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PubMed2003

The evolution of inquilinism, host-plant use and mitochondrial substitution rates in Tamalia gall aphids.

We used mitochondrial DNA data to infer phylogenies for 28 samples of gall-inducing Tamalia aphids from 12 host-plant species, and for 17 samples of Tamalia inquilinus, aphid 'inquilines' that obligately inhabit galls of the gall inducers and do not form their own galls. Our phylogenetic analyses indicate that the inquilines are monophyletic and closely related to their host aphids. Tamalia coweni aphids from different host plants were, with one exception, very closely related to one another. By contrast, the T. inquilinus aphids were strongly genetically differentiated among most of their host plants. Comparison of branch lengths between the T. coweni clade and the T. inquilinus clade indicates that the T. inquilinus lineage evolves 2.5-3 times faster for the cytochrome oxidase I gene. These results demonstrate that: (1) Tamalia inquilines originated from their gall-inducing hosts, (2) communal (multi-female) gall induction apparently facilitated the origin of inquilinism, (3) diversification of the inquilines has involved rapid speciation along host-plant lines, or the rapid evolution of host-plant races, and (4) the inquilines have undergone accelerated molecular evolution relative to their hosts, probably due to reduced effective population sizes. Our findings provide insight into the behavioural causes and evolutionary consequences of transitions from resource generation to resource exploitation.

Animals↗
PubMed2004

Historical biogeography of two cosmopolitan families of flowering plants: Annonaceae and Rhamnaceae.

Annonaceae are a pantropically distributed family found predominantly in rainforests, so they are megathermal taxa, whereas Rhamnaceae are a cosmopolitan family that tend to be found in xeric regions and may be classified as mesothermal. Phylogenetic analyses of these families are presented based on rbcL and trnL-F plastid DNA sequences. Likelihood ratio tests revealed rate heterogeneity in both phylogenetic trees and they were therefore made ultrametric using non-parametric rate smoothing and penalized likelihood. Divergence times were then estimated using fossil calibration points. The historical biogeography of these families that are species rich in different biomes is discussed and compared with other published reconstructions. Rhamnaceae and most lineages within Annonaceae are too young to have had their distribution patterns influenced by break-up of previously connected Gondwanan landmasses. Contrasts in the degree of geographical structure between these two families may be explained by differences in age and dispersal capability. In both groups, long-distance dispersal appears to have played a more significant role in establishing modern patterns than had previously been assumed. Both families also contain examples of recent diversification of species-rich lineages. An understanding of the processes responsible for shaping the distribution patterns of these families has contributed to our understanding of the historical assembly of the biomes that they occupy.

Annonaceae↗
PubMed1990

Combinatorial control of cell differentiation by cAMP and DIF-1 during development of Dictyostelium discoideum.

At least three distinct types of cell arise from a population of similar amoebae during Dictyostelium development: prespore, prestalk A and prestalk B cells. We report evidence suggesting that this cellular diversification can be brought about by the combinatorial action of two diffusible signals, cAMP and DIF-1. Cells at different stages of normal development were transferred to shaken suspension, challenged with various combinations of signal molecules and the expression of cell-type-specific mRNA markers measured 1-2 h later. pDd63, pDd56 and D19 mRNAs were used for prestalk A, prestalk B and prespore cells respectively. We find the following results. (1) Cells first become responsive to DIF-1 for prestalk A differentiation and to cAMP for prespore differentiation at the end of aggregation, about 2 h before these cell types normally appear. (2) At the first finger stage of development, when the rate of accumulation of the markers is maximal, the expression of each is favoured by a unique combination of effectors: prespore differentiation is stimulated by cAMP and inhibited by DIF-1; prestalk A differentiation is stimulated by both cAMP and DIF-1 and prestalk B differentiation is stimulated by DIF-1 and inhibited by cAMP. (3) Half-maximal effects are produced by 10-70 nM DIF-1, which is in the physiological range. (4) Ammonia and adenosine, which can affect cell differentiation in other circumstances, have no significant pathway-specific effect in our conditions. These results suggest that cell differentiation could be brought about in normal development by the localized action of cAMP and DIF-1.

Adenosine↗
PubMed2004

Contrasting modes of natural selection acting on pigmentation genes in the Drosophila dunni subgroup.

Genes that encode for divergent adaptive traits may have genealogies that contrast with those from loci that are not functionally involved in differentiation. Here, we examine DNA sequence variation among the species of the eastern Caribbean Drosophila dunni subgroup at two loci, yellow and dopa decaboxylase (Ddc), which both play integral roles in pigmentation patterning of adult Drosophila. Phylogenetic analyses of these loci produce gene genealogies with topologies that mirror those described for other nuclear genes: the six morphologically distinct species within the subgroup are divided into only three lineages, with one lineage containing four species that share extensive ancestral polymorphism. At the Ddc locus these major lineages are delineated only by silent site variation. We observe a significantly higher rate of synonymous site divergence than non-synonymous divergence, consistent with strong purifying selection acting on the locus. In contrast, the yellow locus exhibits patterns of amino acid divergence and nucleotide diversity that are consistent with recent diversifying selection acting in two different lineages. This selection appears to be targeting amino acid variants in the signal sequence of the Yellow protein, a region which is tightly constrained among members of the larger D. cardini radiation. This result highlights not only the potential importance of yellow in the evolution of divergent pigmentation patterns among members of the D. dunni subgroup, but also hints that variation in signal peptide sequences may play a role in phenotypic diversification.

Amino Acid Sequence↗
PubMed2004

Duplication and diversifying selection among termite antifungal peptides.

We have identified and analyzed the mRNA sequence of 20 new defensin-like peptides from 11 Australian termite species of Nasutitermes and from an outgroup, Drepanotermes rubriceps. The sequence was amplified by reverse transcriptase PCR with a degenerate primer designed from termicin, an antifungal peptide previously characterized from the termite Pseudocanthotermes spiniger. All 20 genes show high sequence identity with P. spiniger termicin and have duplicated repeatedly during the radiation of Nasutitermes. Comparison of the relative fixation rates of synonymous (silent) and nonsynonymous (amino acid altering) mutations indicates that the Nasutitermes termicins are positively selected. This positive selection appears to drive a decrease in termicin charge. In termites with two genes, the decrease in charge is predominantly restricted to one termicin. Furthermore, the spread of charge is significantly greater within species than across species among amino acid sites that appear to be under strong positive selection and this spread is attributable to only three sites. Our results suggest that after termicin duplication, certain critical sites have maintained a positive charge in one duplicate and evolved towards neutrality in the other and that positive selection has directed these changes repeatedly and independently. This diversification among duplicated genes may be a counter-response to the evolution of fungal resistance in social insects that are particularly vulnerable to fungal epidemics.

Amino Acid Sequence↗
PubMed2004

Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants.

We explore the phylogeny of the polygrammoid ferns using nucleotide sequences derived from three plastid loci for each of 98 selected species. Our analyses recovered four major monophyletic lineages: the loxogrammoids, two clades consisting of taxa restricted to the Old World, and a largely neotropical clade that also includes the pantropical Grammitidaceae. The loxogrammoid lineage diverges first and is sister to a large clade comprising the three remaining species-rich lineages. One paleotropical clade includes the drynarioid and selligueoid ferns, whereas the second paleotropical clade includes the platycerioids, lepisoroids, microsoroids, and their relatives. The grammitids nest within the neotropical clade, although the sister taxon of this circum-tropic, epiphytic group remains ambiguous. Microsorum and Polypodium, as traditionally defined, were recovered as polyphyletic. The relatively short branch lengths of the deepest clades contrast with the long branch lengths leading to the terminal groups. This suggests that the polygrammoid ferns arose through an old, rapid radiation. Our analysis also reveals that the rate of substitution in the grammitids is remarkably higher relative to other polygrammoids. Disparities in substitution rate may be correlated with one or more features characterizing grammitids, including species richness, chlorophyllous spores, and an extended gametophytic phase.

Classification↗
PubMed2005

Diversification before the most recent glaciation in Balanus glandula.

A deep genetic cline between southern populations of the barnacle Balanus glandula (from about Monterey Bay southward) and northern populations (from northern California through Alaska) has recently been described. If this pattern is due to historical isolation and genetic drift, we expect it to have formed recently and represent a transient, nonequilibrium state. However, this cline appears to have formed well before the last glacial maximum. Our assays of sequence diversity at a region of mitochondrial cytochrome oxidase I, combined with coalescent estimators of the time of separation for these two regions, suggest that a late Pleistocene event more than 100 thousand years ago may be responsible for the initial separation. This suggests that either strong oceanographic mechanisms or natural selection have maintained the cline, because there has clearly been adequate time for this cline or polymorphism to resolve itself by genetic drift and migration. However, reliance on only a single mitochondrial marker for which the substitution rate has been estimated still limits the resolution of our analysis.

Animals↗
PubMed2005

Quail-duck chimeras reveal spatiotemporal plasticity in molecular and histogenic programs of cranial feather development.

The avian feather complex represents a vivid example of how a developmental module composed of highly integrated molecular and histogenic programs can become rapidly elaborated during the course of evolution. Mechanisms that facilitate this evolutionary diversification may involve the maintenance of plasticity in developmental processes that underlie feather morphogenesis. Feathers arise as discrete buds of mesenchyme and epithelium, which are two embryonic tissues that respectively form dermis and epidermis of the integument. Epithelial-mesenchymal signaling interactions generate feather buds that are neatly arrayed in space and time. The dermis provides spatiotemporal patterning information to the epidermis but precise cellular and molecular mechanisms for generating species-specific differences in feather pattern remain obscure. In the present study, we exploit the quail-duck chimeric system to test the extent to which the dermis regulates the expression of genes required for feather development. Quail and duck have distinct feather patterns and divergent growth rates, and we exchange pre-migratory neural crest cells destined to form the craniofacial dermis between them. We find that donor dermis induces host epidermis to form feather buds according to the spatial pattern and timetable of the donor species by altering the expression of members and targets of the Bone Morphogenetic Protein, Sonic Hedgehog and Delta/Notch pathways. Overall, we demonstrate that there is a great deal of spatiotemporal plasticity inherent in the molecular and histogenic programs of feather development, a property that may have played a generative and regulatory role throughout the evolution of birds.

Animals↗
PubMed1997

Sequence and expression patterns of two forms of the middle molecular weight neurofilament protein (NF-M) of Xenopus laevis.

The middle molecular weight neurofilament protein (NF-M) is relevant to our understanding of vertebrate neurofilaments in growing axons, both because it exists in all vertebrates and because it undergoes characteristic changes in its phosphorylation state during axonal development. Indeed, all vertebrate neurofilament proteins are believed to have originated by gene duplication from an ancestral, NF-M-like protein. The role of NF-M in axonal development has been studied extensively in the frog, Xenopus laevis, through the use of monoclonal antibodies. To acquire a better understanding of the relationship of X. laevis NF-M to that of other vertebrates and to obtain additional reagents to study and perturb neurofilaments in developing axons, we isolated cDNA clones from the nervous system. These clones encoded two forms of NF-M, which exhibited 93% amino acid identity overall and 94%, 96% and 90% identity over their head, rod, and tail domains, respectively. Synonymous nucleotide substitution rates between the two forms tied their origin to an ancestral duplication of the Xenopus genome, which occurred approximately 30 million years ago. Non-synonymous substitution rates indicated that the tail domain is evolving more rapidly than the rod domain. Both forms shared structural features in common with other vertebrate NF-Ms but had only a single example of the KSP phosphorylation motif that is repeated multiple times in the NF-Ms of bird, goldfish and mammal. In post-metamorphic frogs, the NF-M(1) transcript was consistently expressed at higher levels than that of NF-M(2), although their anatomical patterns of expression were qualitatively similar. During development, however, only NF-M(2) was detectable in retinal ganglion cells through stage 42. We speculate that the differences observed between these two forms may represent early stages of protein diversification akin to what occurred after the gene duplications that gave rise to other vertebrate neurofilament proteins.

Amino Acid Sequence↗
PubMedDate not supplied

Assessing the cultures of medical group practices.

BACKGROUND: The culture of medical group practices is gaining increasing attention as one of the most important organizational factors influencing the costs and quality of health care. Based on organizational theory, we propose that the culture of the practice differs depending on size, ownership, location, and the number of medical specialties. METHODS: A survey was sent to 1223 physicians in 191 clinics in the upper Midwest. The clinic response rate was 77%. The survey instrument identifies 9 culture dimensions, each with 3 to 6 measurement statements. RESULTS: Smaller clinics had higher scores on 6 of the 9 dimensions. Physician-owned clinics had higher scores on 4 of the 9 dimensions, whereas system-owned clinics had a higher score on only 1 dimension. Only 1 dimension differed among the locations. Single-specialty clinics had higher scores on 4 dimensions and multispecialty clinics had higher scores on 2 dimensions. CONCLUSION: Our data confirm the contention that the culture of medical group practices varies considerably; to a degree, this variance is as predicted by organizational theory. The culture changes as group practices become larger and more complex through diversification into multispecialty practices or become part of larger health care systems.

Ambulatory Care Facilities↗
PubMed2004

Phylogenetic investigations of Antarctic notothenioid fishes (Perciformes: Notothenioidei) using complete gene sequences of the mitochondrial encoded 16S rRNA.

The Notothenioidei dominates the fish fauna of the Antarctic in both biomass and diversity. This clade exhibits adaptations related to metabolic function and freezing avoidance in the subzero Antarctic waters, and is characterized by a high degree of morphological and ecological diversity. Investigating the macroevolutionary processes that may have contributed to the radiation of notothenioid fishes requires a well-resolved phylogenetic hypothesis. To date published molecular and morphological hypotheses of notothenioids are largely congruent, however, there are some areas of significant disagreement regarding higher-level relationships. Also, there are critical areas of the notothenioid phylogeny that are unresolved in both molecular and morphological phylogenetic analyses. Previous molecular phylogenetic analyses of notothenioids using partial mtDNA 12S and 16S rRNA sequence data have resulted in limited phylogenetic resolution and relatively low node support. One particularly controversial result from these analyses is the paraphyly of the Nototheniidae, the most diverse family in the Notothenioidei. It is unclear if the phylogenetic results from the 12S and 16S partial gene sequence dataset are due to limited character sampling, or if they reflect patterns of evolutionary diversification in notothenioids. We sequenced the complete mtDNA 16S rRNA gene for 43 notothenioid species, the largest sampling to-date from all eight taxonomically recognized families. Phylogenetic analyses using both maximum parsimony and maximum likelihood resulted in well-resolved trees with most nodes supported with high bootstrap pseudoreplicate scores and significant Bayesian posterior probabilities. In all analyses the Nototheniidae was monophyletic. Shimodaira-Hasegawa tests were able to reject two hypotheses that resulted from prior morphological analyses. However, despite substantial resolution and node support in the 16S rRNA trees, several phylogenetic hypotheses among closely related species and clades were not rejected. The inability to reject particular hypotheses among species in apical clades is likely due to the lower rate of nucleotide substitution in mtDNA rRNA genes relative to protein coding regions. Nevertheless, with the most extensive notothenioid taxon sampling to date, and the much greater phylogenetic resolution offered by the complete 16S rRNA sequences over the commonly used partial 12S and 16S gene dataset, it would be advantageous for future molecular investigations of notothenioid phylogenetics to utilize at the minimum the complete gene 16S rRNA dataset.

Animals↗
PubMed2001

Which Is Better: Simultaneous or Sequential Choice?

The term "diversification bias" refers to the tendency for people to take more variety when choosing several items simultaneously than when choosing them sequentially. In this article, we investigate whether this really is a bias by measuring evaluations of sets chosen simultaneously or sequentially. In Experiment 1 participants made two choices between audio tracks for consecutive consumption. Participants liked low-variety sets most and were more likely to choose high-variety sets in simultaneous choice. In Experiment 2 participants chose between three gambles which varied in the probability of winning and their expected value. Again, simultaneous choices seemed worse than sequential ones: The simultaneous-choice groups took far more low expected value gambles than did sequential-choice subjects and rated their enjoyment as lower. We conclude that simultaneous choice often leads to outcomes that are worse than sequential choice and discuss the circumstances when this is likely to be true.

Journal Article↗
PubMed2005

Greater contribution of N-methyl-D-aspartic acid receptors in ventral compared to dorsal hippocampal slices in the expression and long-term maintenance of epileptiform activity.

Functional segregation along the dorso-ventral axis of the hippocampus is a developing concept. The higher susceptibility of the ventral hippocampus to epileptic activity compared with dorsal hippocampus is one of the main features, which still has obscure mechanisms. Using the model of magnesium-free medium and field recordings, single epileptiform discharges displayed higher incidence (77% vs 57%), rate (41.7+/-3.1 vs 13.5+/-0.7 events/min), duration (173.9+/-17.7 vs 116.8+/-13.6 ms) and intensity (coastline, 25.4+/-2.5 vs 9.5+/-1.8) in ventral compared with dorsal rat hippocampal slices. In addition, the decay phase of the evoked synaptic potentials was 110% slower in ventral slices. The N-methyl-D-aspartate (NMDA) receptor antagonist d-(-)-2-amino-5-phosphonopentanoic acid (50-100 microM) decreased the discharge rate and coastline similarly in ventral and dorsal slices, but it shortened the discharges in ventral slices (by 40%) only. The NMDA receptor antagonist 3-((R)-2-carboxypiperazin-4-yl)-propyl-1-phosphonic acid (10 microM) decreased the rate in both groups and additionally shortened discharges in both kinds of slices, an effect which was greater in ventral ones (31% vs 13%). Furthermore, both drugs shortened the evoked potentials more in ventral (77%) than in dorsal slices (52%). On the other hand, 1 microM of 3-((R)-2-carboxypiperazin-4-yl)-propyl-1-phosphonic acid shortened the discharges and evoked synaptic potentials only in ventral slices, and slowed down the discharge rate only in dorsal slices. Addition of NMDA, in the magnesium-free medium, enhanced activity in both kinds of slices. At 5 and 10 microM of NMDA 51% of the ventral but only 9% of the dorsal slices displayed persistent epileptiform discharges, which were recorded for at least one hour after reintroduction of magnesium in the medium. At 10-20 microM the enhancement of activity was transient, followed by suppression of discharges in 40% and 76% of the ventral and dorsal slices, respectively. Most of the slices having experienced suppression did not develop persistent activity. We propose that the NMDA receptors contribute to the higher susceptibility of the ventral hippocampus to expression and long-term maintenance of epileptiform discharges. This diversification may be related to other aspects of hippocampal dorso-ventral functional segregation.

2-Amino-5-phosphonovalerate↗
PubMed2003

Assemblage of signaling DNA enzymes with intriguing metal-ion specificities and pH dependences.

We report a group of new DNA enzymes that possess a synchronized RNA-cleavage/fluorescence-signaling ability and exhibit wide-ranging metal-ion and pH dependences. These DNA catalysts were derived from a random-sequence DNA pool in a two-stage process: (1) establishment of a catalytic DNA population through repetitive rounds of in vitro selection at pH 4.0, and (2) sequence-diversification and catalytic-activity optimization through five parallel paths of in vitro evolution conducted at pH 3.0, 4.0, 5.0, 6.0, and 7.0, respectively. The deoxyribozymes were evolved to cleave the phosphodiester bond of a single ribonucleotide embedded in DNA and flanked immediately by two deoxyribonucleotides modified with a fluorophore and a quencher, respectively--a setting that synchronizes catalysis with fluorescence signaling. The most dominant catalyst from each pool was examined for metal-ion specificity, catalytic efficiency, pH dependence, and fluorescence-signaling capability. Individual catalysts have different metal-ion requirements and can generate as much as a 12-fold fluorescence enhancement upon RNA cleavage. Most of the DNA enzymes have a pH optimum coinciding with the selection pH and exhibit a rate constant approximating 1 min(-)(1) under optimal reaction conditions. The demonstration of DNA enzymes that are functional under extremely high acidity (such as pH 3 and 4) indicates that DNA has the ability to perform efficient catalysis even under harsh reaction conditions. The isolation of many new signaling DNA enzymes with broad pH optima and metal-ion specificities should facilitate the development of diverse deoxyribozyme-based biosensors.

Base Sequence↗
PubMedDate not supplied

Phylogeny of Saxifragales (angiosperms, eudicots): analysis of a rapid, ancient radiation.

Rapid, ancient radiations pose one of the most difficult challenges for phylogenetic estimation. We used DNA sequence data of 9,006 aligned base pairs from five genes (chloroplast atpB, matK, rbcL, and 18S and 26S nrDNA) to elucidate relationships among major lineages of Saxifragales (angiosperms, eudicots). These relationships were poorly supported in previous studies, apparently because the lineages originated in rapid succession. Using an array of methods that explicitly incorporate assumptions about evolutionary process (weighted maximum parsimony, maximum likelihood, LogDet/paralinear transformed distances), we show that the initial diversification of Saxifragales was indeed rapid. We suggest that the poor resolution of our best phylogenetic estimate is not due to violations of assumptions or to combining data partitions having conflicting histories or processes. We show that estimated branch lengths during the initial diversification are exceedingly short, and we estimate that acquiring sufficient sequence data to resolve these relationships would require an extraordinary effort (approximately 10(7) bp), assuming a linear increase in branch support with branch length. However, our simulation of much larger data sets containing a distribution of phylogenetic signal similar to that of the five sampled gene sequences suggests a limit to achievable branch support. Using statistical tests of differences in the likelihoods of topologies, we evaluated whether the initial radiation of Saxifragales involved the simultaneous origin of major lineages. Our results are consistent with predictions that resolving the branching order of rapid, ancient radiations requires sampling characters that evolved rapidly at the time of the radiation but have since experienced a slower evolutionary rate.

Base Composition↗
PubMed2005

Putting the brakes on snake venom evolution: the unique molecular evolutionary patterns of Aipysurus eydouxii (Marbled sea snake) phospholipase A2 toxins.

Accelerated evolution of toxins is a unique feature of venoms, with the toxins evolving via the birth-and-death mode of molecular evolution. The venoms of sea snakes, however, are remarkably simple in comparison to those of land snakes, which contain highly complex venoms. Aipysurus eydouxii (Marbled sea snake) is a particularly unique sea snake, feeding exclusively upon fish eggs. Secondary to this ecological change, the fangs have been lost and the venom glands greatly atrophied. We recently showed that the only neurotoxin (a three-finger toxin) gene found in the sea snake A. eydouxii has a dinucleotide deletion, resulting in the loss of neurotoxic activity. During these studies, we isolated and identified a number of cDNA clones encoding isozymes of phospholipase A(2) (PLA(2)) toxins from its venom gland. Sixteen unique PLA(2) clones were sequenced from the cDNA library and TA cloning of reverse transcription-polymerase chain reaction products. Phylogenetic analysis of these clones revealed that less diversification of the PLA(2) toxins has occurred in the A. eydouxii venom gland in comparison to equivalent terrestrial and other marine snakes. As there is no longer a positive selection pressure acting upon the venom, mutations have accumulated in the toxin-coding regions that would have otherwise had a deleterious effect upon the ability to use the venom for prey capture. Such mutations include substitutions of highly conserved residues; in one clone, the active site His(48) is replaced by Arg, and in two other clones, highly conserved cysteine residues are replaced. These mutations significantly affect the functional and structural properties of these PLA(2) enzymes, respectively. Thus, in A. eydouxii, the loss of the main neurotoxin is accompanied by a much slower rate of molecular evolution of the PLA(2) toxins as a consequence of the snake's shift in ecological niche. This is the first case of decelerated evolution of toxins in snake venom.

Amino Acid Sequence↗
PubMed1997

Dissecting global diversity patterns: examples from the Ordovician Radiation.

Although the history of life has been characterized by intermittent episodes of radiation that can be recognized in global compilations of biodiversity, it does not necessarily follow that these episodes are caused by processes that occurred uniformly around the world. Major diversity increases could be generated by the cumulative effects of different mechanisms operating simultaneously at several geographic or environmental scales. The purpose of this review is to describe ongoing research on the manifestations, at several scales, of the Ordovician Radiation, which was among the most extensive intervals of diversification in the history of life. Through much of the period, diversity was concentrated most heavily near regions of active mountain building and volcanism; differences in diversity patterns from continent to continent, and among regions within continents, reflect this overprint. While this suggests a linkage of the Radiation and tectonic activity, this is by no means the only mediating agent. Outcrop-based research in North America has demonstrated that tectonic activity was detrimental to some biotic elements, in contrast to its effects on other organisms. Moreover, in the Great Basin of North America where the local stratigraphic record is of particularly high quality, biotic transitions characteristic of the period occurred far more rapidly than observed in global compilations of diversity, suggesting that the global rate of transition may represent the aggregate sum of transitions that occurred abruptly, but at different times, around the world. Finally, it has been demonstrated that, in concert with an increase in average age, the environmental and geographic ranges of Ordovician genera both increased significantly through the period, indicating a role for intrinsic factors in producing Ordovician biotic patterns.

Animals↗
PubMed2005

Evolution of the heart from bacteria to man.

This review provides an overview of the evolutionary path to the mammalian heart from the beginnings of life (about four billion years ago ) to the present. Essential tools for cellular homeostasis and for extracting and burning energy are still in use and essentially unchanged since the appearance of the eukaryotes. The primitive coelom, characteristic of early multicellular organisms ( approximately 800 million years ago), is lined by endoderm and is a passive receptacle for gas exchange, feeding, and sexual reproduction. The cells around this structure express genes homologous to NKX2.5/tinman, and gradual specialization of this "gastroderm" results in the appearance of mesoderm in the phylum Bilateria, which will produce the first primitive cardiac myocytes. Investment of the coelom by these mesodermal cells forms a "gastrovascular" structure. Further evolution of this structure in the bilaterian branches Ecdysoa (Drosophila) and Deuterostoma (amphioxus) culminate in a peristaltic tubular heart, without valves, without blood vessels or blood, but featuring a single layer of contracting mesoderm. The appearance of Chordata and subsequently the vertebrates is accompanied by a rapid structural diversification of this primitive linear heart: looping, unidirectional circulation, an enclosed vasculature, and the conduction system. A later innovation is the parallel circulation to the lungs, followed by the appearance of septa and the four-chambered heart in reptiles, birds, and mammals. With differentiation of the cardiac chambers, regional specialization of the proteins in the cardiac myocyte can be detected in the teleost fish and amphibians. In mammals, growth constraints are placed on the heart, presumably to accommodate the constraints of the body plan and the thoracic cavity, and adult cardiac myocytes lose the ability to re-enter the cell cycle on demand. Mammalian cardiac myocyte innervation betrays the ancient link between the heart, the gut, and reproduction: the vagus nerve controlling heart rate emanates from centers in the central nervous system regulating feeding and affective behavior.

Animals↗