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Evolution of the vertebral formulae in mammals: a perspective on developmental constraints.

Developmental constraints refer to biases that limit phenotypic changes during evolution. To examine the contribution of developmental constraints in the evolution of vertebrate morphology, we analyzed the distribution pattern of mammalian vertebral formulae. Data on mammalian vertebral formulae were collected from the Descriptive Catalogue of the Osteological Series Contained in the Museum of the Royal College of Surgeons of England by Richard Owen (1853) and were plotted onto the most reliable mammalian phylogenetic tree based on recent molecular studies. In addition to the number of cervical vertebrae that is almost fixed to 7, we found that the number of thoracolumbar vertebrae tends to be 19 in many groups of mammals. Since fidelity of the number of thoracolumbar vertebrae was also completely maintained in Monotremata and Marsupialia, we presumed that thoracolumbar vertebral number as well as cervical vertebral number might have been fixed in the primitive mammalian lineage. On the basis of primitive vertebral formulae, we could clarify the polarity of evolution and identify several deviations from the primitive states during the mammalian evolution. The changes in the vertebral formulae in eutherian mammals seem to be lineage-specific, such that most species in Carnivora have 20 instead of 19 thoracolumbar vertebrae. Because such lineage-specific vertebral formulae contrast with the estimated distribution pattern on the assumption of evolution only through the selective pressure, we concluded that developmental constraints played an important role in the evolution of mammalian vertebral formulae.

Animals↗

Ancient Wings: animating the evolution of butterfly wing patterns.

Character optimization methods can be used to reconstruct ancestral states at the internal nodes of phylogenetic trees. However, seldom are these ancestral states visualized collectively. Ancient Wings is a computer program that provides a novel method of visualizing the evolution of several morphological traits simultaneously. It allows users to visualize how the ventral hindwing pattern of 54 butterflies in the genus Bicyclus may have changed over time. By clicking on each of the nodes within the evolutionary tree, the user can see an animation of how wing size, eyespot size, and eyespot position relative the wing margin, have putatively evolved as a collective whole. Ancient Wings may be used as a pedagogical device as well as a research tool for hypothesis-generation in the fields of evolutionary, ecological, and developmental biology.

Animals↗

Ancestors and variants: tales from the cryptic.

Those who work at the interface of development and evolution are united by the conviction that developmental comparisons can shed light on both the evolution of specific morphologies and the macroevolutionary process itself. In practice, however, the field comprises a diversity of approaches. As the field grows and practitioners attempt to digest a growing mountain of comparative data, the various approaches of "Evo Devo" have themselves evolved. A meeting organized by the authors and held at the University of Chicago in the Spring of 1999 illustrated some of these changes. This review will draw on its content to discuss recent developments in two areas: the reconstruction of common ancestors and the developmental basis of evolutionary change.

Animals↗

Gene-balanced duplications, like tetraploidy, provide predictable drive to increase morphological complexity.

Controversy surrounds the apparent rising maximums of morphological complexity during eukaryotic evolution, with organisms increasing the number and nestedness of developmental areas as evidenced by morphological elaborations reflecting area boundaries. No "predictable drive" to increase this sort of complexity has been reported. Recent genetic data and theory in the general area of gene dosage effects has engendered a robust "gene balance hypothesis," with a theoretical base that makes specific predictions as to gene content changes following different types of gene duplication. Genomic data from both chordate and angiosperm genomes fit these predictions: Each type of duplication provides a one-way injection of a biased set of genes into the gene pool. Tetraploidies and balanced segments inject bias for those genes whose products are the subunits of the most complex biological machines or cascades, like transcription factors (TFs) and proteasome core proteins. Most duplicate genes are removed after tetraploidy. Genic balance is maintained by not removing those genes that are dose-sensitive, which tends to leave duplicate "functional modules" as the indirect products (spandrels) of purifying selection. Functional modules are the likely precursors of coadapted gene complexes, a unit of natural selection. The result is a predictable drive mechanism where "drive" is used rigorously, as in "meiotic drive." Rising morphological gain is expected given a supply of duplicate functional modules. All flowering plants have survived at least three large-scale duplications/diploidizations over the last 300 million years (Myr). An equivalent period of tetraploidy and body plan evolution may have ended for animals 500 million years ago (Mya). We argue that "balanced gene drive" is a sufficient explanation for the trend that the maximums of morphological complexity have gone up, and not down, in both plant and animal eukaryotic lineages.

Animals↗

Mechanisms of pattern formation in development and evolution.

We present a classification of developmental mechanisms that have been shown experimentally to generate pattern and form in metazoan organisms. We propose that all such mechanisms can be organized into three basic categories and that two of these may act as composite mechanisms in two different ways. The simple categories are cell autonomous mechanisms in which cells enter into specific arrangements ('patterns') without interacting, inductive mechanisms in which cell communication leads to changes in pattern by reciprocal or hierarchical alteration of cell phenotypes ('states') and morphogenetic mechanisms in which pattern changes by means of cell interactions that do not change cell states. The latter two types of mechanism can be combined either morphostatically, in which case inductive mechanisms act first, followed by the morphogenetic mechanism, or morphodynamically, in which case both types of mechanisms interact continuously to modify each other's dynamics. We propose that this previously unexplored distinction in the operation of composite developmental mechanisms provides insight into the dynamics of many developmental processes. In particular, morphostatic and morphodynamic mechanisms respond to small changes in their genetic and microenvironmental components in dramatically different ways. We suggest that these differences in 'variational properties' lead to morphostatic and morphodynamic mechanisms being represented to different extents in early and late stages of development and to their contributing in distinct ways to morphological transitions in evolution.

Animals↗

Creating high-level components with a generative representation for body-brain evolution.

One of the main limitations of scalability in body-brain evolution systems is the representation chosen for encoding creatures. This paper defines a class of representations called generative representations, which are identified by their ability to reuse elements of the genotype in the translation to the phenotype. This paper presents an example of a generative representation for the concurrent evolution of the morphology and neural controller of simulated robots, and also introduces GENRE, an evolutionary system for evolving designs using this representation. Applying GENRE to the task of evolving robots for locomotion and comparing it against a non-generative (direct) representation shows that the generative representation system rapidly produces robots with significantly greater fitness. Analyzing these results shows that the generative representation system achieves better performance by capturing useful bias from the design space and by allowing viable large scale mutations in the phenotype. Generative representations thereby enable the encapsulation, coordination, and reuse of assemblies of parts.

Algorithms↗

First steps in eukaryogenesis: physical phenomena in the origin and evolution of chromosome structure.

Our present understanding of the origin and evolution of chromosomes differs considerably from current understanding of the origin and evolution of the cell itself. Chromosome origins have been less prominent in research, as the emphasis has not shifted so far appreciably from the phenomenon of primeval nucleic acid encapsulation to that of the origin of gene organization, expression, and regulation. In this work we discuss some reasons why preliminary steps in this direction are being taken. We have been led to examine properties that have contributed to raise the ancestral prokaryotic programmes to a level where we can appreciate in eukaryotes a clear departure from earlier themes in the evolution of the cell from the last common ancestor. We shift our point of view from evolution of cell morphology to the point of view of the genes. In particular, we focus attention on possible physical bases for the way transmission of information has evolved in eukaryotes, namely, the inactivation of whole chromosomes. The special case of the inactivation of the X chromosome in mammals is discussed, paying particular attention to the physical process of the spread of X inactivation in monotremes (platypus and echidna). When experimental data is unavailable some theoretical analysis is possible based on the idea that in certain cases collective phenomena in genetics, rather than chemical detail, are better correlates of complex chemical processes.

Animals↗

Rapid radiation and cryptic speciation in squat lobsters of the genus Munida (Crustacea, Decapoda) and related genera in the South West Pacific: molecular and morphological evidence.

Squat lobsters (genus Munida and related genera) are among the most diverse taxa of western Pacific crustaceans, though several features of their biology and phylogenetic relationships are unknown. This paper reports an extensive phylogenetic analysis based on mitochondrial DNA sequences (cytochrome c oxidase subunit I and 16S rRNA) and the morphology of 72 species of 12 genera of western Pacific squat lobsters. Our phylogenetic reconstruction using molecular data supports the recent taxonomic splitting of the genus Munida into several genera. Excluding one species (M. callista), the monophyly of the genus Munida was supported by Bayesian analysis of the molecular data. Three moderately diverse genera (Onconida, Paramunida, and Raymunida) also appeared monophyletic, both according to morphological and molecular data, always with high support. However, other genera (Crosnierita and Agononida) seem to be para- or polyphyletic. Three new cryptic species were identified in the course of this study. It would appear that the evolution of this group was marked by rapid speciation and stasis, or certain constraints, in its morphological evolution.

Animals↗

Functional comparison of the nematode Hox gene lin-39 in C. elegans and P. pacificus reveals evolutionary conservation of protein function despite divergence of primary sequences.

Hox transcription factors have been implicated in playing a central role in the evolution of animal morphology. Many studies indicate the evolutionary importance of regulatory changes in Hox genes, but little is known about the role of functional changes in Hox proteins. In the nematodes Pristionchus pacificus and Caenorhabditis elegans, developmental processes can be compared at the cellular, genetic, and molecular levels and differences in gene function can be identified. The Hox gene lin-39 is involved in the regulation of nematode vulva development. Comparison of known lin-39 mutations in P. pacificus and C. elegans revealed both conservation and changes of gene function. Here, we study evolutionary changes of lin-39 function using hybrid transgenes and site-directed mutagenesis in an in vivo assay using C. elegans lin-39 mutants. Our data show that despite the functional differences of LIN-39 between the two species, Ppa-LIN-39, when driven by Cel-lin-39 regulatory elements, can functionally replace Cel-lin-39. Furthermore, we show that the MAPK docking and phosphorylation motifs unique for Cel-LIN-39 are dispensable for Cel-lin-39 function. Therefore, the evolution of lin-39 function is driven by changes in regulatory elements rather than changes in the protein itself.

Amino Acid Sequence↗

Evolution of the mdx mouse cardiomyopathy: physiological and morphological findings.

Heart failure is a major cause of death in boys with Duchenne muscular dystrophy. In order to determine if the cardiac function of the mdx mouse is similarly disturbed, we performed murine echocardiograms and left heart catheterization studies, along with morphometric analysis of cardiac fibrosis. Serial echocardiograms in mdx mice revealed the evolution from normal cardiac function in young mice to a dilated cardiomyopathy in adult mice. Very old mdx mice exhibited a widespread but patchy increase in ventricular wall fibrosis. These results show that the mdx cardiac function is more impaired than was previously thought and shares important clinical features with the cardiomyopathy of Duchenne muscular dystrophy.

Aging↗

Evolution of the Pt layer deposited on MgO(001) by pulsed laser deposition as a function of the deposition parameters: a scanning tunneling microscopy and energy dispersive X-ray diffractometry/reflectometry study.

A combined ultrahigh vacuum scanning tunneling microscopy (STM-UHV) and energy dispersive X-ray diffractometry/reflectometry (EDXD/EDXR) study of the evolution of face-centered cubic (fcc) Pt layer growth on MgO(100) by pulsed laser deposition as a function of the process parameters such as deposition temperature and deposition duration has been carried out. The aim of this study is to define the best experimental conditions to obtain a controlled film deposition selective on the Pt growth direction (either [111] or [002]). The evolution of the Pt surface morphology as a function of the deposition temperature (T(dep)) from 300 to 700 degrees C has been studied with STM and ED techniques. Results show that the Pt surface, characterized at T(dep) = 300 degrees C by a 3D island morphology, evolves at higher temperatures to a morphology in which the original islands coexist with a distribution of orthogonal 2D stripes. The two features can be associated with the [111] and [002] Pt growth directions of the fcc phase, respectively. For T(dep) = 700 degrees C, the island morphology of the (111) face completely disappears, while the merging process of the (002) stripes reaches completion. The evolution of the morphology at T(dep) = 600 degrees C as a function of the deposition time and thickness has then been studied with STM-UHV, revealing an initial growth of mosaic-like 3D islands. These independent islands, already interconnected, expand along two orthogonal directions and, for longer deposition times, lead to the texture of orthogonal stripes. The EDXR characterization providing the morphological parameters of the films, i.e., thickness and roughness, confirms the above observation and quantifies the effect of such morphological changes on the surface roughness of the Pt film, an important parameter for applications of Pt films as underlayer in magnetic recording media.

Journal Article↗

Exploring developmental, functional, and evolutionary aspects of amphioxus sensory cells.

Amphioxus has neither elaborated brains nor definitive sensory organs, so that the two may have evolved in a mutually affecting manner and given rise to the forms seen in extant vertebrates. Clarifying the developmental and functional aspects of the amphioxus sensory system is thus pivotal for inferring the early evolution of vertebrates. Morphological studies have identified and classified amphioxus sensory cells; however, it is completely unknown whether the morphological classification makes sense in functional and evolutionary terms. Molecular markers, such as gene expression, are therefore indispensable for investigating the developmental and functional aspects of amphioxus sensory cells. This article reviews recent molecular studies on amphioxus sensory cells. Increasing evidence shows that the non-neural ectoderm of amphioxus can be subdivided into molecularly distinct subdomains by the combinatorial code of developmental cues involving the RA-dependent Hox code, suggesting that amphioxus epithelial sensory cells developed along positional information. This study focuses particularly on research involving the molecular phylogeny and expression of the seven-transmembrane, G protein-coupled receptor (GPCR) genes and discusses the usefulness of this information for characterizing the sensory cells of amphioxus.

Animals↗

Evolution of the dentition in prehistoric Ohio Valley Native Americans: II. Morphology of the deciduous dentition.

In order to evaluate the microevolutionary dynamics of morphological features of the deciduous dentition, I collected data on the variation of 57 features (33 crown and 24 root) from prehistoric Ohio Valley populations. I sampled a total of 370 individuals from 26 populations representing a lineage that inhabited the middle and upper Ohio valley region from approximately 3000 to 350 BP. Evolutionary changes in the frequencies of morphological features of the deciduous teeth in this lineage were limited. Over 80% of the features show no significant differences among the populations. The relatively few features that show consistent differences separate pre- and postmaize agricultural populations. I discuss explanations for this change in terms of selection differences or gene flow. The general pattern of morphological trait expression in the deciduous teeth of this Ohio Valley lineage corresponds to what has been termed the Mongoloid dental complex (sinodonty in the permanent teeth). I suggest additional features that, with further study, may be added to this morphological complex.

Biological Evolution↗

Variation in neuromuscular activity during prey capture by trophic specialists and generalists (Pisces: Labridae).

Members of the marine teleost family Labridae are among the most abundant and morphologically diverse fish on coral reefs. A quantitative analysis was conducted of the neuromuscular activity patterns controlling movement of the jaws during prey capture by 4 labrid species ranging from trophic specialists to trophic generalists. A total of more than 800 captures of 3 prey types was analyzed. All 4 species showed significant modulation of electromyographic parameters in response to different prey types. Significant variation was also found between replicate experiments on the same individuals. To obtain valid assessments of interspecific variability, statistical analyses must take into account this potentially high degree of intraspecific variability. By partitioning the variance in a nested analysis of variance, a lack of significant differences in electromyographic parameters between species became apparent. In contrast to the closely related Cichlidae, trophic diversification in the Labridae has not been accompanied by the acquisition of unique neuromuscular activity patterns for prey capture. The dramatic adaptive radiation that has occurred in these 2 families has involved different processes of evolutionary diversification. Neuromuscular stereotypy of labrids may be associated with the lack of structural flexibility in their 'coupled jaw'. Additional study is needed to establish the extent to which labrid radiation into various trophic niches is related to the evolution of specialized morphologies and foraging behaviors.

Animals↗

[Diagnosis of subependymal cysts by brain ultrasound].

OBJECTIVE: The aim of this study was to describe the clinical data and sonographic evolution of subependymal cysts (SEC) and to compare alone SEC versus SEC combined with other brain abnormalities. METHODS: The diagnosis of SEC relied on the discovery of one/several cysts at the outer angle of one/both lateral ventricles detected by routine brain ultrasound tests in the first days of life. During the 1981-2000 period, 66 neonates had evidence of SEC in Port-Royal Department of Neonatal Medicine. The cases were divided into two groups: G-I, alone SEC; G-II, SEC combined with other brain abnormalities. Hospital charts were reviewed for gestational and delivery data and ultrasound scans were analyzed according to a preset list of items (dates, measurements, other findings). RESULTS: Patients with alone SEC had a higher gestational and birthweight, better birth conditions, and less respiratory morbidity. High rates of associated congenital anomalies were observed in both groups (19% in G-I, 13% in G-II, NS). Rare pre/perinatal infections did not show any association with SEC. SEC were uni or bilateral, single or multiple (string), with no difference between G-I and G-II. Unilateral SEC predominated in the left side (26/28, P<0.01). Serial examinations were carried out in 49/66 (74%), showing an increase in size of SEC in 21/49 (45%) in the first month of life, while 12 SEC (24%) disappeared. The 11 deaths occurred in neonates with other severe brain injuries (five peri-intraventricular hemorrhages, four periventricular leukomalacias) except one (diaphragmatic hernia). CONCLUSIONS: Alone SEC and SEC combined with other brain abnormalities follow the same morphologic pattern and evolution. The striking predominance in the left side may suggest another etiology. Both are in favor of a developmental deviation and not a viral fetopathy.

English Abstract↗

Variation and selection at the CAULIFLOWER floral homeotic gene accompanying the evolution of domesticated Brassica oleracea.

The evolution of plant morphologies during domestication events provides clues to the origin of crop species and the evolutionary genetics of structural diversification. The CAULIFLOWER gene, a floral regulatory locus, has been implicated in the cauliflower phenotype in both Arabidopsis thaliana and Brassica oleracea. Molecular population genetic analysis indicates that alleles carrying a nonsense mutation in exon 5 of the B. oleracea CAULIFLOWER (BoCAL) gene are segregating in both wild and domesticated B. oleracea subspecies. Alleles carrying this nonsense mutation are nearly fixed in B. oleracea ssp. botrytis (domestic cauliflower) and B. oleracea ssp. italica (broccoli), both of which show evolutionary modifications of inflorescence structures. Tests for selection indicate that the pattern of variation at this locus is consistent with positive selection at BoCAL in these two subspecies. This nonsense polymorphism, however, is also present in both B. oleracea ssp. acephala (kale) and B. oleracea ssp. oleracea (wild cabbage). These results indicate that specific alleles of BoCAL were selected by early farmers during the domestication of modified inflorescence structures in B. oleracea.

Base Sequence↗

Cluster-cluster aggregation kinetics and primary particle growth of soot nanoparticles in flame by light scattering and numerical simulations.

The agglomeration kinetics of growing soot generated in a diffusion atmospheric flame are here studied in situ by light scattering technique to infer cluster morphology and size (fractal dimension D(f) and radius of gyration R(g)). SEM analysis is used as a standard reference to obtain primary particle size D(P) at different residence times. The number N(P) of primary particles per aggregate and the number concentration n(A) of clusters are evaluated on the basis of the measured angular patterns of the scattered light intensity. The major finding is that the kinetics of the coagulation process that yields to the formation of chain-like aggregates by soot primary particles (size 10 to 40 nm) can be described with a constant coagulation kernel beta(c,exp)=2.37x10(-9) cm3/s (coagulation constant tau(c) approximately = 0.28 ms). This result is in nice accord with the Smoluchowski coagulation equation in the free molecular regime, and, vice versa, it is in contrast with previous studies conducted by invasive (ex situ) techniques, which claimed the evidence in flames of coagulation rates much larger than the kinetic theory predictions. Thereafter, a number of numerical simulations is implemented to compare with the experimental results on primary particle growth rate and on the process of aggregate reshaping that is observed by light scattering at later residence times. The restructuring process is conjectured to occur, for not well understood reasons, as a direct consequence of the atomic rearrangement in the solid phase carbon due to the prolonged residence time within the flame. Thus, on one side, it is shown that the numerical simulations of primary size history compare well with the values of primary size from SEM experiment with a growth rate constant of primary diameter about 1 nm/s. On the other side, the evolution of aggregate morphology is found to be predictable by the numerical simulations when the onset of a first-order "thermal" restructuring mechanism is assumed to occur in the flame at about 20 ms residence time leading to aggregates with an asymptotic fractal dimension D(f,infinity) approximately = 2.5.

Journal Article↗

Complexities in ETS-domain transcription factor function and regulation: lessons from the TCF (ternary complex factor) subfamily. The Colworth Medal Lecture.

The ETS-domain transcription factor family can be divided into a series of subfamilies. Elk-1 represents the founding member of the ternary complex factor (TCF) subfamily. By focusing on the TCF subfamily, we can demonstrate the complexities that exist in the function and regulation of ETS-domain transcription factors. This article focuses on Elk-1 in detail and summarizes the functions of other TCFs. The key themes covered include the domain structure of the TCFs, the mechanisms of complex formation with serum response factor, regulation of TCFs by mitogen-activated protein kinase cascades, and transcriptional regulatory properties of the TCFs. Finally, the emerging role of the TCFs in vivo is discussed. A picture is developing indicating that, while these proteins exhibit significant sequence and functional conservation, key differences in their structure and regulation are being identified which may relate to unique functions of these proteins in vivo.

Amino Acid Sequence↗