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PubMedDate not supplied

Why are arthropods segmented?

Segmentation as an attribute of organisms is being increasingly discussed in the recent literature because (1) new phylogenies suggest that organisms classically considered to be segmented may lie in separate clades; (2) the molecular basis of segmental development has been much studied; (3) various theories of bilaterian origins place weight on segmentation as a primitive character; (4) there has been recent stress on the importance of modularity as an evolutionary topic. However, the definition and extent of segmentation are highly ambiguous and usually typological. Here, segmentation is regarded as an attribute of organs, not organisms. The evolution of just one system, the arthropod epidermis, is examined on the basis of the fossil record and the extant euarthropods, tardigrades, and onychophorans. It may be seen to have become segmented in a complex pathway that necessitated shifts in function, redundancy, and changes in associated organs. This complexity must inevitably reflect on, and to an extent have primacy over, the genetic basis for the changes involved. Evolutionary functional morphology has been relatively little considered in the context of the evolution of development, but may play an important role in defining the framework within which this evolution occurs.

Animals↗
PubMedDate not supplied

The evolutionary history of crustacean segmentation: a fossil-based perspective.

The evolution of segmentation in Crustacea, that is, the formation of sclerotized and jointed body somites and arrangement of somites into tagmata, is viewed in light of historical traits and functional constraints. The set of Early to Late Cambrian 'Orsten' arthropods have informed our current views of crustacean evolution considerably. These three-dimensionally preserved fossils document ancient morphologies, as opposed to purely hypothetical models and, because of the unusual preservation of larval stages, provide us with unparalleled insight into the morphogenesis of body somites and their structural equipment. The variety of evolutionary levels represented in the 'Orsten' including lobopodians, tardigrades, and pentastomids also allows phylogenetic interpretations far beyond the Crustacea. The 'Orsten' evidence and data from representatives of the Lower Cambrian Chengjiang biota in southwestern China, including phylogenetically earlier forms, form the major source of our morphology-based review of structural and functional developments that led toward the Crustacea. The principal strategy of arthropods is the simultaneous development of head somites, as expressed in a basal "head larva," and a successive addition of postcephalic somites from a preterminal budding zone with progressive maturation of metameric structures. This can be recognized in the developmental patterns of extant and fossil representatives of several euarthropod taxa, particularly crustaceans, trilobites, and chelicerates (at least basally). The development of these taxa points to an early somite-poor and free-living hatching stage. Embryonic development to a late stage within an egg, as occurring in recent onychophorans and certain in-group euarthropods, is regarded as achieved several times convergently.

Animals↗
PubMed2000

Asymmetry of configuration space induced by unequal crossover: implications for a mathematical theory of evolutionary innovation.

Evolution can be regarded as the exploration of genetic or morphological state space by populations. In traditional models of population and quantitative genetics, the state space can be formally represented as a configuration space with clearly defined concepts of neighborhood and distance, defined by the action of variational operators such as mutation and/or recombination. In this paper, we describe a process where no genetic configuration space closure (and hence, no non-arbitrary notion of distance and neighborhood) exists. The process is gene duplication by means of unequal crossover, which we regard as an example of an "innovation" process that changes the state space of the system rather than exploring a closed state space. We assert that such processes are qualitatively distinct from representations of the adaptation process, which occur on regular configuration spaces.

Biological Evolution↗
PubMed2007

How to tweak a beak: molecular techniques for studying the evolution of size and shape in Darwin's finches and other birds.

A flurry of technological advances in molecular, cellular and developmental biology during the past decade has provided a clearer understanding of mechanisms underlying phenotypic diversification. Building upon such momentum, a recent paper tackles one of the foremost topics in evolution, that is the origin of species-specific beak morphology in Darwin's finches.1 Previous work involving both domesticated and wild birds implicated a well-known signaling pathway (i.e. bone morphogenetic proteins) and one population of progenitor cells in particular (i.e. cranial neural crest), as primary factors for establishing beak size and shape. But these results were limited in their ability to explain fully the morphogenetic bases of patterned outgrowth. So in a quest to identify novel genes whose expression correlated with differences in beak anatomy among Darwin's finches, a DNA microarray approach was undertaken using tissues harvested from the Galápagos Islands. The results are striking and point to a protein called calmodulin, which is a mediator of cellular calcium signaling, as a key determinant of beak length.

Animals↗
PubMed1991

Masticatory-stress hypotheses and the supraorbital region of primates.

The purpose of this study is to test various masticatory-stress hypotheses about the evolution and function of well-developed browridges of higher primates. This was done by measuring and analyzing patterns of in vivo bone strain recorded from three-element rosette strain gages bonded to the supraorbital region and to other portions of the bony face of Macaca fascicularis and Papio anubis during mastication and incision. The magnitude and direction of the principal strains recorded support Endo's hypothesis that the supraorbital region during mastication and incision is bent in the frontal plane (Endo, 1966). Our data do not, however, support his hypothesis that the supraorbital region is bent more during incision than during mastication. The data also demonstrate that overall levels of supraorbital strain are not larger in more prognathic subjects. Most importantly, the data indicate that the supraorbital region of nonhuman catarrhines is strained very little during mastication and incision. This indicates that there is much more supraorbital bone than is necessary both to counter masticatory loads and to provide an adequate safety factor to failure for these loads. This in turn suggests that the macaque and baboon browridges can be considerably reduced in size and still maintain these required structural characteristics. Thus, our experiments provide no support whatsoever for those hypotheses that directly link browridge morphology to masticatory stress (cf. Endo, 1966; Russell, 1983, 1985). A recent review of Endo's original work indicates that this latter statement is also true for humans (Picq and Hylander, 1989). We conclude, therefore, that there is no good reason to believe that enlarged browridges in living and/or fossil primates are structural adaptations to counter intense masticatory forces. The evolution of browridge morphology in primates is best explained on the basis of factors related to the position of the brain relative to the orbits (Moss and Young, 1960). When these structures are widely separated, as in gorillas, the large intervening space must be bridged with bone. In addition, enough bone must be present within the supraorbital and bridged regions to prevent structural failure due to non-masticatory external forces associated with highly active primates (e.g., accidental traumatic forces applied to the orbits and neurocranium). This requirement results in both pronounced browridges and in much more supraorbital bone than is necessary to counter routine cyclical stress during mastication and incision. This in turn explains why bone strains recorded from the supraorbital region are extremely small relative to other portions of the primate face during mastication and incision.

Animals↗
PubMed2006

Assigning sex to pre-adult stalk-eyed flies using genital disc morphology and X chromosome zygosity.

BACKGROUND: In stalk-eyed flies (Diopsidae) the eyes and antennae are laterally displaced at the ends of elongated eyestalks. Eyespan and the degree of sexual dimorphism in eyespan vary considerably between species and several sexually dimorphic species show sexual selection through female mate preference for males with exaggerated eyespan. The genes on which selection acts to regulate eyespan remain to be identified. This could be achieved by comparing gene expression during eyestalk development in males and females if the sex of pre-adult flies could be reliably assigned. Here we describe two techniques, one morphological and one microsatellite-based, that identify the sex of stalk-eyed fly larvae and pupae. RESULTS: We showed that genital discs of the stalk-eyed fly Teleopsis dalmanni have two highly distinct morphologies, compact ("C") and lobed ("L"). Segment composition (revealed by Engrailed expression) was consistent with C morphology being typical of males and L morphology of females. We confirmed the proposed association between disc morphology and sex by evaluating the combined heterozygosity of four X-linked microsatellite markers. We demonstrated that individuals with C genital discs had hemizygous (male) genotypes while those with L discs were heterozygous (female) genotypes. Similar dimorphism in genital disc morphology was observed in eight other species spanning three representative Diopsid genera. In every case the segment composition supported C morphology being male and L morphology female. We assigned larval sex by C or L morphology and compared cell division frequencies in male and female eye-antennal discs in two species (T. dalmanni and Diasemopsis meigenii) sexually dimorphic for eyespan. The number of mitotic (anti-H3-labelled) cells did not differ between the sexes in either species. CONCLUSION: We have made novel use of two complementary techniques for identifying the sex of pre-adult stalk-eyed flies. These procedures will facilitate studies of the evolution of sexually dimorphic development in a variety of other species. Morphology and En expression in male and female genital discs are highly conserved within each genus of Diopsidae. Finally, sexual dimorphism for eyespan in two Diopsid species is unlikely to be due to an increased rate of cell division at the third larval instar in males.

Animals↗
PubMed2005

The dynamics of genetic and morphological variation on volcanic islands.

Oceanic archipelagos of volcanic origin have been important in the study of evolution because they provide repeated natural experiments allowing rigorous tests of evolutionary hypotheses. Ongoing volcanism on these islands may, however, affect the evolutionary diversification of species. Analysis of population structure and phylogeographic patterns in island populations can provide insight into evolutionary dynamics on volcanic islands. We analysed genetic and morphological variation in the gecko Tarentola boettgeri on the island of Gran Canaria and compared it with Tarentola delalandii on Tenerife, a neighbouring volcanic island of similar age but distinctly different geological past. Intraspecific divergence of mitochondrial haplotypes indicates long-term persistence of Tarentola on each island, with a phylogeographic signal left by older volcanic events. More recent volcanic eruptions (approximately 0.2 million years ago on Tenerife, approximately 2.2 million years ago on Gran Canaria) have left a signature of population expansion in the population genetic structure, the strength of which depends on the time since the last major volcanic eruption on each island. While these stochastic events have left traces in morphological variation in Tenerife, in Gran Canaria geographical variation was solely associated with environmental variables. This suggests that historically caused patterns in morphology may be overwritten by natural selection within 2 million years.

Animals↗
PubMed1991

Paleozoic Foraminifera.

The approximately 300 million years that make up Paleozoic time saw the evolution of eight of the fifteen recognized suborders of Foraminifera. Of the suborders present in the Paleozoic, seven are morphologically relatively simple, slowly evolving, and continued into Mesozoic and Cenozoic times to become the ancestoral lineages from which evolved several additional post-Paleozoic suborders. In contrast, an eighth Paleozoic suborder, the Fusulinina, was an abundant, ecologically dominant group that evolved from simple to highly specialized forms and had a history of rapid evolution with diverse lineages. Fusulinines became extinct at the end of the Paleozoic. Their early representatives may have given rise to three and eventually four post-Paleozoic suborders. A number of suborders in the Paleozoic have similar, supposedly independent, early evolutionary patterns with the following series of morphological steps: (1) single chambers with or without apertures depending on the amount of wall cement; (2) groups of chambers that appear to be buds or aggregations of individuals rather than true chambers; (3) a proloculus followed by a tubular second chamber that is first erect and gradually evolves into enrolled free-living individuals; (4) development of constrictions in the tubular chamber; and finally (5) evolution of true chambers. These morphological steps, which are basic organizational steps with evolutionary significance, appear in lineages with quite different test compositions and, therefore, are considered only distantly related in the present classification.

Animals↗
PubMed2006

Eukaryotic organisms in Proterozoic oceans.

The geological record of protists begins well before the Ediacaran and Cambrian diversification of animals, but the antiquity of that history, its reliability as a chronicle of evolution and the causal inferences that can be drawn from it remain subjects of debate. Well-preserved protists are known from a relatively small number of Proterozoic formations, but taphonomic considerations suggest that they capture at least broad aspects of early eukaryotic evolution. A modest diversity of problematic, possibly stem group protists occurs in ca 1800-1300 Myr old rocks. 1300-720 Myr fossils document the divergence of major eukaryotic clades, but only with the Ediacaran-Cambrian radiation of animals did diversity increase within most clades with fossilizable members. While taxonomic placement of many Proterozoic eukaryotes may be arguable, the presence of characters used for that placement is not. Focus on character evolution permits inferences about the innovations in cell biology and development that underpin the taxonomic and morphological diversification of eukaryotic organisms.

Biological Evolution↗
PubMedDate not supplied

Diverse developmental mechanisms contribute to different levels of diversity in horned beetles.

An ongoing challenge to evolutionary developmental biology is to understand how developmental evolution on the level of populations and closely related species relates to macroevolutionary transformations and the origin of morphological novelties. Here we explore the developmental basis of beetle horns, a morphological novelty that exhibits remarkable diversity on a variety of levels. In this study, we examined two congeneric Onthophagus species in which males develop into alternative horned and hornless morphs and different sexes express marked sexual dimorphism. In addition, both species differ in the body region (head vs. thorax) that develops the horn. Using a comparative morphological approach we show that prepupal growth of horn primordia during late larval development, as well as reabsorption of horn primordia during the pupal stage, contribute to horn expression in adults. We also show that variable combinations of both mechanisms are employed during development to modify horn expression of different horns in the same individual, the same horn in different sexes, and different horns in different species. We then examine expression patterns of two transcription factors, Distal-less (Dll) and aristaless (al), in the context of prepupal horn growth in alternative male morphs and sexual dimorphisms in the same two species. Expression patterns are qualitatively consistent with the hypothesis that both transcription factors function in the context of horn development similar to their known roles in patterning a wide variety of arthropod appendages. Our results suggest that the origin of morphological novelties, such as beetle horns, rests, at least in part, on the redeployment of already existing developmental mechanisms, such as appendage patterning processes. Our results also suggest, however, that little to no phylogenetic distance is needed for the evolution of very different modifier mechanisms that allow for substantial modulation of trait expression at different time points during development in different species, sexes, or tissue regions of the same individual. We discuss the implications of our results for our understanding of the evolution of horned beetle diversity and the origin and diversification of morphological novelties.

Animals↗
PubMedDate not supplied

Male pregnancy in seahorses and pipefishes (family Syngnathidae): rapid diversification of paternal brood pouch morphology inferred from a molecular phylogeny.

In contrast to the majority of vertebrate species, primary male parental care is common in fishes and encompasses a remarkable diversity of adaptations. Seahorses and pipefishes (Family Syngnathidae) exhibit some of the most specialized forms of paternal care in animals and so are ideally suited to the study of the evolution of male parental care. During mating, female syngnathids transfer eggs to specialized morphological structures that are located on either the abdomen or tail of the male. The male provides all postfertilization parental care and has morphological and physiological adaptations to osmoregulate, aerate, and even nourish the developing embryos. While all syngnathid species are adapted for paternal care, the brooding structure with which this is accomplished varies between species, from simple ventral gluing areas to much more complex structures such as the completely enclosed pouches of the seahorses. Our combined cytochrome b-, 12S rDNA-, and 16S rDNA-based molecular phylogeny of syngnathid fishes demonstrates that rapid diversification of male brooding structures has been associated with the major evolutionary radiation of the group, suggesting that development and diversification of structures involved in paternal care may have been key evolutionary innovations of the Syngnathidae. Molecular analyses also highlight geographical centers of biodiversity and suggest interoceanic migration of Syngnathus pipefishes from their center of origin in the Pacific.

Animals↗
PubMed2001

The evolution of sexual size dimorphism in the house finch. III. Developmental basis.

Sexual size dimorphism of adults proximately results from a combination of sexually dimorphic growth patterns and selection on growing individuals. Yet, most studies of the evolution of dimorphism have focused on correlates of only adult morphologies. Here we examined the ontogeny of sexual size dimorphism in an isolated population of the house finch (Carpodacus mexicanus). Sexes differed in growth rates and growth duration; in most traits, females grew faster than males, but males grew for a longer period. Sexual dimorphism in bill traits (bill length, width, depth) and in body traits (wing, tarsus, and tail length; mass) developed during different periods of ontogeny. Growth of bill traits was most different between sexes during the juvenile period (after leaving the nest), whereas growth of body traits was most sexually dimorphic during the first few days after hatching. Postgrowth selection on juveniles strongly influenced sexual dimorphism in all traits; in some traits, this selection canceled or reversed dimorphism patterns produced by growth differences between sexes. The net result was that adult sexual dimorphism, to a large degree, was an outcome of selection for survival during juvenile stages. We suggest that previously documented fast and extensive divergence of house finch populations in sexual size dimorphism may be partially produced by distinct environmental conditions during growth in these populations.

Animals↗
PubMed2000

What it takes to fly: the structural and functional respiratory refinements in birds and bats.

In absolute terms, flight is a highly energetically expensive form of locomotion. However, with respect to its cost per unit distance covered, powered flight is a very efficient mode of transport. Birds and bats are the only extant vertebrate taxa that have achieved flight. Phylogenetically different, they independently accomplished this elite mode of locomotion by employing diverse adaptive schemes and strategies. Integration of functional and structural parameters, a transaction that resulted in certain trade-offs and compromises, was used to overcome exacting constraints. Unique morphological, physiological and biochemical properties were initiated and refined to enhance the uptake, transfer and utilization of oxygen for high aerobic capacities. In bats, exquisite pulmonary structural parameters were combined with optimal haematological ones: a thin blood-gas barrier, a large pulmonary capillary blood volume and a remarkably extensive alveolar surface area in certain species developed in a remarkably large lung. These factors were augmented by, for example, exceptionally high venous haematocrits and haemoglobin concentrations. In birds, a particularly large respiratory surface area and a remarkably thin blood-gas (tissue) barrier developed in a small, rigid lung; a highly efficient cross-current system was fabricated within the parabronchi. The development of flight in only four animal taxa (among all the animal groups that have ever evolved; i.e. insects, the now-extinct pterosaurs, birds and bats) provides evidence for the enormous biophysical and energetic constraints that have stymied volancy. Bats improved a fundamentally mammalian lung to procure the large amounts of oxygen needed for flight. The lung/air sac system of birds is not therefore a prescriptive morphology for flight: the essence of its design can be found in the evolution of the reptilian lung, the immediate progenitor stock from which birds arose. The attainment of flight is a classic paradigm of the remarkable adaptability inherent in organismal and organic biology for countering selective pressures by initiating elegant morphologies and physiologies.

Altitude↗
PubMed1980

Evolution and gene transfer in purple photosynthetic bacteria.

A concern voiced in connection with recent sequencing of cytochrome c from the Rhodospirillaceae or purple non-sulphur photosynthetic bacteria is that molecular information might be of little use in deciphering bacterial phylogeny because of the possibility of lateral transfer of genes and the consequent scrambling of the genetic record. This could be true for many proteins, of course, but the immediate question is, is it true for cytochroms available for comparison? The evidence suggests that this is probably not so. The disagreement between cytochrome c sequences and the standard taxonomy of the Rhodospirillaceae in Bergey's Manual is not a problem. That reference, as its title indicates, is a manual of determinative rather than evolutionary bacteriology. Its goal is a reproducible system for identification of bacteria. If one claims that these determinative categories also have phylogenetic or evolutionary meaning, this is an assertion that must be proven. The argument outlined below suggests that molecular traits may eventually become more a dependable basis for classification of the Rhodospirillaceae than is gross morphology.

Base Sequence↗