Using Avida-ED for Teaching and Learning About Evolution in Undergraduate Introductory Biology Courses

Evolution is a complex subject that requires knowledge of basic biological concepts and the ability to connect them across multiple scales of time, space, and biological organization. Avida-ED is a digital evolution educational software environment designed for teaching and learning about evolution and the nature of science in undergraduate biology courses. This study describes our backward design approach to developing an instructional activity using Avida-ED for teaching and learning about evolution in a large-enrollment introductory biology course. Using multiple assessment instruments, we measured student knowledge and understanding of key principles of natural selection before and after instruction on evolution (including the Avida-ED activity). Assessment analysis revealed significant post-instruction learning gains, although certain evolutionary principles (most notably those including genetics concepts, such as the genetic origin of variation) remained particularly difficult for students, even after instruction. Students, however, demonstrated a good grasp of the genetic component of the evolutionary process in the context of a problem on Avida-ED. We propose that: (a) deep understanding of evolution requires complex systems thinking skills, such as connecting concepts across multiple levels of biological organization, and (b) well designed use of Avida-ED holds the potential to help learners build a meaningful and transferable understanding of the evolutionary process. An erratum to this article can be found at     

Phylogenetic approaches to nomenclature: a comparison based on a nemertean case study

Phylogenetic approaches to biological nomenclature are becoming increasingly common. Here I compare the behaviour of two such approaches, the phylogenetic system of definition and the phylogenetic system of reference, when there is a shift in the preference of phylogenetic hypotheses. The comparison is based on a case study from nemertean systematics and is the first to compare two different phylogenetic approaches throughout three stages of change, including two stages of phylogenetic nomenclature. It is concluded that a phylogenetic system of reference in combination with uninomials is superior in conveying phylogenetic information.     

Stems,nodes, crown clades,and rank‐free lists: is Linnaeus dead?

Recent radical proposals to overhaul the methods of biological classification are reviewed. The proposals of phylogenetic nomenclature are to translate cladistic phylogenies directly into classifications, and to define taxon names in terms of clades. The method has a number of radical consequences for biologists: taxon names must depend rigidly on the particular cladogram favoured at the moment, familiar names may be reassigned to unfamiliar groupings, Linnaean category terms (e.g. phylum, order, family) are abandoned, and the Linnaean binomen (e.g. Homo sapiens) is abandoned. The tenets of phylogenetic nomenclature have gained strong support among some vocal theoreticians, and rigid principles for legislative control of clade names and definitions have been outlined in the PhyloCode. The consequences of this semantic maelstrom have not been worked out. In pratice, phylogenetic nomenclature will bc disastrous, promoting confusion and instability, and it should be abandoned. It is based on a fundamental misunderstanding of the difference between a phylogeny (which is real) and a classification (which is utilitarian). Under the new view, classifications are identical to phlylogenies, and so the proponents of phylogenetic nomenclature will end up abandoning classifications altogether.     

Naming diversity in an evolutionary context: Phylogenetic definitions of the Roucela clade (Campanulaceae/Campanuloideae) and the cryptic taxa within

In recent times, evolution has become a central tenet of taxonomy, but nomenclature has consistently been decoupled from the tree‐thinking process, often leading to significant issues in reconciling traditional (Linnaean) names with clades in the Tree of Life. Recent evolutionary studies on the Roucela clade, a group of endemic plants found in the Mediterranean Basin, motivated the establishment of phylogenetic concepts to formally anchor clade names on the Campanuloideae (Campanulaceae) tree. These concepts facilitate communication of clades that approximate traditionally defined groups, in addition to naming newly discovered cryptic diversity in a phylogenetic framework.     

Ancient signals: comparative genomics of plant MAPK and MAPKK gene families

MAPK signal transduction modules play crucial roles in regulating many biological processes in plants, and their components are encoded by highly conserved genes. The recent availability of genome sequences for rice and poplar now makes it possible to examine how well the previously described Arabidopsis MAPK and MAPKK gene family structures represent the broader evolutionary situation in plants, and analysis of gene expression data for MPK and MKK genes in all three species allows further refinement of those families, based on functionality. The Arabidopsis MAPK nomenclature appears sufficiently robust to allow it to be usefully extended to other well-characterized plant systems.     

Common evolutionary trends underlie the four‐bar linkage systems of sunfish and mantis shrimp

Comparative biomechanics offers an opportunity to explore the evolution of disparate biological systems that share common underlying mechanics. Four‐bar linkage modeling has been applied to various biological systems such as fish jaws and crustacean appendages to explore the relationship between biomechanics and evolutionary diversification. Mechanical sensitivity states that the functional output of a mechanical system will show differential sensitivity to changes in specific morphological components. We document similar patterns of mechanical sensitivity in two disparate four‐bar systems from different phyla: the opercular four‐bar system in centrarchid fishes and the raptorial appendage of stomatopods. We built dynamic linkage models of 19 centrarchid and 36 stomatopod species and used phylogenetic generalized least squares regression (PGLS) to compare evolutionary shifts in linkage morphology and mechanical outputs derived from the models. In both systems, the kinematics of the four‐bar mechanism show significant evolutionary correlation with the output link, while travel distance of the output arm is correlated with the coupler link. This common evolutionary pattern seen in both fish and crustacean taxa is a potential consequence of the mechanical principles underlying four‐bar systems. Our results illustrate the potential influence of physical principles on morphological evolution across biological systems with different structures, behaviors, and ecologies.     

Evolutionary Medicine: A Key to Introducing Evolution

Science teachers can use examples and concepts from evolutionary medicine to teach the three concepts central to evolution: common descent, the processes or mechanisms of evolution, and the patterns produced by descent with modification. To integrate medicine into common ancestry, consider how the evolutionary past of our (or any) species affects disease susceptibility. That humans are bipedal has produced substantial changes in our musculoskeletal system, as well as causing problems for childbirth. Mechanisms such as natural selection are well exemplified in evolutionary medicine, as both disease-causing organism and their targets adapt to one another. Teachers often use examples such as antibiotic resistance to teach natural selection: it takes little alteration of the lesson plan to make explicit that evolution is key to understanding the principles involved. Finally, the pattern of evolution can be illustrated through evolutionary medicine because organisms sharing closer ancestry also share greater susceptibility to the same disease-causing organisms. Teaching evolution using examples from evolutionary medicine can make evolution more interesting and relevant to students, and quite probably, more acceptable as a valid science.     

Evolutionary concepts meet the neck of penguins (Aves: Sphenisciformes), towards a “survival strategy” for evo-devo

Evolutionary developmental biology (or evo-devo) is the scientific connectivity that allowed a more comprehensive and practical completeness in the contemporary conceptualisation of evolution. The links between genetics, developmental mechanics and evolution led to a better understanding of evolutionary mechanisms. An analysis of evolutionary concepts such as homology, homeoses, constraints, novelties, modularity, and selection is given through the recurring example of the variations identified in the modular repartition of the cervical vertebrae in extant and fossil penguins. The inclusion of this study about penguins in the evolutionary system also involves a reflection on the current state and the future of evo-devo. Three principles of assessment and method, applicable to many natural and conceptual scales, are introduced to define a ??survival strategy?? for evo-devo. The above-mentioned principles are intended to strengthen and continue the connectivity induced de facto. These current and future investigation challenges are discussed and connected to three main naturalist names related directly to the conceptualisation of evolution: Charles Darwin, étienne Geoffroy Saint-Hilaire, and Lamarck.     

Emergent insights from the synthesis of conceptual frameworks for biological invasions

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

The role of subspecies in obscuring avian biological diversity and misleading conservation policy

Subspecies are often used in ways that require their evolutionary independence, for example as proxies for units of conservation. Mitochondrial DNA sequence data reveal that 97% of continentally distributed avian subspecies lack the population genetic structure indicative of a distinct evolutionary unit. Subspecies considered threatened or endangered, some of which have been targets of expensive restoration efforts, also generally lack genetic distinctiveness. Although sequence data show that species include 1.9 historically significant units on average, these units are not reflected by current subspecies nomenclature. Yet, it is these unnamed units and not named subspecies that should play a major role in guiding conservation efforts and in identifying biological diversity. Thus, a massive reorganization of classifications is required so that the lowest ranks, be they species or subspecies, reflect evolutionary diversity. Until such reorganization is accomplished, the subspecies rank will continue to hinder progress in taxonomy, evolutionary studies and especially conservation.     

Variation in immune defence as a question of evolutionary ecology

The evolutionary-ecology approach to studying immune defences has generated a number of hypotheses that help to explain the observed variance in responses. Here, selected topics are reviewed in an attempt to identify the common problems, connections and generalities of the approach. In particular, the cost of immune defence, response specificity, sexual selection, neighbourhood effects and questions of optimal defence portfolios are discussed. While these questions still warrant further investigation, future challenges are the development of synthetic concepts for vertebrate and invertebrate systems and also of the theory that predicts immune responses based on a priori principles of evolutionary ecology.     

A theoretical framework for defining some concepts in evolution.

We present a theoretical framework for biological evolution with the intention of giving precise mathematical definitions of some concepts in evolutionary biology such as fitness, evolutionary pressure, specialization and natural selection. In this framework, such concepts are identified with well-known mathematical terms within the theory of dynamical systems. We also discuss some more general implications in evolution; for instance, the fact that our model naturally exhibits a frequency spectrum of the type 1/f for low frequencies of evolutionary events.     

On the integrated frameworks of species concepts: Mayden’s hierarchy of species concepts and de Queiroz’s unified concept of species

Richard L. Mayden and Kevin de Queiroz have devised and developed ‘a hierarchy of species concepts’ and ‘a unified species concept’, respectively. Although their integrated frameworks of species concepts are rather different as to how to integrate the diverse modern concepts of species, the end result is that they are likely to agree on species recognition in nature, because they virtually share the same major components (i.e. evolutionary or lineage concept of species; same way of delimiting species), and have the same important consequences. Both the hierarchical and unified frameworks, however, are interpreted to have shortcoming regarding the way of integrating the modern species concepts. I reformulate these ideas into a framework of species concepts as follows: It treats the idea of species as population‐level evolutionary lineages (sensu Wiley 1978 ) as the concept for species category, and it adopts the contingent biological properties of species (e.g. internal reproductive isolation, diagnosability, monophyly) as operational criteria in delimiting species. I also suggest that existing and revised versions of the integrated framework of species concepts all are not new species concepts, but versions of the evolutionary species concept, because they treat the evolutionary (or lineage) species concept as the concept for species category.     

On the relationship between content, ancestor, and ancestry in phylogenetic nomenclature

In this paper I draw attention to the concepts of content and ancestry in phylogenetic nomenclature. I argue that these concepts are tightly linked and that they cannot be separated as suggested by Bryant and Cantino [Biol. Rev. 77 (2002) 39] in their recent response to a critique of phylogenetic nomenclature. In addition, I argue that the basic assumption in phylogenetic nomenclature that a taxon name always refers to the same ancestor or ancestry is questionable.     

Museum behind the scenes–an inquiry-based learning unit with biological collections in the classroom

The aim of this study was to design and evaluate an inquiry- and activity-based learning unit for the classroom that uses biological collections to teach key evolutionary concepts and to support the understanding and appreciation of the work of a museum. The unit consisted of three parts that focused on the most important tasks of museums: collecting and conserving, researching and exhibiting. The students created their own collection, performed research surrounding it and then designed an exhibition. Seventy-six secondary sixth- and seventh-grade students participated in the testing of the prototype unit. For evaluation, we carried out a pre-/post-test design using a questionnaire that assessed content knowledge and learning enjoyment. The mean knowledge score of the post-test indicated significant learning gains compared to the pre-test results. The test on learning enjoyment showed best mean values for actions that included the collection, compared to results of the theoretical parts of the unit. This approach demonstrates that the learning unit offers the opportunity to experience the tasks of a museum at first hand and to acquire knowledge about evolutionary science and evolutionary principles.     

Phylogeny,classification and nomenclature: a reply to F. Pleijel and G. W. Rouse

In systematics, the uncovering of monophyletic units, of sister group relationships and also of paraphyla is an important part of primary research. The hypotheses derived are thus subject to falsification and are subject to change. In contrast, classifications are a secondary step, as they are derived from such hypotheses. Classifications are based on different philosophies, which permit different solutions as to how results in the fields of taxonomy and phylogenetics can be transposed into a ‘system’. The function of classifications is at least partly utilitarian, and this is even more true for the names and principles of nomenclature. Nomenclature is simply a tool for information retrieval and for safeguarding understanding. Directly linking names and cladograms or nodes, respectively – making them subject to changes by falsification – would deliberately ignore the primary, strictly utilitarian function of long‐established principles of nomenclature and would endanger an instrument that functions almost perfectly. Approaches to introduce a so‐called PhyloCode should therefore not be pursued, as there is no chance at all that this kind of code could be generally accepted.     

The Origin of Cellular Life and Biosemiotics

Recent successes of systems biology clarified that biological functionality is multilevel. We point out that this fact makes it necessary to revise popular views about macromolecular functions and distinguish between local, physico-chemical and global, biological functions. Our analysis shows that physico-chemical functions are merely tools of biological functionality. This result sheds new light on the origin of cellular life, indicating that in evolutionary history, assignment of biological functions to cellular ingredients plays a crucial role. In this wider picture, even if aggregation of chance mutations of replicator molecules and spontaneously self-assembled proteins led to the formation of a system identical with a living cell in all physical respects but devoid of biological functions, it would remain an inanimate physical system, a pseudo-cell or a zombie-cell but not a viable cell. In the origin of life scenarios, a fundamental circularity arises, since if cells are the minimal units of life, it is apparent that assignments of cellular functions require the presence of cells and vice versa. Resolution of this dilemma requires distinguishing between physico-chemical and biological symbols as well as between physico-chemical and biological information. Our analysis of the concepts of symbol, rule and code suggests that they all rely implicitly on biological laws or principles. We show that the problem is how to establish physico-chemically arbitrary rules assigning biological functions without the presence of living organisms. We propose a solution to that problem with the help of a generalized action principle and biological harnessing of quantum uncertainties. By our proposal, biology is an autonomous science having its own fundamental principle. The biological principle ought not to be regarded as an emergent phenomenon. It can guide chemical evolution towards the biological one, progressively assigning greater complexity and functionality to macromolecules and systems of macromolecules at all levels of organization. This solution explains some perplexing facts and posits a new context for thinking about the problems of the origin of life and mind.