control or process dependence), con- cer either form or function features, and occur in developmental time (within a single generation) or evolutionary time (across generations). Modified from Love (2006) hierarchy for a novelty (form) requires understanding part-whole relations, whereas a procedural hierarchy for an innovation (function) draws attention to activities operating sequentially. Adding both developmental and evolutionary “time” enlarges the space of possibilities. For example, a procedural function hierarchy in developmental time for the origin of neural crest cell migration requires comprehending gene expression involved in the folding of the neural tube prior to the gene expression involved in the detachment or migration of neural crest cells—specific function features must activate serially or jointly prior to the operation of the innovation during the developmental process in a’ single generation. Eight areas of possibility can be distinguished (Table 2) and must be addressed to explain particular novelties or assess whether a general explanatory framework is, adequate. ‘The final criterion of adequacy—degree of generalization—pertains to how different explanatory answers given to individual questions within the problem agenda relate to each other. Can investigations of particular novelties be generalized to other research on different innovations or novelties? Can investigations of model systems be generalized to the phylogenetic juncture relevant to the innovation or novelty under scrutiny? The former question is about generalization from one case to another, and the latter about applying generalizations from contemporary model systems used in developmental research to putative taxa at the phylogenetic juncture for a particular novelty or innovation. These abstract questions can be linked to more specific questions. Do explanatory principles invoked to explain the vertebrate jaw also apply to the origin of cephalopod appendages? Is. the developmental patterning of the branchial arches in zebrafish a relevant model for comprehending jaw origins? When combined with the other criteria, the explanatory burden becomes concrete: are the compositional and procedural hierarchies related to various form and function features (cell types, bones, gene expression, cell migration, muscle firing), observed in the developing zebrafish jaw, appropriate models for explaining, how phenotypic variation originated at the appropriate phylogenetic juncture ~425 mil- lion years ago? ‘The problem agenda of innovation and novelty is complex but its criterion of adequacy show that the complexity is structured. Once these criteria of adequacy are made explicit, they act as a template for indicating where and how different explanatory contributions are made. If we distinguish evolutionary pattem from process, we produce a two-step explanatory architecture. For the pattern component we establish the phylogenetic context, and non-homology of the trait in question. For the process component we elucidate causes that generate phenotypic variation during ontogeny relevant to the origination of the innovation or novelty in question. Some of the interdisciplinary prerequisites of an D SpringerInterdisciplinary Lessons for the Teaching of Biology 265 PATTERN PROCESS Level of Explanation Non homology ‘Mechanistic Accoum | Organization Component | Phyfogonote Juncruro or Variation Behavior! Biology vucven ‘uontative Genetics sleontotogy/Morphe Functional Morpholo STRUCTURES Pateontology Morphology ohology rucrur Comparative Dovoiopmont omens siacptnary | Siena (Morolescad Elgon Tissues Approaches Phenotype Plastcty Lowen cen syste (Molen CQuanttaive Genetics | ORGANELLE Ans eel Developmental Genetics | GENETIC Developmenal ones Molecular Bology/Gonetcs | BIOCHEMICAL Fig. 1A schematic representation of the disciplinary contributions required to explain evolutionary novelties. Criteria of adequacy specify how different disciplines make their explanatory contribution, This schematic does not capture all ofthe criteria because it represents only form features (novelties); the origin ‘of novel functions (innovations) is ignored (from Love 2008a, 20103) explanatory framework can be visualized by mapping aspects of the criteria of adequacy onto this two-step structure (Fig. 1). This problem agenda anatomy, abstracted from empirical and conceptual research questions, partitions explanatory responsibilities and thereby identifies the different disciplinary contributions required. It also demonstrates why a reductionist epistemology cannot account for the origin of innovations and novelties—the criteria of explanatory adequacy demand more. The origination of form and function features via ontogenetic processes at many different, interacting, hierarchical levels in different dimensions of time with sufficient generality cannot be adequately explained without conceptual resources and methodologies that encompass form and function at those different levels and their many causal interactions in more than a few isolated cases. Explanatory generalizations at lower levels of organization do not propagate ‘upwards’ and break down at higher levels of organization as a consequence of dissociations between gene expression and morphology (Brigandt 2010a; Love 2008a; Minelli 1998; Wray 1999). Since many innovations and novelties exist at higher levels of organization (c.g., the vertebrate jaw), a reductionist epistemology concentrating on lower levels of organization alone cannot get purchase on the very phenomena motivating the explanatory project in the first place. ‘The interdisciplinary lesson to take away from this analysis is that complex problems structure research rather than hypothesis testing or theory confirmation. We must teach students the anatomy of problems in biology because this is what guides and motivates investigation even as theoretical perspectives change: “The continuing commitment on the part of scientists to certain procedures, goals, [and] problems, ... supply most of the con- tinuity in science” (Hull 1976, 656). The problem agenda of evolutionary innovation and novelty is just one of many in evolutionary theory that demands interdisciplinarity. Although hypotheses are constantly being tested and evolutionary theory is well confirmed, scientific inquiry is animated by the aim of integrating contributions to address key questions in these different problem agendas. We can even represent evolutionary theory as a D Springer266 A.C. Love Innovation CEO Novelty and Development Variation Form and Function Classification Inheritance Phylogeny Heredity Evolvability Fitness History of Life Coevolution ar Biogeogr Diversity Speciation Systematics Fig. 2 An idealized picture of the problem agendas in evolutionary theory along with indications of their {nterelations, and overlaid with three major areas of biology: (1) systematics, (2) genetics, cell, and development biology, and (3) ecology (from Love 2010a) constellation of interrelated problem agendas (Fig. 2), clustered by major divisions in biological science (Genetics, Cell, and Developmental Biology: Ecology: Systematics). ‘This helps to explain why problem agendas are tightly associated with disciplinary approaches within these divisions, and why explanatory success in one problem agenda is, compatible with stagnation in another: empirical and conceptual progress in evolutionary biology is regional, rather than global. Combine this with the finding that problem oriented teaching methods have generated significant advances in student learning and concept, mastery (Cooper et al. 2006), and we have a trump card to the Red Queen's pessimistic proverb, 4 Methodological Blind Spots: Idealization and Interdisciplinarity ‘Thus far we have seen that style differences in and around Evo-devo's interdisciplinarity remind us of the heterogeneity of reasoning in biology, and that evolutionary explanations involve different combinations of disciplinary resources depending on the problem agendas D SpringerInterdisciplinary Lessons for the Teaching of Biology 267 in view—biological research is structured by problems. The emphasis has been on how these approaches can produce a richer overall understanding of biological phenomena despite their potential incompatibility. But sometimes our descriptions and explanations are successful because they ignore aspects of natural phenomena or involve approximations. Idealization is a reasoning strategy that scientists use to describe and explain that purposefully departs from features known to be present in nature; it knowingly ignores Variation in properties and excludes values for variables (Jones 2005; Weisberg 2007). One type of idealization found in some Evo-devo investigations impinges on our understanding of phenotypic plasticity—the capacity of a genotype to generate phenotypic variation in response to differential environmental cues (Pigliucci 2001; Gilbert and Epel 2009). ‘The study of ontogeny in model organisms is usually executed with a set of normal stages for embryonic development, which enables researchers in different laboratories to have standardized pictorial comparisons of experimental results (Hopwood 2005, 2007). These stages are a type of idealization because they intentionally ignore kinds of variation when representing development, including variation associated with phenotypic plasticity via strict control of environmental variables. Evo-devo studies of phenotypic plasticity concentrate on identifying its underlying molecular genetic mechanisms in ontogeny (Pigliucci 2001; Kirschner and Gerhart 2005; Frankino and Raff 2004). There is a tension between the phenomenon of plasticity and the practice of staging because developmental variation that might be relevant to evolution is “removed” in the process of rendering ontogeny experimentally tractable and representing it in a standardized fashion (Love 2010b), ‘The practices of developmental biology were not tailored to study phenotypic plasticity. Variation is often viewed as ‘noise’ that must be reduced or eliminated (Frankino and Raff 2004), Animals and plants that exhibit less plasticity are chosen, which is correlated with other experimentally useful features such as short generation time and rapid development (Bolker 1995), and then raised under stable environmental conditions to minimize variation. Due to the ineliminability of variation and differences in laboratory contexts, absolute chronology is not a reliable measure of time when studying ontogeny. Therefore, the trajectory from fertilized zygote to adult is partitioned into distinct temporal periods (stages) by reference to the occurrence of morphological events that are easily identifiable by the naked eye or light microscope, and can be depicted with one or two iconic representations. ‘These developmental stages compose a ‘periodization’ that intentionally ignores variation associated with phenotypic plasticity by judgments of embryonic typicality. Material Practices (choosing model organisms, strictly controlling variables during ontogeny) and conceptual practices (using normal stages to represent typical development) positively reinforce each other and make phenotypic plasticity that much more difficult to observe. Once constructed, it is possible to use stages as a visual standard against which to describe variation as a deviation from the norm. But, more typically, variation that is ignored in the making of stages is also ignored or treated as noise (a problem to be eliminated) when encountered in day-to-day research (Frankino and Raff 2004). Normal stages fulfill a number of goals related to the descriptive and explanatory endeavors of developmental biologists. They provide a way to secure reproducibility, enable consistent and unambiguous communication among researchers, facilitate accurate predictions of developmental phenomena, and aid in making comparisons or generalizations across species.” But sometimes the morphological indicators overlap temporally, terminology that * A staging series is a tool that provides accuracy in developmental studies ..because different embryos, even together within a single clutch, develop at slightly different rates. ...Comparisons reveal more of this variability among embryos from different clutches ... Genetic uniformity may alleviate but does not © Springer268 A.C. Love is useful for one purpose may be misleading for another (especially in cross-species comparisons), and normal stages can encourage overlooking the significance of variation. ‘Thus, there are good reasons for adopting normal stages and these help explain why their continued use yields empirical success. Since developmental stages involve ignoring types of known variation, they are a form of idealization: representations of developmental phenomena based on observational features and measurement techniques that intentionally set aside variation to depict a typical case for concrete descriptive and explanatory purposes. These idealizations of ontogeny allow for a classification of developmental events, that is comprehensive, manageable. reasonably homogeneous with sharp boundaries between stages, and stable under different investigative conditions (Dupré 2001; Griesemer 1996). They accomplish these goals by treating particular forms of variation as less relevant to explaining (because invariant features are more explanatory) or not in need of explanation (because variation is akin to “noise"). Idealizations also can facilitate extrap- lating findings from the investigative context of a model organism to other domains (Love 2008b, 2009a; Steel 2008). ‘We can summarize the tension between the practice of staging and the phenomenon of plasticity as follows: (a) Variation due to phenotypic plasticity is a normal feature of ontogeny; (b) the developmental staging of model organisms downplays variation associated with the effects of environmental variables (e.g., phenotypic plasticity) by strictly limiting the range of their values and by not representing variation in characters utilized to establish the periodization; and, therefore (c) studying model organisms via normal stages will make it, difficult, if not impossible, to observe patterns of variation due to phenotypic plasticity. The fact that phenotypic plasticity is sparsely documented could be a spur to investigate it more thoroughly, but the opposite conclusion is often drawn: absence of evidence is taken to imply evidence of its developmental insignificance. But now the difficulty for Evo-devo is evident. ‘The documentation of these patterns of variation is precisely what is required to gage whether phenotypic plasticity explains evolutionary pattems or the origin of novelties (Robert 2004; West-Eberhard 2003). The practice of developmental staging in model organisms can retard our ability to assess the evolutionary significance of phenotypic plasticity. A negative assessment of its significance is likely because the variation will not be manifested or documented, and therefore is unlikely to be reckoned as substantive. This tension is interdisciplinary in character. The most powerful experimental systems for studying development (from one disciplinary perspective) are set up to overlook variation that may be critical to comprehending how evolutionary processes occur (from another disciplinary perspective). If Evo-devo investigations revolve around a character that was assessed for typicality to underwrite the temporal partitions in normal stages, then much of the variation in this character was conceptually removed in advance. For example, the conventional periodization for post-embryonic ontogeny in arthropods isin terms of molt-to- molt intervals subsequent to hatching, which are then grouped into stages (e.g.. larva, pupa, and imago), Although this periodization has a legitimate function in studying arthropod ontogeny, and has been a key ingredient of empirically fruitful reasoning in past investigations, a molt-molt periodization is unhelpful to probe molt-timing evolution because variation with respect to time has been eliminated (Minelli et al. 2006; Love 2009). The advantages of idealization for studies of ontogeny are accompanied by blindness to variation. that might be relevant to Evo-devo investigations of phenotypic plasticity. Footnote & continued climinate this problem; even embryos of a clonal stain... develop asynchronously (Kimmel et al. 1995, 253). D SpringerInterdisciplinary Lessons for the Teaching of Biology 269 Is there a way to address these blind spots? In answering we must remember that reasoning strategies involving idealization, such as normal stages, are necessary to the successful prosecution of biological investigations. Thus, any compensatory tactics should be chosen to redress the specific blind spots arising from the idealizations utilized—they must be related to the effects of ignoring variation due to phenotypic plasticity that result from the developmental staging of model organisms. We need practices that promote the observation of variation due to phenotypic plasticity that is ignored when normal stages are constructed, One tactic is to adopt an alternate periodization: choose different characters to construct new temporal partitions that facilitate observing variation with respect to char- acteristics previously stabilized in the normal stages (Minelli 2003, 57ff). These alternative periodizations can divide developmental events according to processes or landmarks that differ from those used to construct normal stages. They are useful not because of being more comprehensive but because they involve different idealizations that overlap and crosscut the temporal partitions in standardized stagings of embryogenesis. A new periodization was constructed for final-instar wing disk ontogeny in the buckeye butterfly in order to uncover variability due to phenotypic plasticity because the temporal measures of larval ontogeny do not correlate tightly with the state of the wing disks (Reed et al. 2007). ‘Substantial variability has been uncovered in this and other cases (¢.g., Mabee et al. 2000). A third interdisciplinary lesson is now at hand: we must convey to students that different disciplinary approaches in biology use reasoning strategies that simultaneously reveal some features of living systems and systematically obscure or overlook others. Normal stages are a productive idealization for studying ontogeny but this empirical success is purchased at a price. That this occurs becomes more apparent in interdisciplinary research like Evo-devo where tensions between different strategies are manifested (similar to what wwe observed for form and function reasoning styles). Instead of giving up proven reasoning strategies (such as normal stages) because they overlook aspects of biological phenomena (like variation due to phenotypic plasticity), new idealizations (alternative periodizations) are formulated that exhibit different trade-offs between what is revealed and what is, obscured, There is no skeleton key methodology for comprehending living systems and we should not expect one (Wimsatt 2007). Students need to know that our diverse and sometimes incompatible reasoning strategies are fallible and illuminating simultaneously. 5 Interdisciplinary Lessons for Pedagogy It is now time to take stock. We have identified three broad lessons from the interdisciplinary practices in and around Evo-devo. © Reasoning in biological science is not homogeneous; biological science is composed of multiple perspectives that correspond to diverse explanatory aims and exhibit divergent reasoning styles. We ‘must teach the heterogeneity of reasoning in biology. (I) Problem (agendas) structure and guide biological research rather than hypothesis testing of theory confirmation. Scientific inquiry is animated by key questions in these different problem agendas that requize integrating different contributions, We must teach students the anatomy of problems in biology. (AD Different disciplinary approaches use reasoning strategies that simultaneously reveal some features and systematically obscure or overlook others. Looking at living systems from one per ‘spective means overlooking living systems in other ways. We must teach that biological reasoning, Sirategies can be incompatible and illuminating at the same time. © Springer270 A.C. Love Each of these lessons points towards a non-reductionist epistemology in biology: instead of a single disciplinary approach being the most explanatory or fundamental, different methodologies from biological sub-disciplines must be synthesized to generate empirically adequate explanations. The lessons also fall broadly within the domain of nature of science (NOS) discussions in science education. NOS themes include the theory- adenness of observations and the provisional character of scientific knowledge (Efi et al 1999; Lederman 1992; Kampourakis and McComas 2010), but are not exhausted by abstract philosophical considerations (Allchin, in press). Rather, rich case studies (historical and contemporary) need to be constantly under a student's intellectual microscope. One advantage of these lessons from Evo-devo is that they emerge from concrete conceptual and material practices (the legitimacy of functional homology; explaining evolutionary novelties; and overlooking phenotypic plasticity due to normal stages). They also Tend themselves to a more explicit discussion of NOS in the curriculum. These lessons are not merely a laundry list but result from analyzing scientific inquiry—they are not simply declarative but functional in the sense of being transferable to new circumstances (Ford 2008). Interdisciplinary practices touch multiple aspects of NOS, including the existence of alternative explanations and methods across disciplinary approaches, the roles of consil- ence and collaboration, the variability in experimental practices (e.g., model organism. choices), and different standards for what counts as evidence in interdisciplinary situations (Allchin 2010, in press). ‘A well-chosen example can go a long way toward fulfilling these goals in the classroom context. Consider the age-old question, ‘how did the leopard get its spots’ (Allen et al. 2010)? The inter inary lessons suggest teaching this case from multiple perspectives: (I) what is the difference between reasoning about felid coat patterns from the perspective of function (advantages of camouflage) and the perspective of form. (shape or arrangement of coat pattern)? (II) what problem agendas structure the different explanatory approaches (morphogenesis from developmental biology to understand pattem generation; adaptation from evolutionary biology to understand the fitness affects of camouflage; phylogenetic reconstruction from systematics to understand the coat pattern distributions across felids)? and, (III) how do the successful methodologies used inhibit us from seeing other features due to idealizations? Some of these idealizations include the values and variables used to code camouflage patterns (simple/complex; regular! irregular; large/small) and their developmental generation (modeled as a reaction-diffusion mechanism with five dimensions such as degree of pattern regularity or complexity), as well as the values and variables for habitats (c.g., temperate forest; riparian zones; grasslands and plains) and habitat preferences (time spent in habitat [absent/ sometimes present/preferemtially present]). All three interdisciplinary lessons can be displayed concretely in this one case study T observed at the outset that an emphasis on the practice of science is in accord with teaching science as inquiry. Thus far I have construed inquiry primarily as content, something to be conveyed to students, But these interdisciplinary lessons can be extended to teaching methods as well. Self-directed assignments that probe multidimensional case studies in and out of the classroom can be sensitive to all three lessons. Students can be given character traits in organisms and assigned inquiry tasks with a forced orient toward form or function (or different senses of function), and then challenged to reconcile differences between them. Specific problem agendas can be dissected to ascertain associated criteria of adequacy and the different conceptual resources required for empirically adequate explanations. Particular periodizations can be scrutinized for blind spots and students can propose compensatory tactics for the disciplinary approaches where those D SpringerInterdisciplinary Lessons for the Teaching of Biology am periodizations are utilized.” These kinds of assignments hold the potential to interweave the content and practices of biology into a single package because empirical content must be learned along the way: the fitness effects of coat patterns, the nature of reaction— diffusion models in development, and the methods for reconstructing phylogenies (inter alia). There are substantive connections between these interdisciplinary lessons and the National Science Education Standards (NSES] (1996) and Benchmarks for Scientific Lit- eracy [BSL] (2009). Throughout the NSES there is a stress on scientific inquiry generating knowledge via “different types of investigation” (176) that harmonizes with the existence of diverse practices (reasoning styles or idealizations). This fits hand and glove with scientific inquiry involving the recognition and analysis of alternative explanations and models (148, 175) because these diverse practices often derive from different disciplinary approaches. Different disciplinary contexts can have distinct evidential standards, which means the evaluation of explanations involves more than empirical confirmation (“Evi- dence, Models, and Explanation”). This is illustrated nicely in the example of how physics. chemistry, biology, and medicine investigate the process of respiration: “understanding takes different perspectives...each is likely to give greater emphasis to concepts that have special significance in their particular discipline” (91),"° Inquiry tasks with a forced orientation toward form or function can foster the idemtifi- cation and development of individual student strengths and weaknesses (a changing emphasis in the NSES). These tasks are enhanced by social dynamics, thereby providing a platform for nurturing collaboration among students (50); groups adopt different reasoning, styles and then attempt to reconcile them subsequently. And these reasoning styles link directly to unifying concepts (Form and Function) and principles that undergird content standards (e.g,, “Structure and Function in Living Systems” [156]). Finally, the concepts of “Systems, Order, and Organization” behind content standards at all levels reinforce a problem-structured view of inquiry and the value of idealizations: “The natural...world is, complex; it is too large and complicated to investigate and comprehend all at once. Students and scientists lear to define small portions for the convenience of investigation” (116). For the BSL (AAAS Project 2061 (2009), Chap. 1), the limitations of conclusions drawn from science are represented in idealizations, whose success involves ignoring features ‘known to obtain in nature. They exemplify how believing in the unity of nature is subtle because “the belief that knowledge gained by studying one part of the universe can be applied to other parts ... turns out to be true only part of the time.” The BSZ. picks up on reasons that underlie normal stages—the control of variables to unambiguously interpret, experimental outcomes—and the advantages of scientific practices (“tools...often give more information about things than can be obtained by just observing things unaided”). Problem-structured aspects of scientific inquiry and its accompanying interdisciplinarity reinforce a case study approach when teaching biology, as well as highlight the complexity of scientific inquiry." ° Tt is worth noting that these teaching methods resonate with perspectives offered by John Dewey on empirical inquiry, and hence his philosophical outlook on education: "Selective emphasis, choice, is inevitable whenever reflection occu. ..Deception comes only when the presence and operation of choice is concealed, disguised, denied. ...Honest empirical method will state when and where and why the act of selection took place, and thus enable others to repeat it and {est its worth” (Dewey 1981/1925, 34), °© ‘The professional development of teachers also requires the ability “to make conceptual connections within and across science disciplines” (59). 44 “Case studies provide opportunities to examine such matters as the theoretical and practical limitations of science, [and] the differences in the character of the knowledge the different sciences generate”; © Springer2m A.C. Love The interdisciplinary lessons from Evo-devo's practices portray how “sometimes similar investigations give different results because of differences in the things being investigated, [and] the methods used,” and that “scientific investigations may take many different forms [and)...are conducted for different reasons... There are different traditions in science about what is investigated and how.” Diversity in scientific inquiry is depicted bby form versus function reasoning styles, the different problem agendas that exercise biologists, the different disciplinary contributions required to address these agendas, and the sometimes incompatible practices that emerge from different explanatory aims (e, functional homology or normal stages and phenotypic plasticity). “No matter how the curriculum is organized, it should provide students with opportunities to become aware of the great range of scientific disciplines that exist...the focus of such studies should be substantive (what are typical studies like in the discipline).” In addition to providing this substance, these lessons also justify organismal biology amidst predominant molecular methodologies because “some scientific knowledge is very old and yet is still applicable today.” Although there is a healthy synergy between these interdisciplinary lessons and the expectations set forth in the NSES and BSL, there are important discrepancies.'* The NSES content standards emphasize “having students reflect on the concepts that guide the inquiry” (174), but there is no hint of how problems provide this structure. Rather, the framing is in terms of hypotheses.'’ It is “theories [that] compete for acceptance” and science is construed as “the testing, revising, and occasional discarding of theories” (BSL). ‘There is repeated mention of students learning how to “identify questions that can be answered through scientific investigations” (NSES, 145), but next to no discussion of how different questions might be interrelated or cluster into broader domains. If one of the changing emphases is “assessing rich, well-structured knowledge” (100), a key question concerns what is structuring the knowledge; the anatomy of problems is a part of this assessment. The internal heterogeneity and interdisciplinarity of evolutionary biology (Love 2010a) can be used to teach why controversy in one area does not infect others and why the pace of progress differs across disciplines today and throughout history. Inquiry is ofien organized by problem agendas whose criteria of adequacy demand coordinating multiple theories in a variety of configurations, not a linear movement from puzzlement through hypothesis to theory: “We don’t solve the problem by choosing one or another of the theories—we must bring all of them to bear” (Wimsatt 2007, 163). Another discrepancy pertains to the assumption that form and function describe “complementary aspects of objects, organisms, and systems in the natural...world” (NSES, 119). This misses the potential for conflicts between reasoning styles. It is not just students who need to be informed: “teachers need to experience the value and benefits of coop- erative work as well as the struggles and tensions that it can produce” (NSES, 61). Another discrepancy is between idealization and “describing things as accurately as possible” (BSL). Idealizations are intentionally “inconsistent” with what we already know and Footnote 1 continued “disciplines differ from one another in what is studied, techniques used, and outcomes sought...many problems are studied by scientists using information and skills from many disciplines.” " The interdisciplinary lessons also challenge particular perspectives on NOS. If problems structure and fuide biological research rather than hypothesis testing or theory confirmation, then science should not be construed primarily as a hypothetica deductive endeavor (et. Allehin 200%; Lawson 2003). % “Students should...demonstrate the logical connections between the scientific concepts guiding a hypothesis and the design of an experiment” (WSES, 175); "Hypotheses are widely used in science for choosing what data to pay attention to and what additional data to seek” (BSL). D SpringerInterdisciplinary Lessons for the Teaching of Biology 23 therefore in tension with the stress on consistency and the unity of nature (BSE). These inconsistencies are not progressively resolved but rather proliferate as inquiry succeeds. ‘There is one important caveat surrounding the affirmations and challenges of the NSES and BSL by the three interdisciplinary lessons from Evo-devo, The level of life science education for much of my discussion is undergraduate college biology. Many of the highlighted features from the NSES and BSL concern lower levels where these lessons might seem less germane. While this disparity is genuine, itis also important to observe that my affirmations and challenges occur across the spectrum of the NSES and BSL, and therefore refer to systematically embedded themes. Generating knowledge using different investigatory perspectives, the differences between form and function reasoning styles, and the use of idealization are relevant at all levels of biology education. The hypothesis, testing, theory confirmation paradigm that is in conflict with the interdisciplinary lesson that problem agendas guide and structure biological investigation is promulgated throughout, possibly more egregiously at the undergraduate level than at younger grades. ‘Numerous questions remain, especially about assessment: what measures will reliably ascertain whether students have grasped these lessons (NSES, Chap. 5; Allchin, in press)? One possibility is to involve students in assessment while they adopt the role of disciplinary representative. Criticism of work from groups of students working as representa- tives of a different perspective can lead to novel, illuminating feedback. Another possibility is to propose biological questions of different kinds to see if students can categorize them in terms of their problem agenda or whether they encourage a particular reasoning style. But the difficulty of these kinds of assessments accompany the complexity of teaching science and prompt us to recall the gap between the practice of biology and what is necessary for students of different ages to Team, Ironically, the dissimilarities between educational demands and authentic inquiry might be lessened through heeding these interdisciplinary lessons, and thereby help to correct naive views of NOS that persist, even among undergraduates. 6 Concluding Remarks My discussion has been oriented around philosophical considerations that emerge from the interdisciplinary practices of Evo-devo: the differences between form and function reasoning styles, illustrated with the problematic status of functional homology (Sect. 2); the structuring of scientific investigation and explanation by problems agendas, displayed in the case of evolutionary innovations and novelties (Sect. 3); and the role of successful idealizations in systematically overlooking important biological phenomena, exemplified by how normal stages used to study model organism ontogeny obscure phenotypic plasticity (Sect. 4). As noted, there is more philosophical discussion about Evo-devo than I have canvassed here, including analyses of key concepts like developmental constraints (Amundson 1994), modularity (Winther 2001), and evolvability (Brigandt 2007; Love 2003b), or discussions of the nature of typological thinking (Brigandt 2007; Love 2009a). ‘The three interdisciplinary lessons for the teaching of biology from the practice of Evo- devo are aimed at dissolving the dilemma of either teaching Evo-devo content as con- troversial or maintaining a finger on the unruly pulse of current research, As a consequence, the illogical proclamations of the Red Queen can be blunted: our efforts can draw us beyond the status quo without taxing us to the point of exhaustion by having to run twice as fast. As Alice replied to the Red Queen “I'd rather not try, please!” To which we can add—there is a better way. © Springer214 A.C. Love Acknowledgments I am grateful to Douglas Allchin, Ingo Brigandt, Tom Doyle, Kostas Kampourakis, ‘Alessandro Minelli, and Molly Paxton for incisive comments and helpful suggestions on earlier versions of | this paper. Although I could not incorporate all of their recommendations, the resulting paper benefited tremendously from those that I did, In particular, Tom Doyle convinced me that much more shold be ssid about how my analysis bears on conventional models of teaching and translates into different classroom conlexis. My ignorance of the former and lack of experience with the latter necessitate postponing the requisite discussion to a furure date References AAAS Project 2061. (2009). Benchmarks for scientific literacy. New York: Oxford University Press. http://www project2061.org/publications/bsVonline). Abouheif, E. (1997). Developmental genetics and homology: A hierarchical approach. Trends in Ecology de Evolution, 12, 405-408, Abouheif, E., Akam, M., Dickinson, W. 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