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Proximodistal patterning of the limb: insights from evolutionary morphology.

There is an active debate about how skeletal elements are encoded along the proximodistal (PD) axis of the developing limb. Our aim here is to see whether consideration of the evolutionary morphology of the limb can contribute to our understanding of patterning mechanisms. Of special interest in this context are animals showing reiterated skeletal elements along the PD axis (e.g., dolphins and plesiosaurs with hyperphalangy). We build on previous hypotheses to propose a two-step model of PD patterning in which specification of broad domains in the early limb bud is distinct from subsequent processes that divides an initial anlage into a segmental pattern to yield individual skeletal elements. This model overcomes a major evolutionary problem with the progress zone model, which has not previously been noted: pleiotropy. Parallels with other developmental systems are briefly discussed.

Animals↗

Latent homologues for the neural crest as an evolutionary novelty.

The neural crest is a craniate synapomorphy and a bona fide evolutionary novelty. Recently, researchers considering intriguingly similar patterns of gene expression, cell behaviors, and embryogenetic processes in noncraniate deuterostomes have suggested that cephalochordates, urochordates, and echinoderms or their ancestors might have possessed cells that were precursors to the neural crest or its constituent cells. To emphasize the caution with which similarities at genetic, cellular, or embryological levels should be interpreted as substantiations for cell, germ layer, or tissue homologies, we present and evaluate additional tantalizing evidence that could be considered as documenting neural crest precursors in precraniates. Furthermore, we propose an evolutionary context--latent homologue--within which these data should be interpreted.

Animals↗

Evolutionary basis of parallelism in North American scincid lizards.

This study uses a phylogenetic framework to explore the causes of parallelism in two North American scincid lizard assemblages: the skiltonianus and fasciatus species groups of the genus Plestiodon. Each group consists of several closely related species with conserved neonate morphology; features that distinguish species become accentuated during ontogeny, and these differences often resemble different endpoints along a developmental continuum. This continuum is believed to be an expression of the ancestral ontogeny, and has led to the hypothesis that evolutionary change in development has generated much of the observed morphological diversity. However, progress on understanding these mechanisms is limited by a lack of well-supported phylogenetic data for the fasciatus group, and for Plestiodon in general. Recent phylogenetic studies on the skiltonianus group have revealed previously undetected cases of parallelism, and raise the possibility that similar cases have yet to be discovered in the fasciatus group. Here, I estimate a phylogeny to test the monophyly of the fasciatus group and infer its relationship with other North American Plestiodon using 2537 bp from six mtDNA genes. I use the phylogeny to reconstruct the mode (graduated vs. punctuated) and direction of body size evolution, to map the evolution of two predominant color morphs, and to test whether size and color pattern evolve concertedly. The results show that the morphotypes of the traditional fasciatus group constitute good species, but that the species group is rendered paraphyletic by several geographically overlapping species that deviate from the fasciatus-like ontogeny. Body size evolution has occurred gradually and bi-directionally, and shifts to large body size have been consistently associated with the loss of the striped color pattern during ontogeny. I show that parallelism, a lack of rigorous phylogenetic analysis, and a reliance on shared ontogenetic features for predicting phylogenetic relatedness, has misled the traditional systematics of these lizards, but that general ideas concerning the role of development in their morphological evolution remain supported. I close by proposing that the processes influencing repeated phyletic patterns in the skiltonianus and fasciatus groups represent adherence to an ancestral ground state, and discuss the importance of using phylogenies for the initial characterization of evolutionary changes in development.

Animals↗

Evolutionary origins of Hsp90 chaperones and a deep paralogy in their bacterial ancestors.

The 82-90 kD family of molecular chaperone proteins has homologs in eukaryotes (Hsp90) and many eubacteria (HtpG) but not in Archaebacteria. We used representatives of all four different eukaryotic paralogs (cytosolic, endoplasmic reticulum (ER), chloroplast, mitochondrial) together with numerous eubacterial HtpG proteins for phylogenetic analyses to investigate their evolutionary origins. Our trees confirm that none of the organellar Hsp90s derives from the endosymbionts of early eukaryotes. Contrary to previous suggestions of distant origins through lateral gene transfer (LGT) all eukaryote Hsp90s are related to Gram-positive eubacterial HtpG proteins. The nucleocytosolic, ER and chloroplast Hsp90 paralogs are clearly mutually related. The origin of mitochondrial Hsp90 is more obscure, as these sequences are deeply nested within eubacteria. Our trees also reveal a deep split within eubacteria into a group of mainly long-branching sequences (including the eukaryote mitochondrial Hsp90s) and another group comprising exclusively short-branching HtpG proteins, from which the cytosolic/ER versions probably arose. Both versions are present in several eubacterial phyla, suggesting gene duplication very early in eubacterial evolution and multiple independent losses thereafter. We identified one probable case of LGT within eubacteria. However, multiple losses can simply explain the evolutionary pattern of the eubacterial HtpG paralogs and predominate over LGT. We suggest that the actinobacterial ancestor of eukaryotes harbored genes for both eubacterial HtpG paralogs, as the actinobacterium Streptomyces coelicolor still does; one could have given rise to the mitochondrial Hsp90 and the other, following another duplication event in the ancestral eukaryote, to the cytosolic and ER Hsp90 homologs.

Animals↗

At least four MHC class I genes are transcribed in the horse: phylogenetic analysis suggests an unusual evolutionary history for the MHC in this species.

Nineteen horse MHC class I specificities have been serologically identified previously at a single locus (ELA-A), and two other specificities appear to be coded at other loci. Biochemical studies indicate that there are at least two expressed loci. In order to establish the number of transcribed horse MHC class I genes, we made a cDNA library from a heterozygous animal (ELA-A3/A7), and screened for positive clones using a bovine class I probe. More than 200 class I clones were isolated in this way, and so far seven unique full length sequences have been identified. All of the sequences are predicted to code for surface expressed, functional molecules. The number of different sequences identified demonstrate that at least four genes are transcribed, although variations in transmembrane length (which is generally conserved in class I loci) suggest that five genes could be represented. Evolutionary analysis of these sequences (and two additional sequences known to represent different horse class I loci) reveals no firm relationships, such that the division between the different loci cannot be discerned. These results suggest an unusual evolutionary history for the horse MHC, the precise nature of which may be revealed only following further cross-species comparisons.

Amino Acid Sequence↗

Structural diversity of eukaryotic small subunit ribosomal RNAs. Evolutionary implications.

The phylogenetic diversity of the eukaryotic kingdom was assessed by comparing the structural and evolutionary diversity of 18-20S ribosomal RNA genes. The coding regions for cytoplasmic small subunit ribosomal RNA genes vary in length from 1753 to 2305 nucleotides, and they appear to be evolutionary mosaics in which highly and partially conserved sequences are interspersed among regions that display very high rates of genetic drift. Structural similarities between these gene sequences were used to establish a phylogenetic framework for the eukaryotes. The extent of sequence variation within the eukaryotes exceeds that displayed within the eubacterial or archaebacterial lines of descent. The kinetoplastids and euglenoids represent the earliest branchings among the eukaryotes. These branchings preceded the divergence of lineages leading to the slime molds and apicomplexans and far antedate a radiative period that gave rise to the plants, animals, fungi, and other protists.

Animals↗

Freud: the first evolutionary psychologist?

An evolutionary perspective on attachment theory and psychoanalytic theory brings these two fields together in interesting ways. Application of the evolutionary principle of parent-offspring conflict to attachment theory suggests that attachment styles represent context-sensitive, evolved (adaptive) behaviors. In addition, an emphasis on offspring counter-strategies to adult reproductive strategies leads to consideration of attachment styles as overt manifestations of psychodynamic mediating processes, including the defense mechanisms of repression and reaction formation.

Adult↗

Evolutionary psychopathology: why isn't the mind designed better than it is?

This paper outlines some of the basic issues in evolutionary thinking when applied to psychopathology. It aims to give some background and context to the papers that follow. Particular attention is given to the way evolution is constrained by previous design features, the importance of trade offs between advantages and disadvantages of different traits, the way social dilemmas and conflicts have shaped the evolution of human motivational systems, and the psychology that supports them. The paper concludes with a plea to avoid thinking of evolutionary approaches as a reductionist science, when the focus needs to be on the way evolution theory informs the study of the social-biological interface.

Biological Evolution↗

Emotional disorders in evolutionary perspective.

Understanding emotional disorders requires understanding the evolutionary origins and functions of normal emotions. They are special states, shaped by natural selection to adjust various aspects of the organism in ways that have tended to give a selective advantage in the face of the adaptive challenges characteristic of a particular kind of situation. They are designed to maximize reproductive success, not happiness. Negative emotions such as anxiety and low mood are not disorders, but, like the capacity for pain, evolved defences. Excessive anxiety or low mood is abnormal, but we will not have confidence about what is excessive until we understand their functions better than we do. Emotional disorders arise often from social emotions because of the conflicts inherent in social life, and because of the strategic advantages of demonstrating commitments to follow through on threats and promises. An evolutionary understanding of individuals in terms of their relationship strategies and the social emotions offers great promise for psychotherapists.

Affective Symptoms↗

Eco-Evolutionary Genomics Reveal Mountain Range-Specific Adaptation and Intraspecific Variation in Vulnerability to Climate Change of Alpine Endemics.

Alpine plants restricted to rocky habitats exhibit intraspecific diversification due to range fragmentation during Holocene warming, complicating predictions of their climate vulnerability. A lack of understanding of eco-evolutionary mechanisms driving their response to climate change results in ineffective conservation efforts. To uncover the genomic basis of their diversification and explain spatial patterns of their vulnerability, we combine landscape genomics and species distribution modelling. Our model, the Campanula lehmanniana complex, occurs in three distinct central Asian mountain ranges, considered both a biodiversity hotspot and a vascular plant diversity darkspot. Genome-environment association confirmed the adaptive basis of intraspecific diversification, driven by numerous loci of small effect. Genomic and ecological data indicate mountain range-specific climate sensitivity driven by altitude, temperature and precipitation. The cold-dry adapted group from Zeravshan-Hissar Mts will face niche decline but show a higher degree of preadaptation to future climate, while the temperate-humid group from Tian Shan shows an opposite response, with a higher risk of maladaptation despite predicted niche expansion. Maladapted populations at northern margins may require an influx of adaptive variation to cope with predicted changes. However, limited landscape connectivity between island-like habitats, combined with long migration distances required to minimise genotype-environment disruption, highlights the role of human-assisted migration in enabling evolutionary rescue. These results underscore the need to facilitate gene flow from pre- to maladapted populations and the importance of population-specific approaches to inform effective conservation strategies in heterogeneous mountain ecosystems. The results may be relevant to numerous Central Asian mountain species that show similar phylogeographic patterns.

Climate Change↗

Evolutionary legacy of the "living fossil" genus Parrotia (Hamamelidaceae): genomic insights into species divergence and polygenic adaptation.

Despite their long evolutionary history, the genomic basis of adaptation and speciation in "living fossil" plants remain largely unexplored. Parrotia, a Tertiary relict tree genus with two extant species, P. subaequalis and P. persica, exhibits a disjunct distribution between East Asia and West Asia. Here, we present the first chromosome-level assemblies for both species, confirmed their sibling relationship, and dated the speciation event to the early Miocene. The recent proliferation of long-terminal repeat retrotransposons has driven the genome expansion in P. subaequalis. We detected widespread heterogeneous genomic differentiation between species. Extensive signals of divergent selection, local adaptation, and elevated Ka/Ks ratios in Parrotia indicate that this genus has undergone adaptive evolution in distinct refugia, challenging the notion of it as an "evolutionary dead end". Our findings provide new insights into the genomic evolution, environmental adaptation, and speciation of this "living fossil" tree genus.

Genome, Plant↗

Assembling genomes of non-model plants: A case study with evolutionary insights from Ranunculus (Ranunculaceae).

Whereas genome sequencing and assembly technologies are improving, cost can still be prohibitive for plant species with large, complex genomes. As a consequence, genomics work on some taxa in evolutionarily pivotal positions in the vascular plant tree of life has been hampered. The species-rich genus Ranunculus (Ranunculaceae) is an important angiosperm group for the study of polyploidy, apomixis, and reticulate evolution. However, neither mitochondrial nor high-quality nuclear genome sequences are available. This limits phylogenomic, functional, and taxonomic analyses thus far. Here, we tested Illumina short-read, Oxford Nanopore Technology (ONT) and PacBio (HiFi) long-read, and hybrid-read assembly strategies. We sequenced the diploid progenitor species R. cassubicifolius (R. auricomus species complex) and selected the best assemblies in terms of completeness, contiguity, and quality scores. We first assembled the plastome (156 kbp, 85 genes) and mitogenome (1.18 Mbp, 40 genes) sequences using Illumina and Illumina-PacBio-hybrid strategies, respectively. We also present an updated plastome and the first mitogenome phylogeny of Ranunculaceae, including studies of gene loss (e.g., infA, ycf15, or rps) with evolutionary implications. For the nuclear genome sequence, we favored a PacBio-based assembly polished three times with filtered short reads and subsequently scaffolded into eight pseudochromosomes by chromatin conformation data (Hi-C). We obtained a haploid genome sequence of 2.69 Gbp, with 94.1% complete BUSCO genes found and 35 482 annotated genes, and inferred ancient gene duplications compared to existing Ranunculales genomes. The genomic information presented here will enable advanced evolutionary-functional analyses for the species complex, but also for the genus and beyond Ranunculaceae.

Ranunculus↗

Nuclear DNA of plastid origin (NUPTs), neglected driver of genome variation and evolutionary innovation.

Plant nuclear genomes contain a variable, though typically minor, fraction of DNA sequences of plastid origin known as NUPTs. Unlike the massive transfer of DNA and genes from the proto-organelle genome to the nucleus that occurred during the endosymbiotic event that gave rise to plastids, the formation of NUPTs is an ongoing process that does not imply concomitant DNA loss. Although NUPTs are generally considered to be potentially deleterious insertions that are continuously generated and rapidly eliminated at near-constant turnover rates, accumulating evidence reveals alternative evolutionary trajectories. In this review, we discuss recent findings that highlight the episodic formation of NUPTs, their subsequent proliferation, and their eventual long-term fixation within the nuclear genome. We also explore their non-random spatial association with specific genomic elements. NUPTs show preferential overlap with specific superfamilies of transposable elements, which may facilitate their proliferation and dispersal throughout the nuclear genome. Regarding protein-coding genes, the contribution of NUPTs varies among species. In contrast, NUPTs are found to be consistently enriched among certain classes of non-coding RNA genes, notably rRNA, tRNA, and specific regulatory RNA families, suggesting that they are involved in the evolution of gene regulation and translational machinery. Overall, these findings underscore the unexpected complexity of the mechanisms underlying NUPT formation and support the idea that they are a significant source of genome variation and evolutionary innovation. Further research is necessary to fully elucidate the mechanisms underlying NUPT formation, as well as to determine their potential adaptive significance in plant genome evolution.

Plastids↗

Quantitative evolutionary design.

The field of quantitative evolutionary design uses evolutionary reasoning (in terms of natural selection and ultimate causation) to understand the magnitudes of biological reserve capacities, i.e. excesses of capacities over natural loads. Ratios of capacities to loads, defined as safety factors, fall in the range 1.2-10 for most engineered and biological components, even though engineered safety factors are specified intentionally by humans while biological safety factors arise through natural selection. Familiar examples of engineered safety factors include those of buildings, bridges and elevators (lifts), while biological examples include factors of bones and other structural elements, of enzymes and transporters, and of organ metabolic performances. Safety factors serve to minimize the overlap zone (resulting in performance failure) between the low tail of capacity distributions and the high tail of load distributions. Safety factors increase with coefficients of variation of load and capacity, with capacity deterioration with time, and with cost of failure, and decrease with costs of initial construction, maintenance, operation, and opportunity. Adaptive regulation of many biological systems involves capacity increases with increasing load; several quantitative examples suggest sublinear increases, such that safety factors decrease towards 1.0. Unsolved questions include safety factors of series systems, parallel or branched pathways, elements with multiple functions, enzyme reaction chains, and equilibrium enzymes. The modest sizes of safety factors imply the existence of costs that penalize excess capacities. Those costs are likely to involve wasted energy or space for large or expensive components, but opportunity costs of wasted space at the molecular level for minor components.

Animals↗

Bayesian inference of phylogeny and its impact on evolutionary biology.

As a discipline, phylogenetics is becoming transformed by a flood of molecular data. These data allow broad questions to be asked about the history of life, but also present difficult statistical and computational problems. Bayesian inference of phylogeny brings a new perspective to a number of outstanding issues in evolutionary biology, including the analysis of large phylogenetic trees and complex evolutionary models and the detection of the footprint of natural selection in DNA sequences.

Algorithms↗

Unifying the epidemiological and evolutionary dynamics of pathogens.

A key priority for infectious disease research is to clarify how pathogen genetic variation, modulated by host immunity, transmission bottlenecks, and epidemic dynamics, determines the wide variety of pathogen phylogenies observed at scales that range from individual host to population. We call the melding of immunodynamics, epidemiology, and evolutionary biology required to achieve this synthesis pathogen "phylodynamics." We introduce a phylodynamic framework for the dissection of dynamic forces that determine the diversity of epidemiological and phylogenetic patterns observed in RNA viruses of vertebrates. A central pillar of this model is the Evolutionary Infectivity Profile, which captures the relationship between immune selection and pathogen transmission.

Adaptation, Physiological↗

Evolutionary dynamics of biological games.

Darwinian dynamics based on mutation and selection form the core of mathematical models for adaptation and coevolution of biological populations. The evolutionary outcome is often not a fitness-maximizing equilibrium but can include oscillations and chaos. For studying frequency-dependent selection, game-theoretic arguments are more appropriate than optimization algorithms. Replicator and adaptive dynamics describe short- and long-term evolution in phenotype space and have found applications ranging from animal behavior and ecology to speciation, macroevolution, and human language. Evolutionary game theory is an essential component of a mathematical and computational approach to biology.

Adaptation, Biological↗

The evolutionary origin of cooperators and defectors.

Coexistence of cooperators and defectors is common in nature, yet the evolutionary origin of such social diversification is unclear. Many models have been studied on the basis of the assumption that benefits of cooperative acts only accrue to others. Here, we analyze the continuous snowdrift game, in which cooperative investments are costly but yield benefits to others as well as to the cooperator. Adaptive dynamics of investment levels often result in evolutionary diversification from initially uniform populations to a stable state in which cooperators making large investments coexist with defectors who invest very little. Thus, when individuals benefit from their own actions, large asymmetries in cooperative investments can evolve.

Altruism↗