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The pre-Mendelian, pre-Darwinian world: shifting relations between genetic and epigenetic mechanisms in early multicellular evolution.

The reliable dependence of many features of contemporary organisms on changes in gene content and activity is tied to the processes of Mendelian inheritance and Darwinian evolution. With regard to morphological characters, however, Mendelian inheritance is the exception rather than the rule, and neo-Darwinian mechanisms in any case do not account for the origination (as opposed to the inherited variation) of such characters. It is proposed, therefore, that multicellular organisms passed through a pre-Mendelian, pre-Darwinian phase, whereby cells, genes and gene products constituted complex systems with context-dependent, self-organizing morphogenetic capabilities. An example is provided of a plausible 'core' mechanism for the development of the vertebrate limb that is both inherently pattern forming and morphogenetically plastic. It is suggested that most complex multicellular structures originated from such systems. The notion that genes are privileged determinants of biological characters can only be sustained by neglecting questions of evolutionary origination and the evolution of developmental mechanisms.

Animals↗

Limits to optimization: fluid dynamics, adhesive strength and the evolution of shape in limpet shells.

Limpets are commonly found on wave-swept rocky shores, where they may be subjected to water velocities in excess of 20 m s(-1). These extreme flows can impose large forces (lift and drag), challenging the animal's ability to adhere to the substratum. It is commonly thought that the conical shape of limpet shells has evolved in part to reduce these hydrodynamic forces while providing a large aperture for adhesion. This study documents how lift and drag actually vary with the shape of limpet-like models and uses these data to explore the potential of hydrodynamic forces to serve as a selective factor in the evolution of limpet shell morphology. At a low ratio of shell height to shell radius, lift is the dominant force, while at high ratios of height to radius drag is dominant. The risk of dislodgment is minimized when the ratio of height to radius is 1.06 and the apex is in the center of the shell. Real limpets are seldom optimally shaped, however, with a typical height-to-radius ratio of 0.68 and an apex well anterior of the shell's center. The disparity between the actual and the hydrodynamically optimal shape of shells may be due to the high tenacity of limpets' adhesive system. Most limpets adhere to the substratum so strongly that they are unlikely to be dislodged by lift or drag regardless of the shape of their shell. The evolution of a tenacious adhesion system (perhaps in response to predation) has thus preempted selection for a hydrodynamically optimal shell, allowing the shell to respond to alternative selective factors.

Adaptation, Physiological↗

Emerging trends in the study of spiralian larvae.

Many animals undergo indirect development, where their embryogenesis produces an intermediate life stage, or larva, that is often free-living and later metamorphoses into an adult. As their adult counterparts, larvae can have unique and diverse morphologies and occupy various ecological niches. Given their broad phylogenetic distribution, larvae have been central to hypotheses about animal evolution. However, the evolution of these intermediate forms and the developmental mechanisms diversifying animal life cycles are still debated. This review focuses on Spiralia, a large and diverse clade of bilaterally symmetrical animals with a fascinating array of larval forms, most notably the archetypical trochophore larva. We explore how classic research and modern advances have improved our understanding of spiralian larvae, their development, and evolution. Specifically, we examine three morphological features of spiralian larvae: the anterior neural system, the ciliary bands, and the posterior hyposphere. The combination of molecular and developmental evidence with modern high-throughput techniques, such as comparative genomics, single-cell transcriptomics, and epigenomics, is a promising strategy that will lead to new testable hypotheses about the mechanisms behind the evolution of larvae and life cycles in Spiralia and animals in general. We predict that the increasing number of available genomes for Spiralia and the optimization of genome-wide and single-cell approaches will unlock the study of many emerging spiralian taxa, transforming our views of the evolution of this animal group and their larvae.

Animals↗

Body proportions of Australopithecus afarensis and A. africanus and the origin of the genus Homo.

New discoveries of A. africanus fossils from Member 4 Sterkfontein reveal a body form quite unlike earlier Australopithecus species. The new adult material consists of over 48 fore- and hindlimb specimens and includes an associated partial skeleton, Stw 431. The forelimbs and relatively large: the average size of their joints corresponds to a modern human with body mass of 53 kg. The hindlimbs are much smaller with an average size matching a modern human of only 33 kg. Analyses of the Stw 431 partial skeleton confirm these results. In contrast, A. afarensis and anamensis more closely approximate a human pattern of forelimb joint size. This is an unanticipated complication in our understanding of early human evolution. In general, craniodental morphology tracks time in species of Australopithecus: A. anamensis (3.5-4.1 Ma) is the the most primitive with a strongly sloping symphysis, large canine roots, etc., A. afarensis (3.0-3.6 Ma) is less primitive, and A. africanus (2.6-3.0 Ma) shares many derived characteristics with early Homo (e.g., expanded brain, reduced canine, bicuspid lower third premolar, reduced prognathism, greater flexion of the cranial base, deeper TMJ). the new postcranial material, however, reveals an apparently primitive morphology of relatively large forelimb and small hindlimb joints resembling more the pongid than the human pattern. More pongid-like proportions are also present in the two known associated partial skeletons of H. habilis (OH 62 KNM-ER 3735). This may imply either (1) that A. africanus and H. habilis evolved craniodental characters in parallel with the lineage leading to later Homo, or (2) that fore- to hindlimb proportions of A. afarensis (and perhaps A. anamensis) evolved independent of the lineage leading to Homo and does not imply a close phylogenetic link with Homo. Both of these explanations or any other phylogeny imply homoplasy.

Adult↗

[Chronic myeloproliferative disorders. The new WHO classification].

Except for chronic myelogenous leukemia (CML), chronic myeloproliferative disorders (CMPDs) include as main subtypes polycythemia vera (PV), chronic idiopathic myelofibrosis (IMF), and essential thrombocythemia (ET). A common finding in CMPDs is a clonal evolution associated with a significantly variable course, which may be complicated by thrombocythemia, (secondary) myelofibrosis, and finally acceleration (unstable phase) that merges into blastic crisis. New therapeutic modalities (chemo- and interferon therapy, bone marrow and stem cell transplantation) which were developed in the last decade and the striking differences in survival amongst the different subtypes warrant not only an unequivocal distinction from reactive and allied disorders, but a clear-cut classification as well. For this reason, a synoptical approach is essential including clinical data and, as a major diagnostic tool, a bone marrow biopsy. This concept finds expression in the new WHO classification, which also includes as rare subtypes chronic neutrophilic leukemia, eosinophilic leukemia, chronic hypereosinophilic syndrome, and finally unclassifiable entities. Histopathology of bone marrow biopsies reveals specific findings, in particular concerning megakaryopoiesis, which are characteristic for the different subtypes. These features facilitate the still controversially discussed differentiation of thrombocythemia that is frequently present, as is the case in initial (prefibrotic) IMF from ET. Moreover, in addition to clinical findings,the associated heterogeneity of bone marrow morphology indicates a stepwise evolution of the disease process and thus exerts a significant impact on survival, i.e., in CML regarding erythropoiesis and myelofibrosis and in IMF extent of myeloid metaplasia.

Bone Marrow↗

5500 Phages examined in the electron microscope.

"Phages" include viruses of eubacteria and archaea. At least 5568 phages have been examined in the electron microscope since the introduction of negative staining in 1959. Most virions (96%) are tailed. Only 208 phages (3.7%) are polyhedral, filamentous, or pleomorphic. Phages belong to one order, 17 families, and three "floating" groups. Phages are found in 11 eubacterial and archaeal phyla and infect 154 host genera, mostly of the phyla Actinobacteria, Firmicutes, and Proteobacteria. Of the tailed phages, 61% have long, noncontractile tails and belong to the family Siphoviridae. Convergent evolution is visible in the morphology of certain phage groups.

Bacteria↗

The origins of genome complexity.

Complete genomic sequences from diverse phylogenetic lineages reveal notable increases in genome complexity from prokaryotes to multicellular eukaryotes. The changes include gradual increases in gene number, resulting from the retention of duplicate genes, and more abrupt increases in the abundance of spliceosomal introns and mobile genetic elements. We argue that many of these modifications emerged passively in response to the long-term population-size reductions that accompanied increases in organism size. According to this model, much of the restructuring of eukaryotic genomes was initiated by nonadaptive processes, and this in turn provided novel substrates for the secondary evolution of phenotypic complexity by natural selection. The enormous long-term effective population sizes of prokaryotes may impose a substantial barrier to the evolution of complex genomes and morphologies.

Alleles↗

Evolution of eye size and shape in primates.

Strepsirrhine and haplorhine primates exhibit highly derived features of the visual system that distinguish them from most other mammals. Comparative data link the evolution of these visual specializations to the sequential acquisition of nocturnal visual predation in the primate stem lineage and diurnal visual predation in the anthropoid stem lineage. However, it is unclear to what extent these shifts in primate visual ecology were accompanied by changes in eye size and shape. Here we investigate the evolution of primate eye morphology using a comparative study of a large sample of mammalian eyes. Our analysis shows that primates differ from other mammals in having large eyes relative to body size and that anthropoids exhibit unusually small corneas relative to eye size and body size. The large eyes of basal primates probably evolved to improve visual acuity while maintaining high sensitivity in a nocturnal context. The reduced corneal sizes of anthropoids reflect reductions in the size of the dioptric apparatus as a means of increasing posterior nodal distance to improve visual acuity. These data support the conclusion that the origin of anthropoids was associated with a change in eye shape to improve visual acuity in the context of a diurnal predatory habitus.

Animals↗

Renal injury and proteinuria in female spontaneously hypertensive rats.

This investigation describes the evolution of functional and morphological changes in the kidney of female spontaneously hypertensive rats (SHR; 5-90 weeks) and control animals of the same genetic strain, Wistar-Kyoto (WKY). Systolic blood pressure in SHR was 162.0 +/- (SD) 14.1 mm Hg up to 45 weeks, then increased to 189.4 +/- (SD) 16.6 mm Hg (50-90 weeks). Blood pressure in WKY controls remained constant (117.8 +/- 13.6 mm Hg, 5-100 weeks). Protein excretion in the SHR was higher than controls (1.5 +/- 0.5 mg/24 h x 100 g BW, 5-100 weeks) from week 30 on. At 90 weeks, excretion rose to 26.1 +/- 9.8 mg/24 h x 100 g BW and is predominately an albuminuria. After 70 weeks superficial tubular fluid albumin concentration in the SHR is significantly increased (p less than 0.001) over controls. Microscopic changes were evident in glomeruli, tubules, interstitium and arterial vessels at 72-90 weeks and demonstrated an increasing gradient of severity from outer to inner cortex. Glomeruli show sclerosis, fibrinoid necrosis and pericapsular fibrosis. Glomeruli studied with a polyanionic stain revealed a marked decrease in staining affinity of deep, compared to superficial glomeruli. The arterial lesions consisted of thickening of the intima and hyperplasia of the media. WKY kidneys were unremarkable. The selectivity of renal injury and proteinuria in the female SHR is similar to that in the male SHR. However, the evolution of these changes is delayed and may be related to a lower blood pressure until 50 weeks of age.

Age Factors↗

[DNA-DNA hybridization in several species of Hansenula].

The genus Hansenula was considered a long time ago as a good pattern for phylogenetic research. In 1969, Wickerham proposed an evolutive scheme based upon morphological, physiological and ecological criteria. Recently, relatedness among yeasts were analysed by DNA-DNA hybridization in liquid medium. H. anomala var. anomala (G + C content: 37.1%) was compared with H. anomala var. schneggii (37.6%), H. subpelliculosa (33.8%) line 3, H. sydowiorum (40.1%) and H. muscicola (37.1%). These results showed little relatedness between H. anomala var. anomala/H. ciferrii and H. anomala var. anomala/H. subpelliculosa. On the other hand, H. anomala var. schneggii shared 89.5% of its nucleotide sequences with H. anomala var. anomala. These 2 strains were considered to represent the same species. H. holstii showed 67.1% complementarity with H. anomala var. anomala: this strain is considered to represent valid species, different from H. anomala var. anomala, but H. muscicola with 72.5% relatedness to H. anomala var. anomala could be considered as a 'limit species'. An unexpected finding was that H. beckii was closely related to H. anomala var. anomala (84.8%). These data suggested the inadequacy of current criteria used to establish the phylogenetic lines in genus Hansenula.

Ascomycota↗

The evolution of placental mammals.

Based on morphological, virological, biochemical and molecular biological data, it is proposed that the presence of endogenous retrovirus particles in the placental cytotrophoblasts of many mammals is indicative of some beneficial action provided by the virus in relation to cell fusion, syncytiotrophoblast formation and the creation of the placenta. Further, it is hypothesised that the germ line retroviral infection of some primitive mammal-like species resulted in the evolution of the placental mammals.

Animals↗

Behavior, body types and the irreversibility of evolution.

A functional approach to evolutionary morphology is emphasized in this paper. This perspective differs from the current 'structuralist' trend, which emphasizes the constraining role of developmental paths. In addition, the present approach agrees with the adaptationist paradigm. It is further argued that three types of phenomena are better understood in this light: i. The existence of evolutionary trends, ii. The maintenance of certain structural features within a given taxon, and iii. The irreversibility of evolution.

Animals↗

Quantitative genetics and developmental constraints on evolution by selection.

It has often been argued that the principles of random mutation and selection are insufficient to account for macroevolutionary phenomena, such as the origin of morphological novelty and directionality in evolution. A third, epigenetic, principle is said to be required and this principle is thought not to be included in microevolutionary theory. The third principle has most recently been identified as internal selection and/or non-random phenotypic effects of mutation. It is shown that the genetic variance/covariance matrix of quantitative genetic theory measures developmental constraints due to internal selection and non-random mutation. The genetic variance/covariance matrix causes the response to selection to deviate from the optimal rate and direction as specified by the selection gradient, which measures direct selection on the phenotypes. Therefore, microevolutionary theory takes account of developmental constraints on evolution by natural selection through the genetic variance/covariance matrix. Theories for predicting the pattern of genetic variance and covariance from stabilizing selection and the phenotypic effects of mutation are discussed.

Biological Evolution↗

Shapes in the shadow: evolutionary dynamics of morphogenesis.

This article investigates the evolutionary dynamics of morphogenesis. In this study, morphogenesis arises as a side-effect of maximization of number of cell types. Thus, it investigates the evolutionary dynamics of side-effects. Morphogenesis is governed by the interplay between differential cell adhesion, gene-regulation, and intercellular signaling. Thus, it investigates the potential to generate complex behavior by entanglement of relatively "boring" processes, and the (automatic) coordination between these processes. The evolutionary dynamics shows all the hallmarks of evolutionary dynamics governed by nonlinear genotype phenotype mapping: for example, punctuated equilibria and diffusion on neutral paths. More striking is the result that interesting, complex morphogenesis occurs mainly in the "shadow" of neutral paths which preserve cell differentiation, that is, the interesting morphologies arise as mutants of the fittest individuals. Characteristics of the evolution of such side-effects in the shadow appear to be the following: (1) The specific complex morphologies are unique (or at least very rare) among the set of de novo initiated evolutionary histories. (2) Similar morphologies are reinvented at large temporal distances during one evolutionary history and also when evolution is restarted after the main cell differentiation pattern has been established. (3) A mosaic-like evolution at the morphological level, where different morphological features occur in many combinations, while at the genotypic level recombination is not implemented and genotypes diverge linearly and at a constant rate.

Animals↗

Comprehensive analysis of genomic alterations in gliosarcoma and its two tissue components.

Gliosarcoma is a variant of glioblastoma multiforme characterized by two components displaying gliomatous or sarcomatous differentiation. We investigated 38 gliosarcomas for aberrations of tumor-suppressor genes and proto-oncogenes that are commonly altered in glioblastomas. Amplification of CDK4, MDM2, EGFR, and PDGFRA were found in 11% (4/35), 8% (3/38), 8% (3/38), and 3% (1/35) of the tumors, respectively. Nine of 38 gliosarcomas (24%) carried TP53 mutations. PTEN mutations were identified in 45% (9/20) of the investigated tumors. Twenty gliosarcomas were analyzed by comparative genomic hybridization (CGH). Chromosomal imbalances commonly detected were gains on chromosomes 7 (15/20; 75%), X (4/20; 20%), 9q, and 20q (3/20, 15% each); and losses on chromosomes 10 and 9p (7/20, 35% each), and 13q (3/20, 15%). Five different high-level amplifications were mapped to 4q12-q21 (1 case), 6p21 (1 case), 7p12 (2 cases), proximal 12q (4 cases), and 14q32 (1 case) by CGH. Southern blot and/or differential PCR analyses identified amplification of PDGFRA (4q12), CCND3 (6p21), EGFR (7p12), CDK4 (12q14) and/or MDM2 (12q14.3-q15), and AKT1 (14q32.3) in the respective tumors. Separate analysis of the gliomatous and sarcomatous components of eight gliosarcomas by CGH after microdissection and universal DNA amplification revealed that both components shared 57% of the chromosomal imbalances detected. Taken together, our data indicate that the genomic changes in gliosarcomas closely resemble those found in glioblastomas. However, the number of chromosomes involved in imbalances in gliosarcomas was significantly lower than that in glioblastomas, indicating a higher genomic stability in gliosarcomas. In addition, we provide further support for the hypothesis that the gliomatous and sarcomatous components are derived from a single precursor cell clone, which progressed into subclones with distinct morphological features during tumor evolution. According to our data, gain/amplification of genes on proximal 12q may facilitate the development of a sarcomatous phenotype.

Adult↗

Differential effects of mate competition and mate choice on eastern tiger salamanders

Male tiger salamanders, Ambystoma tigrinum tigrinumare slightly larger in body size and have considerably higher and longer tails than females. To determine how these dimorphic traits affected reproductive performance and success, we conducted breeding trials using 12 males and six females per trial and monitored male-female and male-male interactions. Larger males had an advantage in most aspects of mate competition investigated. Males with higher tails had no advantage in either mate competition or mate choice. Males with longer tails also had no advantage in mate competition but were preferred as mates by females. Larger males interrupted courting males more often than smaller males did. The form of male-male interference was conditional on body size and not on either tail dimension. If the intruder was larger than the courting male, it would shove the female away from the courting male and initiate courtship; if the intruder was smaller, it adopted a female mimicry tactic in which it positioned itself between the courting male and female and performed female behaviours to the courting male while simultaneously courting the female. Our trials indicated that the two components of sexual selection may influence the evolution of different male morphological traits in tiger salamanders. Mate competition may favour increased male body length; mate choice may select for greater male tail length.

Journal Article↗

Recovery of the right ventricle after single-lung transplantation in pulmonary hypertension.

Single-lung transplantation has been successfully performed in patients with pulmonary fibrosis and emphysema. In contrast, patients with end-stage pulmonary hypertension (either primary or secondary to Eisenmenger's syndrome) have conventionally been offered heart-lung transplantation. The rationale underlying this approach is that chronic pulmonary hypertension results in irreversible right ventricular dilatation and failure. Recovery of the right ventricle has previously been reported after thromboendarterectomy for chronic large-vessel pulmonary embolism, correction of atrial septal defect or mitral valve replacement. The evolution of right ventricular morphology and function after lung transplantation has not been previously described. This study examines the reversibility of right ventricle dysfunction following normalization of pulmonary artery pressure after single-lung transplantation in 4 patients with pulmonary hypertension. Cardiac function was assessed using electrocardiography, echocardiography and radionuclide angiography. Pulmonary hemodynamic measurements, including pulmonary artery pressure and pulmonary vascular resistance, decreased in all patients after single-lung transplantation. Electrocardiographic changes observed were leftward shift in the QRS axis, and a decrease in P-wave amplitude and in right ventricular force. Echocardiographic examination revealed decreased right atrial, right ventricular and tricuspid valve annular dimensions, normalization of septal motion, and decreased tricuspid regurgitation. Thus, improved pulmonary hemodynamics after single-lung transplantation for pulmonary vascular disease results in reversal of right heart dilatation and dysfunction, and improved myocardial performance. The extent of right ventricular dysfunction beyond which recovery is unlikely to occur has yet to be determined.

Adult↗

The evolution of experimental Trypanosoma cruzi cardiomyopathy in rabbits: further parasitological, morphological and functional studies.

Young rabbits (1-2 months of age) inoculated with trypomastigote forms of the Colombia strain of Trypanosoma cruzi have been shown to develop cardiac pathological changes (together with parasitological and immunological alterations) which are very similar to those observed in the acute and chronic phases of Chagas' disease in man. The cardiac alterations in the acute phase are characterized grossly by slight cardiomegaly with dilatation of the right-sided chambers. Microscopically they are characterized by mild focal myocarditis. The chronic phase is characterized by moderate to marked cardiomegaly with hypertrophy and dilatation of both ventricular chambers. There is thinning of the apical region (apical aneurysm), particularly of the left ventricle. Focal myocarditis is seen microscopically with areas of myocytolytic necrosis, atrophic and hypertrophic myofibers, an inflammatory response predominantly composed of mononuclear cells and interstitial fibrosis. Cineventriculography in the left ventricle of rabbits during the chronic phase disclosed regional myocardial dysfunction, with typical apical systolic bulging. The pathogenesis of Chagasic cardiomyopathy is briefly discussed in the light of these findings. Our investigation has further shown that this animal model is particularly suitable for studies on on the mechanisms, pathology and treatment of Chagas' heart disease.

Actin Cytoskeleton↗