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Vocal mechanics in Darwin's finches: correlation of beak gape and song frequency.

Recent studies of vocal mechanics in songbirds have identified a functional role for the beak in sound production. The vocal tract (trachea and beak) filters harmonic overtones from sounds produced by the syrinx, and birds can fine-tune vocal tract resonance properties through changes in beak gape. In this study, we examine patterns of beak gape during song production in seven species of Darwin's finches of the Galápagos Islands. Our principal goals were to characterize the relationship between beak gape and vocal frequency during song production and to explore the possible influence therein of diversity in beak morphology and body size. Birds were audio and video recorded (at 30 frames s(-1)) as they sang in the field, and 164 song sequences were analyzed. We found that song frequency regressed significantly and positively on beak gape for 38 of 56 individuals and for all seven species examined. This finding provides broad support for a resonance model of vocal tract function in Darwin's finches. Comparison among species revealed significant variation in regression y-intercept values. Body size correlated negatively with y-intercept values, although not at a statistically significant level. We failed to detect variation in regression slopes among finch species, although the regression slopes of Darwin's finch and two North American sparrow species were found to differ. Analysis within one species (Geospiza fortis) revealed significant inter-individual variation in regression parameters; these parameters did not correlate with song frequency features or plumage scores. Our results suggest that patterns of beak use during song production were conserved during the Darwin's finch adaptive radiation, despite the evolution of substantial variation in beak morphology and body size.

Acoustics↗

Ontogenies in mice selected for high voluntary wheel-running activity. I. Mean ontogenies.

The evolutionary importance of postnatal ontogenies has long been recognized, but most studies of ontogenetic trajectories have focused exclusively on morphological traits. For animals, this represents a major omission because behavioral traits and their ontogenies often have relatively direct relationships to fitness. Here four replicate lines of house mice artificially selected for high early-age wheel running and their four replicate control lines were used to evaluate the effects of early-age directional selection, genetic drift, and activity environment (presence or absence of a running wheel) on variation in the ontogenies of three traits known to be genetically correlated: voluntary wheel running, body mass, and food consumption. Early-age selection significantly changed both the shape and position of the wheel-running and food-consumption ontogenies while influencing the position, but not the shape, of the body mass ontogeny. Genetic drift (as indicated by variation among replicate lines) produced significant changes in both the position and shape of all three ontogenies; however, its effect differed between the selection and control groups. For wheel running and food consumption, genetic drift only influenced the control ontogenies, whereas for body mass, genetic drift had a significant effect in both selection groups. Both body-mass and food-consumption ontogenies were significantly altered by activity environment, with the environment causing significant changes in the shape and position of both ontogenies. Overall the results demonstrate strong effects of early-age selection, genetic drift, and environmental variation on the evolution and expression of behavioral and morphological ontogenies, with selection changing only the position of the morphological ontogeny but both the position and shape of the behavioral ontogenies.

Analysis of Variance↗

The intermetatarsal articular facet of the first metatarsal bone in humans: a derived trait unique within primates.

The occurrence and morphology of an intermetatarsal facet of the first metatarsal bone have been investigated in a series of 306 nonhuman primates representative of 40 genera, and in a series of 412 human metatarsal bones (dried bones) (215 left, 197 right). In nonhuman primates, no case of intermetatarsal facet was observed in the 306 first metatarsal bones studied. In humans, a well-defined intermetatarsal facet was observed in 127 out of the 412 bones (30.8%); no significant difference was observed between the left and right sides. The shape of the facet was elliptical, more or less elongated; it had a mean major axis (height) of 10.7 mm, and a mean minor axis (width) of 6.1 mm (extremes: 5 x 3 mm and 17 x 10 mm). The facet was located in the dorsal third of the lateral side of the first metatarsal in 103 out of 127 cases (81.1%), and in the middle third in 24 out of 127 cases (18.9%); it was never observed in the plantar third. The intermetatarsal facet was in connection with the proximal articular facet for the medial cuneiform in 68 out of 127 cases (53.5%), and it was separated from the proximal facet by a small nonarticular area or groove in 59 out of 127 cases (46.5%). The present observations suggest that the human first intermetatarsal facet is a derived trait unique within primates (autapomorphy) which is present in approximately one third of individuals. The appearance of this new articular facet in human evolution seems clearly related to the morphological modifications of the foot due to the acquisition of bipedalism, and more particularly to the loss of abductability of the hallux and its permanent adduction.

Animals↗

Evolution of the brain developmental plan: Insights from agnathans.

In vertebrate evolution, the brain exhibits both conserved and unique morphological features in each animal group. Thus, the molecular program of nervous system development is expected to have experienced various changes through evolution. In this review, we discuss recent data from the agnathan lamprey (jawless vertebrate) together with available information from amphioxus and speculate the sequence of changes during chordate evolution that have been brought into the brain developmental plan to yield the current variety of the gnathostome (jawed vertebrate) brains.

Animals↗

[Early reception of Darwin's selection theory and its sequelae for comparative morphology today].

It is argued that Darwin's concept of evolutionary change is primarily based on the idea of functional adaptation. Genealogical aspects are seen as a secondary consequence of this hypothesis. Unfortunately, the reception of Darwin's work was concentrated on the genealogical aspects from the very beginning (Huxley, Haeckel) and thus channeled future development of evolutionary morphology in a very one-sided way. This direction of development led to the adoption of cladism as a very sophisticated concept of comparative morphology. Though cladism claims to contribute to our understanding of evolution, it is demonstrated that it suffers in this regard because of the incompatibility of "pure morphology" with the demands of functional thinking as an integrative part of Darwin's proposition.

Anatomy, Comparative↗

Modularity and intrinsic evolvability of Hsp90-buffered change.

Hsp90 controls dramatic phenotypic transitions in a wide array of morphological features of many organisms. The genetic-background dependence of specific abnormalities and their response to laboratory selection suggested Hsp90 could be an 'evolutionary capacitor', allowing developmental variation to accumulate as neutral alleles under normal conditions and manifest selectable morphological differences during environmental stress. The relevance of Hsp90-buffered variation for evolution has been most often challenged by the idea that large morphological changes controlled by Hsp90 are unconditionally deleterious. To address this issue, we tested an Hsp90-buffered abnormality in Drosophila for unselected pleiotropic effects and correlated fitness costs. Up to 120-fold differences in penetrance among six highly related selection lines, started from an initially small number of flies and rapidly selected for and against a deformed eye trait (dfe), did not translate into measurable differences in any of several tests of viability, lifespan or competitive fitness. Nor were 17 dfe Quantitative Trait Loci (QTL) associated with fitness effects in over 1,400 recombinant lines. Our ability to detect measurable effects of inbreeding, media environment and the white mutation in the selection line backgrounds independent of dfe penetrance suggests that, within the limitations of laboratory tests of fitness, this large morphological change controlled by Hsp90 was selectable independent of strong, correlated and unconditionally deleterious effects--abundant, polygenic variation hidden by Hsp90 allows potentially deleterious alleles to be readily replaced during selection by less deleterious alleles with similar phenotypic effects. Hsp90 links environmental stress with the expression of developmental variation controlling unprecedented morphological plasticity. As outlined here and in the companion paper of this issue, the complex genetic architecture of Hsp90-buffered variation supports a remarkable modularity of Hsp90 effects on quantitative and qualitative phenotypes, consistent with the 'Hsp90 capacitor hypothesis' and standard quantitative genetic models of threshold traits.

Alleles↗

Miniaturization and its effects on cranial morphology in plethodontid salamanders, genus Thorius (Amphibia, Plethodontidae): II. The fate of the brain and sense organs and their role in skull morphogenesis and evolution.

Relative size and arrangement of the brain and paired sense organs are examined in three species of Thorius, a genus of minute, terrestrial salamanders that are among the smallest extant tailed tetrapods. Analogous measurements of representative species of three related genera of larger tropical (Pseudoeurycea, Chiropterotriton) and temperate (Plethodon) salamanders are used to identify changes in gross morphology of the brain and sense organs that have accompanied the evolution of decreased head size in Thorius and their relation to associated changes in skull morphology. In adult Thorius, relative size (area measured in frontal plane, and length) of the eyes, otic capsules, and brain each is greater than in adults of all of the larger genera; relative size of the nasal capsules is unchanged or slightly smaller. Interspecific scaling phenomena--negative allometry of otic capsule, eye and brain size, isometry or slight positive allometry of nasal capsule size, all with respect to skull length--also are characteristic of intraspecific (ontogenetic) comparisons in both T. narisovalis and Pseudoeurycea goebeli. Predominance of the brain and eyes in Thorius results in greater contact and overlap among these structures and the nasal capsules in the anterior portion of the head. This is associated with anterior displacement of both the eyes and nasal capsules, which now protrude anterior to the skull proper; a change in eye shape; and medial deformation of anterior braincase walls. Posteriorly, predominance of the otic capsules has effected a reorientation of the jaw suspensorium to a fully vertical position that is correlated with the novel presence of a posteriorly directed squamosal process and shift in origin of the quadropectoralis muscle. Many of these changes in cranial morphology may be explained simply as results of mechanical (physical) interactions among the skeletal, nervous, and sensory components during head development at reduced size. This provides further evidence of the role of nervous, sensory, and other "soft" tissues in cranial skeletal morphogenesis, and reinforces the need to consider these tissues in analyses of skull evolution.

Animals↗

Duplication events and the evolution of segmental identity.

Duplication of genes, genomes, or morphological structures (or some combination of these) has long been thought to facilitate evolutionary change. Here we focus on studies of the teleost fishes to consider the conceptual similarities in the evolutionary potential of these three different kinds of duplication events. We review recent data that have confirmed the occurrence of a whole-genome duplication event in the ray-finned fish lineage, and discuss whether this event may have fuelled the radiation of teleost fishes. We then consider the fates of individual duplicated genes, from both a theoretical and an experimental viewpoint, focusing on our studies of teleost Hox genes and their functions in patterning the segmented hindbrain. Finally, we consider the duplication of morphological structures, once again drawing on our experimental studies of the hindbrain, which have revealed that experimentally induced duplicated neurons can produce functionally redundant neural circuits. We posit that the availability of duplicated material, independent of its nature, can lead to functional redundancy, which in turn enables evolutionary change.

Animals↗

Mammalian sex chromosomes: evolution of organization and function.

Comparisons of chromosome size, morphology and gene arrangements between mammals of different species permit us to deduce the genome characteristics of the common ancestor, and to chart the changes that have occurred during the divergence of the two lineages. The more distantly related are the species compared, the more remote the common ancestor whose characteristics can be deduced. This means that, providing there are sufficient similarities to warrant comparison, the more divergent the species compared, the more significant the contribution to our understanding of the organization of an ancestral mammalian genome and the process of mammalian genome evolution. One of the genetic surprises of the last decade was the discovery that, although gross karyotypes of distantly related orders of eutherian mammals (e.g. cat, cow, rabbit, man) have diverged extensively, gene mapping studies reveal the presence of large chromosome segments conserved across at least 60 million years (O'Brien et al. 1988). This finding makes it worthwhile to extend genetic comparisons to the two groups of mammals most distantly related to eutherian mammals--marsupials and monotremes. Here we will review comparisons of the sex chromosomes in these three major groups of extant mammals, and show how they have led us to a new view of the evolution of mammalian sex chromosome organization and function in sex determination and X chromosome inactivation.

Animals↗

Ductal invasive G2 and G3 carcinomas of the breast are the end stages of at least two different lines of genetic evolution.

Ductal invasive grade (G) 2 and G3 carcinomas represent the majority of invasive breast cancers. Previous morphological and cytogenetic studies have provided evidence that ductal invasive G2 carcinoma may originate from at least two different genetic pathways. The aim of this study was to evaluate further the heterogeneity of G2 breast cancer in comparison with G3 cancers by cytogenetic and quantitative analysis. To this end, 35 cases of ductal invasive G2 and 42 cases of ductal invasive G3 carcinomas were investigated by means of comparative genomic hybridization (CGH) and these findings were correlated with DNA ploidy status, mitotic activity index (MAI), mean nuclear area (MNA), volume per lumen (VPL), and clinico-pathological parameters. The findings of this study demonstrate that ductal invasive G2 carcinomas, in contrast to ductal invasive G3 carcinomas, have to be interpreted as the morphological end stage resulting from two different cytogenetic and morphological pathways; the loss of 16q material is the cytogenetic key event in the evolution of a subgroup of this entity. By correlating genetic alterations with DNA ploidy status, an extended morphology-based cytogenetic progression model is presented, with early and late genetic alterations in the pathogenesis of breast cancer. The correlation with MAI gives rise to the hypothesis that these different genetic pathways significantly differ in their proliferation rate. Further studies will be required to elucidate which genes contribute to an altered proliferation rate in these subgroups and to the associated prognosis.

Breast Neoplasms↗

Spatial memory and hippocampal pallium through vertebrate evolution: insights from reptiles and teleost fish.

The forebrain of vertebrates shows great morphological variation and specialized adaptations. However, an increasing amount of neuroanatomical and functional data reveal that the evolution of the vertebrate forebrain could have been more conservative than previously realized. For example, the pallial region of the teleost telencephalon contains subdivisions presumably homologous with various pallial areas in amniotes, including possibly a homologue of the medial pallium or hippocampus. In mammals and birds, the hippocampus is critical for encoding complex spatial information to form map-like cognitive representations of the environment. Here, we present data showing that the pallial areas of reptiles and fish, previously proposed as homologous to the hippocampus of mammals and birds on an anatomical basis, are similarly involved in spatial memory and navigation by map-like or relational representations of the allocentric space. These data suggest that early in vertebrate evolution, the medial pallium of an ancestral fish group that gave rise to the extant vertebrates became specialized for processing and encoding complex spatial information, and that this functional trait has been retained through the evolution of each independent vertebrate lineage.

Animals↗

Composite lymphoma and related disorders.

Lymphomas evolve over time, usually from the small-cell to the large-cell category, and from follicular histologic characteristics to diffuse architecture. The histologic characteristics of lymphomas may be discordant in patients with multiple sites of involvement. When different types of lymphomas are encountered in a single organ or tissue, they are designated as CLs in the Working Formulation of NHL. The most common CLs consist of different subsets of follicular center cell lymphomas, usually one with low-grade follicular histologic characteristics and another with diffuse architecture and/or more aggressive cytologic features. These lymphomas now are considered to represent different phases of clonal evolution rather than representing a coincidental simultaneous occurrence of two unrelated lymphomas. Whether B-cell large-cell lymphoma coexisting with another B-cell lymphoma, namely, the nodular variant of lymphocyte-predominant Hodgkin's disease, also represents clonal evolution requires further study. Composite lymphomas consisting of a B-cell lymphoma and a T-cell lymphoma are extremely rare. The histogenetic implications for a clonal relationship between component subsets in these and other equally rare combinations remain uncertain. The CLs should continue to be recognized because (1) the component morphologic subsets may have entirely different natural histories, requiring different treatment modalities although they may be clonally related; and (2) the study of such cases may provide us with information regarding the complex interrelationship of the lymphoid system and its clonal evolution. The morphologic definition for CL of the Working Formulation should continue to be used whenever possible, with addition of appropriate immunologic and/or molecular-genetic data.

Hodgkin Disease↗

A multidisciplinary approach reveals cryptic diversity in Western Palearctic Tetramorium ants (Hymenoptera: Formicidae).

Diversity of ants of the Tetramorium caespitum/impurum complex was investigated in a multidisciplinary study. Focusing on morphologically hardly distinguishable Western Palearctic samples, we demonstrate the genetic and phenotypic diversity, demarcate phylogenetic entities, and discuss the clades in terms of biogeography. Sequences of 1113bp of the mitochondrial COI gene revealed 13 lineages. COII data, worker morphometry and male genitalia morphology corroborated the COI results for seven lineages; the remaining six were disregarded because of small sample size. A comparison with published data on cuticular hydrocarbons showed correspondence. The seven entities show different distribution patterns, though some ranges overlap in Central Europe. Since no major discrepancy between the results of the different disciplines became apparent, we conclude that the seven entities within the T. caespitum/impurum complex represent seven species. Geographical evidence allows the identification of T. caespitum and T. impurum, and we therefore designate neotypes and redescribe the two species in terms of morphology and mtDNA. As the revision of about 50 taxon names would go beyond the scope of this study, we refer to the remaining five species under code names. We discuss our findings in terms of plesiomorphy and convergent evolution by visualizing the mtDNA phylogeny in morphological space.

Animals↗

[Conjunctival changes related to topical treatment and effectiveness of surgery in glaucoma].

PURPOSE: The topical treatment for a long run in glaucoma rises histopathological changes of conjunctiva. The study is aimed at assessing the correlations between conjunctival changes bound of the type and duration of medical glaucoma therapy and the efficiency of subsequent filtering surgery. METHOD: For this purpose, we carried out a three years clinical prospective study (01.05.2000-30.04.2003), with a mean follow-up period of 18.5 months, enrolling 22 cases (38 eyes) operated by trabeculectomy for primary open angle glaucoma (POAG), with the same technique and the same surgeon. There were excluded both the patients with previous ophthalmic surgical interventions and ocular/systemic chronic inflammatory disease. The cases were divided according to type and duration of topical treatment. There were recorded the conjunctival changes (by histological exam and immunohistochemistry), dynamic evolution of intraocular pressure (IOP) and the morphological aspects of filtering blebs. Statistical analysis was performed by Wilcoxon test. RESULTS: In all groups, the efficiency of filtering surgery was correlated with the duration of topical medication; the survey presents the correlations between topical treatment, conjunctival changes and postoperative evolution. CONCLUSIONS: These results suggest that exhaustive topical treatment in POAG, for a long period, with many drugs may represent a negative prognostic factor for the subsequent filtering surgery. Therefore it is necessary an individual approach for the first line therapy in POAG.

Administration, Topical↗

Functional morphology of jaw trabeculation in the lesser electric ray Narcine brasiliensis, with comments on the evolution of structural support in the Batoidea.

The design of minimum-weight structures that retain their integrity under dynamic loading regimes has long challenged engineers. One solution to this problem found in both human and biological design is the optimization of weight and strength by hollowing a structure and replacing its inner core with supportive struts. In animals, this design is observed in sand dollar test, avian beak, and the cancellous bone of tetrapod limbs. Additionally, within the elasmobranch fishes, mineralized trabeculae (struts) have been reported in the jaws of durophagous myliobatid stingrays (Elasmobranchii: Batoidea), but were believed to be absent in basal members of the batoid clade. This study, however, presents an additional case of batoid trabeculation in the lesser electric ray, Narcine brasiliensis (Torpediniformes). The trabeculae in these species likely play different functional roles. Stingrays use their reinforced jaws to crush bivalves, yet N. brasiliensis feeds by ballistically protruding its jaws into the sediment to capture polychaetes. In N. brasiliensis, trabeculae are localized to areas likely to experience the highest load: the quadratomandibular jaw joints, hyomandibular-cranial joint, and the thinnest sections of the jaws immediately lateral to the symphyses. However, the supports perform different functions dependent on location. In regions where the jaws are loaded transversely (as in durophagous rays), "load leading" trabeculae distribute compressive forces from the cortex through the lumen of the jaws. In the parasymphyseal regions of the jaws, "truss" trabeculae form cross-braces perpendicular to the long axes of the jaws. At peak protrusion, the jaw arch is medially compressed and the jaw loaded axially such that these trabeculae are positioned to resist buckling associated with excavation forces. "Truss" trabeculae function to maintain the second moment of area in the thinnest regions of the jaws, illustrating a novel function for batoid trabeculation. Thus, this method of structural support appears to have arisen twice independently in batoids and performs strikingly different ecological functions associated with the distribution of extreme loading environments.

Animals↗

Interspecies exchange of a Hoxd enhancer in vivo induces premature transcription and anterior shift of the sacrum.

The precise activation, in space and time, of vertebrate Hox genes is an essential requirement for normal morphogenesis. In order to assess for the functional potential of evolutionary conserved Hox regulatory sequences, a phylogenetically conserved bipartite regulatory element necessary for proper spatial and temporal activation of the Hoxd-11 gene was replaced by its fish counterpart in the HoxD complex of mice, using an ES cell-based targeted exchange. Fetuses carrying this replacement activated Hoxd-11 transcription prematurely, which led to a rostral shift of its expression boundary and a consequent anterior transposition of the sacrum. These results demonstrate the high phylogenetic conservation of regulatory mechanisms acting over vertebrate Hox complexes and suggest that minor time difference (heterochronies) in Hox gene activation may have contributed to important morphological variations in the course of evolution.

Animals↗

Molecules, morphology, fossils, and the relationship of angiosperms and Gnetales.

Morphological analyses of seed plant phylogeny agree that Gnetales are the closest living relatives of angiosperms, but some studies indicate that both groups are monophyletic, while others indicate that angiosperms are nested within Gnetales. Molecular analyses of several genes agree that both groups are monophyletic, but differ on whether they are related. Conflicts among morphological trees depend on the interpretation of certain characters; when these are analyzed critically, both groups are found to be monophyletic. Conflicts among molecular trees may reflect the rapid Paleozoic radiation of seed plant lines, aggravated by the long branches leading to extant taxa. Trees in which angiosperms are not related to Gnetales conflict more with the stratigraphic record. Even if molecular data resolve the relationships among living seed plant groups, understanding of the origin of angiosperm organs will require integration of fossil taxa, necessarily using morphology.

Evolution, Molecular↗