The notion of homology in current comparative biology

Current notions on homology, and its recognition, causation, and explanation are reviewed in this report. The focus is primarily on concepts because the formulation of precise definitions of homology has contributed little to our understanding of the issue. Different aspects or concepts of homology have been contrasted, currently the most important ones being the distinction between systematic and biological concepts. The systematic concept of homology focuses on common ancestry and on taxa; the biological concept tries to explain patterns of conservatism in evolution by shared developmental constraints. Similarity or correspondence is generally accepted as a primary criterion in the delimitation of homologues, albeit that this criterion is not without practical and theoretical problems. Apart from similarity, the biological concept of homology also stresses developmental individuality of putative homologous structures. Structural and positional aspects of homology can be separated, with positional homology acquiring an independent status. Similarity, topographic relationships, and ontogenetic development cannot be tests of homology. Within the cladistic paradigm, the most decisive test of homology is that of congruence; proponents of the biological-homology concept have been less concerned with test implications. Adopting a hierarchical view of nature suggests that characters have to be homologized at their appropriate level of organization. A taxic or systematic approach to homology has precedence over a transformational or biological approach. Nevertheless, pattern analysis and process explanations are not independent of each other.     

The female orgasm and the homology concept in evolutionary biology

The definition of homology and its application to reproductive structures, external genitalia, and the physiology of sexual pleasure has a tortuous history. While nowadays there is a consensus on the developmental homology of genital and reproductive systems, there is no agreement on the physiological translation, or the evolutionary origination and roles, of these structural correspondences and their divergent histories. This paper analyzes the impact of evolutionary perspectives on the homology concept as applied to the female orgasm, and their consequences for the biological and social understanding of female sexuality and reproduction. After a survey of the history of pre-evolutionary biomedical views on sexual difference and sexual pleasure, we examine how the concept of sexual homology was shaped in the new phylogenetic framework of the late 19th century. We then analyse the debates on the anatomical locus of female pleasure at the crossroads of theories of sexual evolution and new scientific discourses in psychoanalysis and sex studies. Moving back to evolutionary biology, we explore the consequences of neglecting homology in adaptive explanations of the female orgasm. The last two sections investigate the role played by different articulations of the homology concept in evolutionary developmental explanations of the origin and evolution of the female orgasm. These include the role of sexual, developmental homology in the byproduct hypothesis, and a more recent hypothesis where a phylogenetic, physiological concept of homology is used to account for the origination of the female orgasm. We conclude with a brief discussion on the social implications for the understanding of female pleasure derived from these different homology frameworks.     

A comparative study of available software for high-accuracy homology modeling: from sequence alignments to structural models

An open question in protein homology modeling is, how well do current modeling packages satisfy the dual criteria of quality of results and practical ease of use? To address this question objectively, we examined homology‐built models of a variety of therapeutically relevant proteins. The sequence identities across these proteins range from 19% to 76%. A novel metric, the difference alignment index (DAI), is developed to aid in quantifying the quality of local sequence alignments. The DAI is also used to construct the relative sequence alignment (RSA), a new representation of global sequence alignment that facilitates comparison of sequence alignments from different methods. Comparisons of the sequence alignments in terms of the RSA and alignment methodologies are made to better understand the advantages and caveats of each method. All sequence alignments and corresponding 3D models are compared to their respective structure‐based alignments and crystal structures. A variety of protein modeling software was used. We find that at sequence identities >40%, all packages give similar (and satisfactory) results; at lower sequence identities (<25%), the sequence alignments generated by Profit and Prime, which incorporate structural information in their sequence alignment, stand out from the rest. Moreover, the model generated by Prime in this low sequence identity region is noted to be superior to the rest. Additionally, we note that DSModeler and MOE, which generate reasonable models for sequence identities >25%, are significantly more functional and easier to use when compared with the other structure‐building software.     

Adipocyte biology: It is time to upgrade to a new model

Globally, the obesity pandemic is profoundly affecting quality of life and economic productivity, but efforts to address this, especially on a pharmacological level, have generally proven unsuccessful to date, serving as a stark demonstration that our understanding of adipocyte biology and pathophysiology is incomplete. To deliver better insight into adipocyte function and obesity, we need improved adipocyte models with a high degree of fidelity in representing the in vivo state and with a diverse range of experimental applications. Adipocyte cell lines, especially 3T3-L1 cells, have been used extensively over many years, but these are limited in terms of relevance and versatility. In this review, I propose that primary adipose-derived stromal/stem cells (ASCs) present a superior model with which to study adipocyte biology ex vivo. In particular, ASCs afford us the opportunity to study adipocytes from different, functionally distinct, adipose depots and to investigate, by means of in vivo/ex vivo studies, the effects of many different physiological and pathophysiological factors, such as age, body weight, hormonal status, diet and nutraceuticals, as well as disease and pharmacological treatments, on the biology of adipocytes and their precursors. This study will give an overview of the characteristics of ASCs and published studies utilizing ASCs, to highlight the areas where our knowledge is lacking. More comprehensive studies in primary ASCs will contribute to an improved understanding of adipose tissue, in healthy and dysfunctional states, which will enhance our efforts to more successfully manage and treat obesity.     

It is time to move: linking flight and foraging behaviour in a diving bird

Although the adaptive value of flight may seem obvious, it is the most difficult behaviour of birds to monitor. Here, we describe a technique to quantify the frequency and the duration of flights over several months by implanting a data logger that records heart rate (fH), hydrostatic pressure (diving depth) and the body angle of a large sea duck species, the common eider (Somateria mollissima). According to the mean fH recorded during flight and the parameters recorded to identify the fH flight signature, we were able to identify all flights performed by 13 individuals during eight months. We cumulated local flight time (outside migrations) and found that activity occurs primarily during dawn and morning and that flying activities are strongly related to diving activities (Pearson's r=0.88, permutation test p<0.001). This relationship was interpreted as a consequence of living in a dynamic environment where sea currents move the ducks away from the food patches. We believe that the technique described here will open new avenues of investigation in the adaptive value of flight.     

A plea for ‘genealogical thinking’ in comparative biology – a rebuttal to the reply of Szucsich,Wirkner, and Pass to my article ‘Deconstructing Morphology’

Scholtz G. in press. A plea for ‘genealogical thinking’ in comparative biology – a rebuttal to the reply of Szucsich, Wirkner, and Pass to my article ‘Deconstructing Morphology’. —Acta Zoologica (Stockholm) 00 : 1–4. Szucsich et al. (in press) claim that – in contrast to my statement – morphological thinking has to be ‘cladistic.’ Based on this premise, they stress the difference between the relationships among states of characters versus those among structures assigned to the same character state as implemented in numerical cladistic reasoning. SEA claim that my approach to the homology concept only deals with the problem of the integration of various character states into the same character, whereas the necessary relationships among structures assigned to the same state are not covered. Based on this distinction, SEA also criticise the application of similarity in my definition of homology. Furthermore, they address the issue of evolutionarily independent units.     

Increase in body size is correlated to warmer winters in a passerine bird as inferred from time series data

Climate change is expected to affect natural populations in many ways. One way of getting an understanding of the effects of a changing climate is to analyze time series of natural populations. Therefore, we analyzed time series of 25 and 20 years, respectively, in two populations of the citril finch (Carduelis citrinella) to understand the background of a dramatic increase in wing length in this species over this period, ranging between 1.3 and 2.9 phenotypic standard deviations. We found that the increase in wing length is closely correlated to warmer winters and in one case to rain in relation to temperature in the summer. In order to understand the process of change, we implemented seven simulation models, ranging from two nonadaptive models (drift and sampling), and five adaptive models with selection and/or phenotypic plasticity involved and tested these models against the time series of males and females from the two population separately. The nonadaptive models were rejected in each case, but the results were mixed when it comes to the adaptive models. The difference in fit of the models was sometimes not significant indicating that the models were not different enough. In conclusion, the dramatic change in mean wing length can best be explained as an adaptive response to a changing climate.     

Hox genes, homology and axis formation—The application of morphological concepts to evolutionary developmental biology

This article focuses on the interphyletic comparison of gene expression patterns. By means of the hypothesis of the inversion of the dorsoventral axis during the evolution of the Bilateria, it is demonstrated, that evolutionary developmental biologists use similarities in spatial and temporal gene expression patterns as evidence for the formulation of hypotheses of homology concerning either developing structures or body regions. The molecular genetic and morphogenetic evidence used is discussed within the framework of a cladistic-phylogenetic analysis based on the phylogenetic tree of the Bilateria. I argue that similarity of spatial and temporal gene expression patterns is not a sufficient criterion for homology inference. Therefore, gene expression patterns should be coded as characters. Their homology should be tested in concert with other characters.

Furthermore, it is demonstrated, that spatial and temporal similar gene expression patterns, indicating similar molecular genetic mechanisms, were interpreted as an analytical criterion of homology, offering the possibility to identify similar structures. In contrast to this, the evolutionary developmental biolgists have not developed a causal-analytically extended concept of shape, from which a causal-analytical concept of homology could be deduced. Instead, the homology concept from evolutionary morphology is used.     

Ontogeny, anatomy, and the problem of homology: Carl Gegenbaur and the American tradition of cell lineage studies

Summary In this paper we analyze Carl Gegenbaur’s conception of the relationship between embryology (“Ontogenie”) and comparative anatomy and his related ideas about homology. We argue that Gegenbaur’s conviction of the primacy of comparative anatomy and his careful consideration of caenogenesis led him to a more balanced view about the relationship between ontogeny and phylogeny than his good friend Ernst Haeckel. We also argue that Gegenbaur’s ideas about the centrality of comparative anatomy and his definitions of homology actually laid the conceptual foundations for Hans Spemann’s (1915) later analysis of homology. We also analyze Gegenbaur’s reception in the United States and how the discussions between E.B. Wilson and Edwin Conklin about the role of the “embryological criterion of homology” and the latter’s argument for an even earlier concept of cellular homology reflect the recurring theme of preformism in ontogeny, a theme that finds its modern equivalent in various genetic definitions of homology, only recently challenged by the emerging synthesis of evolutionary developmental biology. Finally, we conclude that Gegenbaur’s own careful methodological principles can serve as an important model for proponents of present day “evo-devo”, especially with respect to the integration of ontogeny with phylogeny embedded in comparative anatomy.     

Novel computational biology methods and their applications to drug discovery

Computational biology methods are now firmly entrenched in the drug discovery process. These methods focus on modeling and simulations of biological systems to complement and direct conventional experimental approaches. Two important branches of computational biology include protein homology modeling and the computational biophysics method of molecular dynamics. Protein modeling methods attempt to accurately predict three-dimensional (3D) structures of uncrystallized proteins for subsequent structure-based drug design applications. Molecular dynamics methods aim to elucidate the molecular motions of the static representations of crystallized protein structures. In this review we highlight recent novel methodologies in the field of homology modeling and molecular dynamics. Selected drug discovery applications using these methods conclude the review.     

基于比较基因组学方法的丝状子囊菌同源序列分析

为探讨丝状子囊菌基因组的同源保守序列作为标记基因,利用Standalone BLASTN方法将构巢曲霉全基因组基因分别与30种丝状子囊菌基因组比较.构巢曲霉与每个丝状子囊菌基因组之间的同源匹配基因数量似乎可反映子囊菌之间的进化关系,构巢曲霉（10,560个基因）与15种散囊菌纲其他真菌间的匹配基因数量为5,179-7,747个,其中与另外7个同属的种匹配的基因数量为7,434-7,747个,而与亲缘关系较远的2种锤舌菌纲真菌灰葡萄孢和核盘菌的匹配基因数量分别仅有4,318个和4,242个.构巢曲霉的10,560个基因与20余种子囊菌基因组同时匹配的基因数为3,509个,占33.2％,构巢曲霉基因与30种子囊菌共同匹配的基因仅924个.此外,E值大小在10-30_0.1范围的同源序列变异性大,而在0-10-100范围的同源序列高度保守.随着基因组序列数据的增加,比较基因组方法将会在真菌系统学研究领域发挥更大的作用.     

No substitute for real data: A cautionary note on the use of phylogenies from birth–death polytomy resolvers for downstream comparative analyses

The statistical estimation of phylogenies is always associated with uncertainty, and accommodating this uncertainty is an important component of modern phylogenetic comparative analysis. The birth–death polytomy resolver is a method of accounting for phylogenetic uncertainty that places missing (unsampled) taxa onto phylogenetic trees, using taxonomic information alone. Recent studies of birds and mammals have used this approach to generate pseudoposterior distributions of phylogenetic trees that are complete at the species level, even in the absence of genetic data for many species. Many researchers have used these distributions of phylogenies for downstream evolutionary analyses that involve inferences on phenotypic evolution, geography, and community assembly. I demonstrate that the use of phylogenies constructed in this fashion is inappropriate for many questions involving traits. Because species are placed on trees at random with respect to trait values, the birth–death polytomy resolver breaks down natural patterns of trait phylogenetic structure. Inferences based on these trees are predictably and often drastically biased in a direction that depends on the underlying (true) pattern of phylogenetic structure in traits. I illustrate the severity of the phenomenon for both continuous and discrete traits using examples from a global bird phylogeny.     

Orchid biology: from Linnaeus via Darwin to the 21st century

Orchidaceae are the largest family of flowering plants, with at least 24 000 species, and perhaps better than any other family of flowering plants, orchids represent the extreme specializations that are possible. As a result, they have long fascinated luminaries of the botanical world including Linnaeus and Darwin, but the size of the family has historically been an impediment to their study. Specifically, the lack of detailed information about relationships within the family made it difficult to formulate explicit evolutionary hypotheses for such a large group, but the advent of molecular systematics has revolutionized our understanding of the orchids. Their complex life histories make orchids particularly vulnerable to environmental change, and as result many are now threatened with extinction. In this Special Issue we present a series of 20 papers on orchid biology ranging from phylogenetics, floral evolutionary development, taxonomy, mycorrhizal associations, pollination biology, population genetics and conservation.     

Using multiple templates to improve quality of homology models in automated homology modeling

When researchers build high-quality models of protein structure from sequence homology, it is today common to use several alternative target-template alignments. Several methods can, at least in theory, utilize information from multiple templates, and many examples of improved model quality have been reported. However, to our knowledge, thus far no study has shown that automatic inclusion of multiple alignments is guaranteed to improve models without artifacts. Here, we have carried out a systematic investigation of the potential of multiple templates to improving homology model quality. We have used test sets consisting of targets from both recent CASP experiments and a larger reference set. In addition to Modeller and Nest, a new method (Pfrag) for multiple template-based modeling is used, based on the segment-matching algorithm from Levitt's SegMod program. Our results show that all programs can produce multi-template models better than any of the single-template models, but a large part of the improvement is simply due to extension of the models. Most of the remaining improved cases were produced by Modeller. The most important factor is the existence of high-quality single-sequence input alignments. Because of the existence of models that are worse than any of the top single-template models, the average model quality does not improve significantly. However, by ranking models with a model quality assessment program such as ProQ, the average quality is improved by approximately 5% in the CASP7 test set.     

The pseudotympanum in the Gymnotiformes (Teleostei,Ostariophysi, Otophysi): homology and evolution of a previously unexplored system in Neotropical electric fishes

The pseudotympanum, a reduction of the hypaxialis muscle in the body wall lateral to the anterior portion of the swim bladder, is unique to some members of the Otophysi, the largest clade of primarily freshwater fishes. Prior studies documented the presence of the pseudotympanum in a number of groups within the Characiformes, to a lesser extent in subunits of the Siluriformes, and in a few taxa of the Cypriniformes, but with only one record of the aperture in the Gymnotiformes – the Neotropical electric fishes. Surveys across the five families and c. two‐thirds of the genera in the Gymnotiformes revealed the occurrence of a pseudotympanum in all examined taxa. Pronounced variation in the expanse and details of the morphology of the opening and its relationship to the anterior elements of the axial skeleton was documented amongst Neotropical electric fish. Details of pseudotympanal morphology in the different families of the Gymnotiformes are described and the variation evaluated within the context of the two alternative hypotheses of higher‐level phylogenetic relationships within the order. © 2015 The Linnean Society of London     

Xylella fastidiosa comparative genomic database is an information resource to explore the annotation, genomic features, and biology of different strains

The Xylella fastidiosa comparative genomic database is a scientific resource with the aim to provide a user-friendly interface for accessing high-quality manually curated genomic annotation and comparative sequence analysis, as well as for identifying and mapping prophage-like elements, a marked feature of Xylella genomes. Here we describe a database and tools for exploring the biology of this important plant pathogen. The hallmarks of this database are the high quality genomic annotation, the functional and comparative genomic analysis and the identification and mapping of prophage-like elements. It is available from web site http://www.xylella.lncc.br.     

Therapeutic perspectives in HIV-1 infection from recent advances in HIV-1 pathogenesis: it is time to move on

The recent availability of highly active antiretroviral combination therapy (HAART) has significantly influenced the natural history of the infection, delaying the progression to overt AIDS and prolonging survival. At the same time, the increasing knowledge on the pathogenic mechanisms an the use of HAART have challenged the most widely accepted theories about HIV-1 disease, in particular about the feasibility to eradicate the virus. This review will outline what is and what is not achievable by HAART, and will discuss new concepts in the immunopathogenesis of HIV-1 infection that provide the rationale for the design of new therapeutic approaches in the management of HIV-1 disease and AIDS.     

Redescription ofDiaphanosoma volzi Stingelin, 1905 (Crustacea: Daphniiformes: Sididae) with remarks on comparative morphology,biology and geographical distribution

The rareDiaphanosoma volzi Stingelin, long confused withD. sarsi Richard andD. celebensis Stingelin, is redescribed, based on extensive material of parthenogenetic and gamogenetic females and males from Thailand. Indonesia, Malaysia, Philippines, India, Australia and Sudan, and a study of intra- and interpopulation morphological variability. Compared with otherDiaphanosoma species, it shows the greatest degree of oligomerization and specialization. It is widely distributed in the tropics and subtropics of the Eastern hemisphere, and mostly occurs in shallow, vegetated habitats.     

Integrative omics - from data to biology

Introduction: Multi-omic approaches are promising a broader view on cellular processes and a deeper understanding of biological systems. with strongly improved high-throughput methods the amounts of data generated have become huge, and their handling challenging.

Area Covered: New bioinformatic tools and pipelines for the integration of data from different omics disciplines continue to emerge, and will support scientists to reliably interpret data in the context of biological processes. comprehensive data integration strategies will fundamentally improve systems biology and systems medicine. to present recent developments of integrative omics, the göttingen proteomics forum (gpf) organized its 6th symposium on the 23rd of november 2017, as part of a series of regular gpf symposia. more than 140 scientists attended the event that highlighted the challenges and opportunities but also the caveats of integrating data from different omics disciplines.

Expert commentary: The continuous exponential growth in omics data require similar development in software solutions for handling this challenge. Integrative omics tools offer the chance to handle this challenge but profound investigations and coordinated efforts are required to boost this field.     

'Type' in morphology and phylogeny

Comparative morphologists, developmental biologists, as well as paleontologists, recognize the existence of 'types,' or 'Baupl?ne,' in nature, marked out by a certain 'sameness' of structure that prevails through all variation in shape and function. The 'sameness' that marks out 'types' is one of structural correspondence (topology and connectivity), which is believed to be causally rooted in ontogeny (developmental constraints). In an evolutionary context, the structural relations that mark out 'types' are explained as relations of homology. The use of concepts such as 'type' or 'Bauplan' has been criticized from a 'populational thinking' point of view as being incompatible with current evolutionary theory. The present article explores the contrasting viewpoints, and concludes that current evolutionary theory can accommodate the concept of a 'type' in the sense of a 'homeostatic property cluster natural kind.' The 'homeostatic property cluster natural kind' is a nonessentialistic concept that allows the kind to be historically delimited.