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Ultrastructure of the post-corpus of Zeldia punctata (Cephalobina) for analysis of the evolutionary framework of nematodes related to Caenorhabditis elegans (Rhabditina).

The ultrastructure of the post-corpus of Zeldia punctata (Cephalobina) was compared with previous observations of Caenorhabditis elegans (Rhabditina) and Diplenteron sp. (Diplogastrina) with the goal of interpreting the morphological evolution of the feeding structures in the Secernentea. The post-corpus of Z. punctata consists of six marginal, 13 muscle, five gland and seven nerve cells. The most anterior of four layers of muscle cells consists of six mononucleate cells in Z. punctata. The homologous layer in C. elegans and Diplenteron consists of three binucleate cells, suggesting a unique derived character (synapomorphy) shared between the Rhabditina and Diplogastrina. Contrary to Diplenteron sp. where we observed three oesophageal glands, Z. punctata and C. elegans have five oesophageal glands. We question this shared character as reflecting a common evolution between the Cephalobina and Rhabditina, because there are strong arguments for functional (adaptive) convergence of the five glands in these bacterial feeders. Convergence is further suggested by the mosaic distribution of three versus five glands throughout the Nemata; this distribution creates difficulties in establishing character polarity. Although morphological data are often laborious to recover and interpret, we nevertheless view 'reciprocal illumination' between molecular and morphological characters as the most promising and robust process for reconstructing the evolution of the Secernentea and its feeding structures.

Animals↗

Hypothesis testing in evolutionary developmental biology: a case study from insect wings.

Developmental data have the potential to give novel insights into morphological evolution. Because developmental data are time-consuming to obtain, support for hypotheses often rests on data from only a few distantly related species. Similarities between these distantly related species are parsimoniously inferred to represent ancestral aspects of development. However, with limited taxon sampling, ancestral similarities in developmental patterning can be difficult to distinguish from similarities that result from convergent co-option of developmental networks, which appears to be common in developmental evolution. Using a case study from insect wings, we discuss how these competing explanations for similarity can be evaluated. Two kinds of developmental data have recently been used to support the hypothesis that insect wings evolved by modification of limb branches that were present in ancestral arthropods. This support rests on the assumption that aspects of wing development in Drosophila, including similarities to crustacean epipod patterning, are ancestral for winged insects. Testing this assumption requires comparisons of wing development in Drosophila and other winged insects. Here we review data that bear on this assumption, including new data on the functions of wingless and decapentaplegic during appendage allocation in the red flour beetle Tribolium castaneum.

Animals↗

Temperature and Pressure Shaped the Evolution of Antifreeze Proteins in Polar and Deep Sea Zoarcoid Fishes.

Antifreeze proteins (AFPs) have enabled teleost fishes to repeatedly colonize polar seas. Four AFP types have convergently evolved in several fish lineages. AFPs inhibit ice crystal growth and lower tissue freezing point. In lineages with AFPs, species inhabiting colder environments may possess more AFP copies. Elucidating how differences in AFP copy number evolve is challenging due to the genes' tandem array structure and consequently poor resolution of these repetitive regions. Here, we explore the evolution of type III AFPs (AFP III) in the globally distributed suborder Zoarcoidei, leveraging six new long-read genome assemblies. Zoarcoidei has fewer genomic resources relative to other polar fish clades while it is one of the few groups of fishes adapted to both the Arctic and Southern Oceans. Combining these new assemblies with additional long-read genomes available for Zoarcoidei, we conducted a comprehensive phylogenetic test of AFP III evolution and modeled the effects of thermal habitat and depth on AFP III gene family evolution. We confirm a single origin of AFP III via neofunctionalization of the enzyme sialic acid synthase B. We also show that AFP copy number increased under low temperature but decreased with depth, potentially because pressure lowers freezing point. Associations between the environment and AFP III copy number were driven by duplications of paralogs that were translocated out of the ancestral locus at which AFP III arose. Our results reveal novel environmental effects on AFP evolution and demonstrate the value of high-quality genomic resources for studying how structural genomic variation shapes convergent adaptation.

Animals↗

Alu elements support independent origin of prosimian, platyrrhine, and catarrhine Mhc-DRB genes.

The primate major histocompatibility complex (Mhc) genes fall into two classes and each of the classes into several families. Of the class II families, the DRB family has a long and complex evolutionary history marked by gene turnover, rearrangement, and molecular convergence. Because the history is not easily decipherable from sequences alone, Alu element insertions were used as cladistic markers to support the surmised phylogenetic relationships among the DRB genes. Intron 1 segments of 24 DRB genes from five platyrrhine species and five DRB genes from three prosimian species were amplified by PCR and cloned, and the amplification products were sequenced or PCR-typed for Alu repeats. Three Alu elements were identified in the platyrrhine and four in the prosimian DRB genes. One of the platyrrhine elements (Alu50J) is also found in the Catarrhini, whereas the other two (Alu62Sc, Alu63Sc) are restricted to the New World monkeys. Similarly, the four prosimian elements are found only in this taxon. This distribution of Alu elements is consistent with the phylogeny of the DRB genes as determined from their intron 1 sequences in an earlier and the present study. It contradicts the exon 2-based phylogeny and thus corroborates the conclusion that the evolution of DRB exon 2 sequences is, to some extent, shaped by molecular convergence. Taken together, the data indicate that each of the assemblages of DRB genes in prosimians, platyrrhines, and catarrhines is derived from a separate ancestral gene.

Alu Elements↗

Parental Care: The Key to Understanding Endothermy and Other Convergent Features in Birds and Mammals.

Birds and mammals share a number of features that are remarkably similar but that have evolved independently. One of these characters, endothermy, has been suggested to have played a cardinal role in avian and mammalian evolution. I hypothesize that it is parental care, rather than endothermy, that is the key to understanding the amazing convergence between mammals and birds. Endothermy may have arisen as a consequence of selection for parental care because endothermy enables a parent to control incubation temperature. The remarkable ability of many birds and mammals to sustain vigorous exercise may also have arisen as a consequence of selection for parental care because provisioning of offspring often requires sustained vigorous exercise. Because extensive parental care encompasses a wide range of behaviors, morphology, and physiology, it may be a key innovation that accounts for the majority of convergent avian and mammalian characters.

Journal Article↗

Mosaic evolution of rodent B1 elements.

We have determined sequences of PCR-amplified B1 elements from hamster and rat (Myomorpha), chipmunk (Sciuromorpha), and guinea pig (Caviomorpha). Between three and six B1 subfamilies were found in these species. In the phylogenetic analysis B1 sequences of hamster, mouse, and rat clustered separately from those of chipmunk and those of guinea pig. This is consistent with an independent evolution of B1 elements in separate rodent lineages. We exclude the possibility of convergent mutations to explain certain diagnostic characters within the modern B1 quasi-dimers and view these elements as mosaic structures assembling preexisting mutations. Furthermore, the presence of Alu-like structural motifs supports the hypothesis of the monophyletic origin of Alu and B1 repeats, i.e., from a common 7SL RNA-derived retroposing monomeric element.

Animals↗

Properties of isolated systems in external fields.

We investigate evolution of an isolated system in an external field, and compare the ensemble averages of the response on successive constant internal energy surfaces to the ensemble averages of steady-state responses constrained to the same energy. We find that the two ensemble averages converge for sufficiently high energies, irrespective of the field strength and the initial energy from which the adiabatic evolution starts. This rule is satisfied for any phase-space distribution on the initial energy surface that can relax to equilibrium. At sufficiently high energies transport coefficients converge to their equilibrium values, because the effect of a constant field on the behavior of a system decreases with its temperature.

Journal Article↗

Evolution of the electronic structure of Be clusters.

Using a modified symbiotic genetic algorithm approach and many-body interatomic potential derived from first principles, we have calculated equilibrium geometries and binding energies of the ground-state and low-lying isomers of Be clusters containing up to 41 atoms. Molecular-dynamics study was also carried out to study the frequency of occurrence of the various geometrical isomers as these clusters are annealed during the simulation process. For a selected group of these clusters, higher-energy isomers were more often found than their ground-state structures due to large catchment areas. The accuracy of the above ground-state geometries and their corresponding binding energies were verified by carrying out separate ab initio calculations based on molecular-orbital approach and density-functional theory with generalized gradient approximation for exchange and correlation. The atomic orbitals were represented by a Gaussian 6-311G** basis, and the geometry optimization was carried out using the GAUSSIAN 98 code without any symmetry constraint. While the ground-state geometries and their corresponding binding energies obtained from ab initio calculations do not differ much from those obtained using the molecular-dynamics approach, the relative stability of the clusters and the energy gap between the highest occupied and the lowest unoccupied molecular orbitals show significant differences. The energy gaps, calculated using the density-functional theory, show distinct shell closure effects, namely, sharp drops in their values for Be clusters containing 2, 8, 20, 34, and 40 electrons. While these features may suggest that small Be clusters behave free-electron-like and, hence, are metallic, the evolution of the structure, binding energies, coordination numbers, and nearest-neighbor distances do not show any sign of convergence towards the bulk value. We also conclude that molecular-dynamics simulation based on many-body interatomic potentials may not always give the correct picture of the evolution of the structure and energetics of clusters although they may serve as a useful tool for obtaining starting geometries by efficiently searching a large part of the phase space.

Journal Article↗

Evolution of mechanoregulation of bone growth will lead to non-optimal bone phenotypes.

Mechanical forces acting on the bones during growth affect their final shape and strength. Mechanoregulation of bone growth may be recognized in embryogenesis, and also in the adaptation of the adult skeleton to changes in mechanical loading. Mechanoregulatory responses for tissues have arisen during evolution, but does evolution give rise to responses that produce optimal skeletal phenotypes? In this paper, we investigate the emergence of an optimal mechanoregulation response in a population. By combining equations describing long bone growth with a genetic algorithm to describe evolutionary change, we created a computational model to simulate the evolution of mechanoregulation in bone growth. A population of individuals is created where each individual is assigned a diploid gene set which controls the growth and remodelling of the bone. At maturity, each bone is assessed and its 'fitness' calculated; fitness is quantified as bone strength relative to bone mass. The simulation continues for many generations, and includes mutations and a varying environment. The genes present in the population are tracked and the evolution of parameters governing mechanoregulation is calculated. The results indicate that a population may converge to one bone growth algorithm but, more usually, a range of mechanoregulation algorithms for different individuals will persist after many generations. Even if the population converges to one mechanoregulation law, convergence to the 'optimum' bone was never found. Although many researchers propose that natural selection has pushed skeletal structure towards an optimum, our computational model suggests that this is unlikely to be the case.

Adaptation, Physiological↗

The theory of genotypic selection: predicting the direction of evolution as a consequence of G:U base pairing and the existence of non-pathogenic strains of HIV-1.

One aspect of the Theory of Genotypic Selection states that G:U base pairs are selected for function and structure. RNA secondary structures stabilised by G:U base pairs are suggested to be involved in the evolution of attenuated strains of polio virus. Asymmetric mutations (C-->U and A-->G) and convergent evolutionary solutions are explained as a direct consequence of G:U base pairing. The possibility of a naturally occurring attenuated strain of HIV-1 is predicted.

Base Composition↗

The hypocone as a key innovation in mammalian evolution.

The hypocone, a cusp added to the primitively triangular upper molar teeth of therian mammals, has evolved convergently > 20 times among mammals during the Cenozoic. Acquisition of the hypocone itself involves little phenotypic change, but subsequent diversification of groups possessing the hypocone may be greatly enhanced. Our analysis of the Cenozoic mammalian radiations, including the Recent fauna, shows that high species diversity of mammals with hypocones and association of the hypocone with herbivory strongly support recognition of the hypocone as a key innovation that has allowed invasion of, and diversification within, herbivorous adaptive zones. In contrast, mammals lacking hypocones show no marked increase in species diversity during the Cenozoic.

Animals↗

Tubulin sequence conservation.

Various aspects of the primary structure of tubulin are discussed and tubulin sequence data are compared. A hypothesis concerning the evolution of tubulin is also presented. The main points raised are: (1) Identical glycyl rich regions in alpha- and beta-tubulin, which are similar in both sequence and predicted secondary structure to a region in several nucleotide binding enzymes, may be involved in binding GTP. (2) Small regions of homology are present to actin, myosin and troponin T. These homologous regions may have the same function, resulting in a convergence of their sequences, or they may have arisen by a pathway of protein evolution which is still only very poorly understood. (3) The mutation rate between pig and chick brain tubulin is 0.22 PAMs (accepted point mutations/100 residues) per hundred million years, which is comparable to that of the histones. At an early time in its history, however, the tubulin heterodimer appears to have had a relatively high rate of mutation. This may have been during the evolution of the first eukaryotes.

Amino Acid Sequence↗

Evidence for multiple genetic forms with similar eyeless phenotypes in the blind cavefish, Astyanax mexicanus.

A diverse group of animals has adapted to caves and lost their eyes and pigmentation, but little is known about how these animals and their striking phenotypes have evolved. The teleost Astyanax mexicanus consists of an eyed epigean form (surface fish) and at least 29 different populations of eyeless hypogean forms (cavefish). Current alternative hypotheses suggest that adaptation to cave environments may have occurred either once or multiple times during the evolutionary history of this species. If the latter is true, the unique phenotypes of different cave-dwelling populations may result from convergence of form, and different genetic changes and developmental processes may have similar morphological consequences. Here we report an analysis of variation in the mitochondrial NADH dehydrogenase 2 (ND2) gene among different surface fish and cavefish populations. The results identify a minimum of two genetically distinctive cavefish lineages with similar eyeless phenotypes. The distinction between these divergent forms is supported by differences in the number of rib-bearing thoracic vertebrae in their axial skeletons. The geographic distribution of ND2 haplotypes is consistent with roles for multiple founder events and introgressive hybridization in the evolution of cave-related phenotypes. The existence of multiple genetic lineages makes A. mexicanus an excellent model to study convergence and the genes and developmental pathways involved in the evolution of the eye and pigment degeneration.

Animals↗

Evolution of eyes.

Seeing is important for most species and it has been a key selective advantage throughout evolution. Consequently, there is a remarkable diversity among types of eyes. Animals have converged on eight optical solutions for collecting and focusing light; in contrast, all eyes share the same molecular strategy for absorbing photons. Recent studies have identified similarities in the genetic information that is used in the development of eyes, leading to the hypothesis that distinctly different eye types might have had a monophyletic origin. Across many species, there is a remarkable continuity of the developmental genes that participate in the construction of similar--but not necessarily homologous--eyes.

Animals↗

Uses and limitations of mammalian fetal membranes and placenta for phylogenetic reconstruction.

Comparative studies on the development of the mammalian fetal membranes and placenta, and discussions of the possible evolutionary significance of these data, have been carried out for more than a century. Throughout much of this period, however, such data were considered in isolation from cranioskeletal and other biological findings, so that only rarely were placental features incorporated into comparative assessments of mammalian evolution. Evolutionary interpretations of the fetal membrane evidence were commonly based on the concept of overall similarity, and most placentologists were strongly influenced by the widely accepted hypothesis that living "insectivores" were closely related to the ancestral eutherian mammals. Mossman's emphasis on the use of all available developmental data on the fetal membranes and placenta, rather than simply the nature of the definitive placenta, marked a turning point in the history of using such findings for assessing eutherian mammalian evolution. Recent advances in the methodology of phylogenetic inference, with emphasis on evaluating convergence versus homology, and on distinguishing between relatively primitive versus derived homologous similarities shared by two or more taxa, have provided a sound framework for the phylogenetic analysis of fetal membrane data. Character analysis of mammalian fetal membrane traits provides a mechanism for testing phylogenetic hypotheses formulated by cranioskeletal features. It is emphasized in this study that fetal membrane data have their greatest input into assessing mammalian phylogeny when they are evaluated in conjunction with the available cranioskeletal and molecular evidence.

Animals↗

Reviving the superorganism.

Individuals become functionally organized to survive and reproduce in their environments by the process of natural selection. The question of whether larger units such as groups and communities can possess similar properties of functional organization, and therefore be regarded as "superorganisms", has a long history in biological thought. Modern evolutionary biology has rejected the concept of superorganisms, explaining virtually all adaptations at the individual or gene level. We criticize the modern literature on three counts. First, individual selection in its strong form is founded on a logical contradiction, in which genes-in-individuals are treated differently than individuals-in-groups or species-in-communities. Imposing consistency clearly shows that groups and communities can be organisms in the same sense that individuals are. Furthermore, superorganisms are more than just a theoretical possibility and actually exist in nature. Second, the view that genes are the "ultimate" unit of selection is irrelevant to the question of functional organization. Third, modern evolutionary biology includes numerous conceptual frameworks for analyzing evolution in structured populations. These frameworks should be regarded as different ways of analyzing a common process which, to be correct, must converge on the same conclusions. Unfortunately, evolutionists frequently regard them as competing theories that invoke different mechanisms, such that if one is "right" the others must be "wrong". The problem of multiple frameworks is aggravated by the fact that major terms, such as "units of selection", are defined differently within each framework, yet many evolutionists who use one framework to argue against another assume shared meanings. We suggest that focusing on the concept of organism will help dispell this fog of semantic confusion, allowing all frameworks to converge on the same conclusions regarding units of functional organization.

Animals↗

Dispersion of human Y chromosome haplotypes based on five microsatellites in global populations.

We have analyzed five microsatellite loci from the nonrecombining portion of the human Y chromosome in 15 diverse human populations to evaluate their usefulness in the reconstruction of human evolution and early male migrations. The results show that, in general, most populations have the same set of the most frequent alleles at these loci. Hypothetical ancestral haplotypes, reconstructed on the basis of these alleles and their close derivatives, are shared by multiple populations across racial and geographical boundaries. A network of the observed haplotypes is characterized by a lack of clustering of geographically proximal populations. In spite of this, few distinct clusters of closely related populations emerged in the network, which are associated with population-specific alleles. A tree based on allele frequencies also shows similar results. Lack of haplotypic structure associated with the presumed ancestral haplotypes consisting of individuals from almost all populations indicate a recent common ancestry and/or extensive male migration during human evolutionary history. The convergent nature of microsatellite mutation confounds population relationships. Optimum resolution of Y chromosome evolution will require the use of additional microsatellite loci and diallelic genetic markers with lower mutation rates.

Alleles↗

Evolution of a total synthesis of (-)-kendomycin exploiting a Petasis-Ferrier rearrangement/ring-closing olefin metathesis strategy.

A convergent stereocontrolled total synthesis of (-)-kendomycin (1) has been achieved. The synthesis proceeds with a longest linear sequence of 21 steps, beginning with commercially available 2,4-dimethoxy-3-methylbenzaldehyde (12). Highlights of the synthesis include an effective Petasis-Ferrier union/rearrangement tactic to construct the sterically encumbered tetrahydropyran ring, a ring-closing metathesis to generate the C(4a-13-20a) macrocycle, an effective epoxidation/deoxygenation sequence to isomerize the C(13,14) olefin, and a biomimetic quinone-methide-lactol assembly to complete the synthesis.

Alkenes↗