No abstract

The field of morphology has recently seen the arrival of computer‐aided ontologies, tools which permit the semantic organization of defined concepts and which therefore promise to be extremely useful in computer‐mediated approaches involving morphological data, for example in cladistics. The theoretical relationship between ontologies and cladistics, however, has hardly been explored. Here we examine the ontological status of the main terms in morphological cladistics, i.e. morpheme, character, character state and ontological concept. Morphemes are units of the descriptional perspective, whereas character states are units of the evolutionary perspective and refer to identical stages of transformation. Both morphemes and character states represent things, i.e. real entities. However, character state and morpheme denote different perspectives on these entities (description vs. evolution). Characters (transformation series; Hennig's ideographic character concept) and ontological concepts are both classes, but not of the same nature. Ontological concepts which are used to classify morphemes are constructs, i.e. totally man‐made classes that serve only as a way of classifying real entities for human recognition. Characters, however, are classes that encompass all character states of common descent and which therefore have objective, human‐independent properties. Characters, then, are natural kinds, classes which exhibit a natural identity in the same way as monophyla do, for example, which also have common descent as an objective property. Against this background, ontology‐based morphology can be a valuable addition to phylogenetic systematics. Formalized, machine‐parsable descriptions permit the generalization of morphemes in large datasets and can facilitate the recognition of identical character states. However, the expertise of the researcher is indispensable, and full automation of the transfer from the descriptive level to the evolutionary level thus appears impossible.

The morphology of hemolymph circulatory systems has been studied in many arthropod groups over the past decades. In most cases, however, the focus of these studies has been the vascular system, while its counterpart, the lacunar system, has often been neglected. To further understanding of the interrelationships between these two complementary subsystems, we investigated both, the hemolymph vascular system and the hemolymph lacunar system, of the decapod Penaeus vannamei using 3Dimaging techniques (micro-computed tomography and confocal laser scanning microscopy) in combination with 3D reconstruction. Major parts of the vascular and lacunar system are described. Our insights into their morphology are used to derive functional conclusions for a model illustrating the interrelationships between the two subsystems. The morphology of and the functional interaction between the vascular and lacunar systems are discussed in the context of the debate on "open vs. closed circulatory systems." K E Y W O R D S

Morphology, the oldest discipline in the biosciences, is currently experiencing a renaissance in the field of comparative phenomics. However, morphological/phenotypic research still suffers on various levels from a lack of standards. This shortcoming, first highlighted as the "linguistic problem of morphology", concerns the usage of terminology and also the need for formalization of morphological descriptions themselves, something of paramount importance not only to the field of morphology but also when it comes to the use of phenotypic data in systematics and evolutionary biology. We therefore argue, that for morphological descriptions, the basis of all systematic and evolutionary interpretations, ontologies need to be utilized which are based exclusively on structural qualities/properties and which in no case include statements about homology and/or function. Statements about homology and function constitute interpretations on a different or higher level. Based on these "anatomy ontologies", further ontological dimensions (e.g., referring to functional properties or homology) may be exerted for a broad use in evolutionary phenomics. To this end we present the first organ-based ontology for the most species-rich animal group, the Arthropoda. Our Ontology of Arthropod Circulatory Systems (OArCS) contains a comprehensive collection of 383 terms (i.e., labels) tied to 296 concepts (i.e., definitions) collected from the literature on phenotypic aspects of circulatory organ features in arthropods. All of the concepts used in OArCS are based exclusively on structural features, and in the context of the ontology are independent of homology and functional assumptions. We cannot rule out that in some cases, terms are used which in traditional usage and previous accounts might have implied homology and/or function (e.g. heart, sternal artery). Concepts are composed of descriptive elements that are used to classify observed instances into the organizational framework of the ontology. That is, descriptions in ontologies are only descriptions of individuals if they are necessary/and or sufficient representations of attributes (independently) observed and recorded for an individual. In addition, we here present for the first time an entirely new approach to formalizing phenotypic research, a semantic model for the description of a complex organ system in a highly disparate taxon, the arthropods. We demonstrate this with a formalized morphological description of the hemolymph vascular system in one specimen of the European garden spider Araneus diadematus. Our description targets five categories of descriptive statement: "position", "spatial relationships", "shape", "constituents", and "connections", as the corresponding formalizations constitute exemplary patterns useful not only when talking about the circulatory system, but also in descriptions in general. The downstream applications of computer-parsable morphological descriptions are widespread, with their core utility being the fact that they make it possible to compare c...

Arthropods are the most species-rich taxon within Metazoa and have gone through major evolutionary changes with regard to body organization. Arthropod hearts and their associated vascular systems are thus morphologically highly disparate: while some arthropods exhibit very powerful hearts and complex vascular systems, other arthropods do not possess any kind of vascular system or heart at all. A comprehensive study investigating the structure of arthropods hearts has never been undertaken. In this study, we therefore investigate the hearts of 34 species from all major arthropod groups using various imaging techniques (confocal laser scanning microscopy, micro-computed tomography, histology) and describe them by addressing different aspects of heart morphology, e.g. the structure of the myocard or the composition of ostia. In a next step, we conceptualize 18 characters related to heart morphology and their respective character states and–using additional data from the literature–score a matrix for a total of 45 species from 38 supraspecific taxa. We map the characters onto prevailing phylogenetic hypotheses and perform parsimony-based ancestral state reconstruction to trace the evolutionary transformations undergone by arthropod hearts. An exploration of the character concepts (as explanatory hypotheses) reveals ontological peculiarities of character statements that clearly distinguish them in terms of ontological status from descriptive statements (i.e. descriptions of morphemes). The implications of these findings influence the interpretation of ground patterns as explanations. This first phylogenetic approach to heart morphology in the arthropod ground pattern reveals numerous new putative synapomorphies and leads to a reconsideration of the morphology of circulatory systems in early arthropods. Hypotheses on the evolution of hearts in (Pan-) Arthropoda are illustrated and discussed.

The need to protect neural tissue from toxins or other substances is as old as neural tissue itself. Early recognition of this need has led to more than a century of investigation of the blood-brain barrier (BBB). Many aspects of this important neuroprotective barrier have now been well established, including its cellular architecture and barrier and transport functions. Unsurprisingly, most research has had a human orientation, using mammalian and other animal models to develop translational research findings. However, cell layers forming a barrier between vascular spaces and neural tissues are found broadly throughout the invertebrates as well as in all vertebrates. Unfortunately, previous scenarios for the evolution of the BBB typically adopt a classic, now discredited ‘scala naturae’ approach, which inaccurately describes a putative evolutionary progression of the mammalian BBB from simple invertebrates to mammals. In fact, BBB-like structures have evolved independently numerous times, complicating simplistic views of the evolution of the BBB as a linear process. Here, we review BBBs in their various forms in both invertebrates and vertebrates, with an emphasis on the function, evolution, and conditional relevance of popular animal models such as the fruit fly and the zebrafish to mammalian BBB research.

In this study, the hemolymph vascular system (HVS) in two cambarid crayfishes, i.e. the Marbled Crayfish, Procambarus virginalis Lyko, 2017 and the Spiny Cheek Crayfish, Faxonius limosus (Rafinesque, 1817), is investigated in regard of areas of non-genetic phenotypic variation. Despite their genetic identity, specimens of P. virginalis show variability in certain features of the HVS. Thus, we describe varying branching patterns, sporadic anastomoses, and different symmetry states in the vascular system of the marbled crayfish. We visualize our findings by application of classical and modern morphological methods, e.g. injection of casting resin, micro-computed tomography and scanning electron microscopy. By comparing our findings for P. virginalis to the vasculature in sexually reproducing crayfishes, i.e. F. limosus and Astacus astacus, we discuss phenotypic variation of the HVS in arthropods in general. We conclude that constant features of the HVS are hereditary, whereas varying states identified by study of the clonal P. virginalis must be caused by non-genetic factors and, that congruent variations in sexually reproducing F. limosus and A. astacus are likely also non-genetic phenotypic variations. Both common causal factors for non-genetic phenotypic variation, i.e., phenotypic plasticity and stochastic developmental variation are discussed along our findings regarding the vascular systems. Further aspects, such as the significance of non-genetic phenotypic variation for phylogenetic interpretations are discussed.

In the field of phylogenetic systematics, the terms homology and homologue and their relationship to cladistic terms such as character, character state, synapomorphy and symplesiomorphy, as well as their relationships to each other, have been and are still discussed frequently. A recent re‐emergence of concepts of homology/homologue free of any reference to explanatory hypotheses prompted us to explore these concepts and their relationships to each other as well as to the concept of morpheme, as introduced recently. All concepts are examined with regard to their ontological status and their bearing in the epistemological process in morphology and phylogenetic systematics. To us, morphemes, homologues and in partem character states refer to things (concrete objects in the ontological sense). However, although morphemes are exclusively descriptive, the latter two represent objects of causal explanations. Homologue always refers to the things themselves, yet a character state also can be a property or the absence of a thing. In this context, a character as a transformation series of character states does not represent a thing but a natural kind. Character states of one character are connected by homology relationships, i.e. common descent. Synapomorphy and symplesiomorphy represent different states of a single transformation series. A nonexplanatory, purely descriptive, concept of homologues is contradictory to its original as well as the post‐Darwinian, evolutionary, concept which refers to causal relationships between parts of organisms and their correspondences in the archetype or ancestor, respectively. Character states, homologues and synapomorphies/symplesiomorphies can only be approximated in the form of hypotheses. We argue that the high value of the concept of homology and its related concepts for evolutionary biology should be maintained by acknowledging their explanatory nature and that dilution with nonexplanatory (even idealistic) definitions should be avoided.