Early steps in building the insect brain: neuroblast formation and segmental patterning in the developing brain of different insect species

In insects, morphological, molecular and genetic studies have provided a detailed insight into the ontogenetic processes that shape the ventral nerve cord. On the other hand, owing to its complexity and less obvious segmental composition, the knowledge about the development of the brain is still fragmentary. A promising approach towards gaining insight into fundamental processes underlying brain development is the comparison of embryonic brain development among different insect species. However, so far such comparative analyses are scarce. In this review, we summarize and compare data on the early steps in brain formation in different hemi- and holometabolous insects. We show that basic aspects of the spatial and temporal development of the embryonic brain neuroblast pattern are conserved among insects. Furthermore, we compare the number and proliferation patterns of neuroblasts related to major neuropil structures such as mushroom bodies, central complex, and antennal lobe. Finally, comparing the expression patterns of engrailed in different species, and considering new data from Drosophila melanogaster, we discuss the segmental organization of the insect brain.     

Extension and retraction of axonal projections by some developing neurons in the leech depends upon the existence of neighboring homologues. I. The HA cells

The role of homologues in the establishment of the pattern of axonal projections of identified segmentally homologous neurons was investigated by means of selective cell ablation and dye injection. The cells studied were the bilateral pairs of heart accessory (HA) neurons found in the fifth and sixth segmental ganglia of the leech ventral nerve cord. Homologues start their morphological differentiation with identical axonal projections, and segmental differences are manifested later, when specific branches stop growing and disappear. The deletion of single HA cells at early stages, however, permits these branches to survive in their ipsilateral homologues and to grow and take over the projections of the deleted neurons. In addition, if both HA homologues on the same side of the nerve cord, or three of the four HA cells, are deleted in an animal, the remaining HA cells often extend novel projections. These observations suggest that either competition for targets, inputs or growth factors, or direct interactions among homologous cells may play a role in the differentiation of segment specific patterns of axonal projections.     

Morphological variation in Gyrinus sericeolimbatus Rég. in New Guinea and description of a new Gyrinus species (Coleoptera: Gyrinidae)

Abstract. Samples of Gyrinus sericeolimbatus from twenty-six localities in New Guinea vary greatly in metric body characters as well as in elytral reticulation patterns. The variation is smaller within than between populations. Although the characters of adjacent populations are usually similar, certain proximate populations exhibit significant differences, probably resulting from some kind of dispersal barrier. Similarities between populations at comparable latitudes suggest that some of the morphological variation is altitude-related. In addition, evidence is found for habitat-related morphological differentiation. Changes in phenotypes in response to possible environmental factors in ontogeny and constraints imposed by various selection pressures are discussed.
A new species, G.brincki sp.n., from western New Guinea is described.     

Song complexity is associated with habitat quality in an upland passerine

Understanding the influence of intrinsic (genetic and morphological) and extrinsic (geographical, environmental and social) factors on the performance and spatial differentiation of sexual signals, such as bird song, can help identify behavioural indicators of individual quality, habitat degradation and social environment. We used the Iberian Bluethroat Luscinia svecica azuricollis, a migratory bird that breeds in fragmented landscapes dominated by shrublands, as a case study to: (1) assess how a set of acoustic indicators of song performance are driven by intrinsic and extrinsic factors; and (2) contrast deterministic (adaptations to the environmental context and morphological constraints) vs. stochastic (differentiation by geographical isolation) explanations for song differentiation patterns. We explored acoustic indicators of song performance (spectral, temporal and song complexity) in relation to parameters related to genetic structure, body size, habitat type, habitat quality (assessed through a spatially explicit modelling approach) and social context (population abundance and songbird community composition). Then, we explored the contribution of genetic, geographical and environmental dissimilarity to song diversification across space. Our results highlight an association of song spectral variables with genetic structure and a significant connection between song complexity and duration with habitat quality. We found no relationship between social features and acoustic variables, or between song differentiation and genetic or geographical distances. There was, however, a correlation between song differentiation and environmental dissimilarity. We recommend the consideration of song complexity as an indicator of habitat quality.     

Evolution of the morphological innovations of feathers

Feathers are complex assemblages of multiple morphological innovations. Recent research on the development and evolution of feathers has produced new insights into the origin and diversification of the morphological innovations in feathers. In this article, I review and discuss the contribution of three different factors to the evolution of morphological innovations in feathers: feather tubularity, hierarchical morphological modularity, and the co-option molecular signaling modules. The developing feather germ is a tube of epidermis with a central dermal pulp. The tubular organization of the feather germ and follicle produces multiple axes over which morphological differentiation can be organized. Feather complexity is organized into a hierarchy of morphological modules. These morphological modules evolved through the innovative differentiation along multiple different morphological axes created by the tubular feather germ. Concurrently, many of the morphological innovations of feathers evolved through the evolutionary co-option of plesiomorphic molecular signaling modules. Gene co-option also reveals a role for contingency in the evolution of hierarchical morphological innovations.     

A preliminary classification of evolutionary radiations

Rapid increases in taxonomic diversity are generally described as adaptive or evolutionary radiations. Such radiations differ widely in the rate and extent of morphologic innovation, taxonomic diversification and phylogenetic breadth, suggesting that several patterns, and likely processes, are involved. At least four distinct patterns of evolutionary radiation can be identified: novelty events, which generate new morphological complexity (altering the body plan of the group under consideration) but not necessarily with the associated production of many lower taxa; broad diversification events involving many independent lineages that undergo diversification, generate many new species and are driven by new ecological opportunities; economic radiations of a limited group of ecologically (but not necessarily phylogenetically) related clades exploiting a limited new ecologic opportunity; and adaptive radiations that may occur at any taxonomic level, but involve a rapid increase in diversity within a single clade, including “true”; adaptive radiations. Many events produce simple diversity increases with no corresponding increase in genetic/developmental/morphological/behavioral sophistication, but the most evolutionarily interesting events add new levels of complexity.     

Cranial evolution in sakis (Pithecia, Platyrrhini). II: Evolutionary processes and morphological integration

Patterns of interspecific differentiation in saki monkeys (Pithecia) are quantitatively described and possible evolutionary processes producing them are examined. The comparison of species correlation matrices to expected patterns of morphological integration reveal significant and similar patterns of development-based cranial integration among species. Aspects of the facial region are more heavily influenced by general size variation than features of the neural region. The comparison of pooled within- and between-groups V/CV matrices suggests that genetic drift might be a sufficient explanation for saki cranial evolution. Differential natural selection gradients are also reconstructed because selection may also have caused population differentiation through evolutionary time. These gradients illustrate the inherent multivariate nature of selection, being a consequence of the interaction between existing morphological integration (correlation) among traits and the action of natural selection. Yet, our attempt to interpret selection gradients in terms of their functional significance did not result in any clear association between selection and function. Perhaps this is also an indication that morphological evolution in sakis was mostly neutral.     

A study of Aphyosemion schmitti (Romand, 1979) and a survey of the Aphyosemion of Liberia (Pisces, Cyprinodontidae)

Aphyosemion schmitti can be separated from neighbouring species on the basis of colour pattern, crossing experiments, cytogenetics, genetics and zoogeographical distribution. Three groups of Aphyosemion can be distinguished, based on morphological and meristic characters, colour patterns and geographical distribution; i.e. the liberiense group, the guineense group and the occidentalis group. A differentiation of the Liberian Aphyosemion is proposed, and its possible phylogenetic relationships are discussed.     

A metric for the study of evolutionary trends in the complexity of serial structures

Little empirical work has been done to see what sort of patterns of change in morphological complexity occur in evolution, mainly because the complexity of whole organisms has been so hard to define and to measure. For serial structures within organisms, there are fewer difficulties; this paper introduces a set of complexity metrics that are designed especially for serial structures, and then explores some of the properties of the new metrics. Also, a principle proposed in the last century by Herbert Spencer, and offered recently in a new form by the thermodynamic school of evolutionary thought, predicts that complexity should increase in evolution as a consequence of the accumulation of perturbations. Here, simulations in which perturbations are introduced to ideal and real series of vertebral measurements show how the complexity increase predicted by Spencer's principle would be captured by the new metrics.     

Searching for Sex Differences in the Vomeronasal Pathway

The sexual differentiation of brain and behavior is reviewed from the findings of sex differences in the vomeronasal pathway. A motivational approach to sex differences in reproductive behavior is stressed by taking into account that sex differences are present in neural networks: from the receptor organ (the vomeronasal organ) to effector nuclei. Sex differences in the brain appear in two morphological patterns. In one, the male presents greater morphological measurements than the female; in the other, the opposite occurs. These two morphological patterns are actively differentiated by gonadal steroids. The functional significance of these two morphological patterns is addressed. Moreover, since the GABA_Areceptor is involved in the organization of sex differences in vomeronasal structures such as the accessory olfactory bulb and in maternal behavior, the role of membrane mechanisms, 5α reduced hormones, and neurosteroids in the sexual differentiation process is discussed.     

Somatic musculature of Tardigrada: phylogenetic signal and metameric patterns

Although studies describing molecular‐based phylogenies within tardigrades are now frequently being published, this is not the case for studies combining molecular and morphological characters. Tardigrade phylogeny is still based, from a morphological point of view, almost exclusively on chitinous structures and little attention has been given to detecting and using novel morphological data. Consequently, we analysed the musculature of seven tardigrade species belonging to the main phyletic lines by confocal laser scanning microscopy and compared these morphological results with new molecular analyses (18S+28S rRNA genes). Finally, we analysed all the data with a total evidence approach. A consilience in the phylogenetic relationships among orders and superfamilies of tardigrades was obtained among the evolutionary trees obtained from morphological, molecular and total evidence approaches. Comparative analysis on the musculature allowed the identification of serial homologies and repeated metameric patterns along the longitudinal animal body axis. A phenomenon of mosaic evolution was detected in musculature anatomy, as dorsal musculature was found to be highly modified with respect to the other body muscle groups, probably related to the evolution of dorsal cuticular plates. An understanding of tardigrade musculature anatomy will give fundamental information to understand the evolution of segmental pattern within Panarthropoda. © 2013 The Linnean Society of London     

A three-phase model of arthropod segmentation

Molecular and morphological evidence (expression patterns of pair-rule genes and segmental position of the genital openings and other segmental markers) suggest that the segmental units of the arthropod body are specified, in early ontogeny, by three spatially and/or temporally distinct mechanisms and do not appear in a strict antero-posterior sequence. A first anterior set of indivisible segments (naupliar segments, possibly three in all arthropods) is followed by a set of more caudal (post-naupliar) primary units (eosegments, possibly ten in all arthropods) which then undergo a process of secondary segmentation, thus giving rise to a higher number of definitive segments (merosegments). The number of merosegments deriving from each eosegment is characteristic of the different arthropod clades and is mostly stable at the level of the traditional arthropodan classes or subclasses. All their segmentation patterns, however, including those found in the segmental organisation of highly segmented forms (such as centipedes and millipedes, notostracan, lipostracan and anostracan crustaceans, and trilobites) are reducible to the basic groundplan with three naupliar and ten postnaupliar segments. These basic units of arthropod segmentation may also have an equivalent in other Ecdysozoa, despite the lack of any segmentation (nematodes) or, at least, of an overt segmentation (kinorhynchs).     

Geographic patterns of morphological variation in Turnera sidoides subsp. pinnatifida (Turneraceae)

With the objective of contributing to understanding the patterns of variation within the Turnera sidoides complex, a detailed evaluation of morphological variation along the range of T. sidoides subsp. pinnatifida was performed. A multivariate analysis based on leaf traits and flower colour data enabled differentiation of five morphotypes. Common-garden experiments demonstrated that the morphological variants have a strong genetic basis. It was also found that the morphotypes are geographically structured along the subspecies range, display different habitat preferences, and occur in regions with different climatic regimes. Although these results are suggestive of adaptive differentiation of T. sidoides subsp. pinnatifida, comparisons between morphological and bioclimatic ordinations showed that the patterns observed cannot be fully explained by current climatic conditions. It is proposed that Miocene–Pleistocene events may explain the origin of the five morphotypes and that current climatic and ecological factors may be contributing to the maintenance of the extent and patterns of morphological differentiation in T. sidoides subsp. pinnatifida.     

Taxonomic differentiation of Cupressus sempervirens and C. atlantica based on morphometric evidence

We investigated the interspecific morphological relationships of two related Mediterranean Cupressus taxa and the possible geographic differentiation of Eastern Mediterranean C. sempervirens using morphometric analysis. A total of 446 individuals, representing eight C. atlantica (Morocco) and 10 C. sempervirens (Greece, Turkey and Lebanon) populations were examined. The 17 morphological characters of cone, seed and shoot with leaves were measured and analysed statistically. Significant morphological differences between C. atlantica and C. sempervirens were detected for the majority of analysed characters. For C. sempervirens, significant differences were also detected and these were related to its three regions of origin, the Greek Islands, Taurus Mts and Lebanon Mts. Multivariate analyses did not indicate a clear geographic pattern of differentiation in C. sempervirens populations. A relatively low level of phenotypic differentiation was detected between populations originating from the Greek Islands and Taurus Mts. We also observed a moderate multivariate differentiation between the Lebanese and remaining C. sempervirens populations. Phenotypic differentiation detected in the Asian part of the C. sempervirens range reflects patterns of differentiation similar to those described for other taxa occurring in the Taurus and Lebanon Mts.     

Ancient origin of the new developmental superfamily DANGER

Developmental proteins play a pivotal role in the origin of animal complexity and diversity. We report here the identification of a highly divergent developmental protein superfamily (DANGER), which originated before the emergence of animals (approximately 850 million years ago) and experienced major expansion-contraction events during metazoan evolution. Sequence analysis demonstrates that DANGER proteins diverged via multiple mechanisms, including amino acid substitution, intron gain and/or loss, and recombination. Divergence for DANGER proteins is substantially greater than for the prototypic member of the superfamily (Mab-21 family) and other developmental protein families (e.g., WNT proteins). DANGER proteins are widely expressed and display species-dependent tissue expression patterns, with many members having roles in development. DANGER1A, which regulates the inositol trisphosphate receptor, promotes the differentiation and outgrowth of neuronal processes. Regulation of development may be a universal function of DANGER family members. This family provides a model system to investigate how rapid protein divergence contributes to morphological complexity.     

Evolutionary conservation of microRNA regulatory circuits: an examination of microRNA gene complexity and conserved microRNA-target interactions through metazoan phylogeny

During the last decade, a variety of critical biological processes, including early embryo development, cell proliferation, differentiation, apoptosis, and metabolic regularity, have been shown to be genetically regulated by a large gene family encoding a class of tiny RNA molecules termed microRNAs (miRNAs). All miRNAs share a common biosynthetic pathway and reaction mechanisms. The sequence of many miRNAs is found to be conserved, in their mature form, among different organisms. In addition, the evolutionary appearance of multicellular organisms appears to correlate with the appearance of the miRNA pathway for regulating gene expression. The miRNA pathway has the potential to regulate vast networks of gene products in a coordinate manner. Recent evidence has not only implicated the miRNA pathway in regulating a vast array of basic cellular processes but also specialized processes that are required for cellular identity and tissue specificity. A survey of the literature shows that some miRNA pathways are conserved virtually intact throughout phylogeny while miRNA diversity also correlates with speciation. The number of miRNA genes, the expression of miRNAs, and target diversities of miRNAs tend to be positively correlated with morphological complexities observed in animals. Thus, organismal complexity can be estimated by the complexity of the miRNA circuitry. The complexity of the miRNA gene families establishes a link between genotypic complexity and phenotypic complexity in animal evolution. In this paper, we start with the discussion of miRNA conservation. Then we interpret the trends in miRNA conservation to deduce miRNA evolutionary trends in metazoans. Based on these conservation patterns observed in each component of the miRNA regulatory system, we attempt to propose a global insight on the probable consistency between morphological evolution in animals and the molecular evolution of miRNA gene activity in the cell.     

An evaluation of myelomeres and segmentation of the chick embryo spinal cord.

We have investigated whether the neuromeres of the developing chick spinal cord (myelomeres) are manifestations of intrinsic segmentation of the CNS by studying the patterns of cell proliferation and neuronal differentiation. Treatment of 2-day embryos with colchicine does produce exaggerated myelomeres, in confirmation of K?llén (Z. Anat. Entwickl.-Gesch. 123, 309-319, 1962). However, this does not imply that myelomeres are segmental proliferation centres: the undulations caused by colchicine are irregular alongside the unsegmented mesoderm, and another mitotic inhibitor, bromodeoxyuridine, has no such effects. In contrast to lower vertebrate embryos, there is no evidence for segmental groups of primary motor neurons in the chick: the earliest motor neurons express cholinesterase, and project their axons into the adjacent sclerotome, at random positions in relation to the somite boundaries. The population of motor neurons projecting HRP-labelled axons into a single somite lies out of phase with both myelomere and somite, and is placed symmetrically about the anterior half-sclerotome. The earliest intrinsic spinal cord neurons, as stained with zinc iodide-osmium tetroxide or anti-68 x Mr neurofilament antibody, show no segmental patterns of differentiation. We conclude that, in contrast to the rhombomeres of the developing hindbrain, myelomeres are not matched by segmental groupings of differentiating nerve cells, and result from mechanical moulding of the neuroepithelium by the neighbouring somites.     

Fractal analysis of sagittal suture morphology

Quantifying shape is a broad problem in the morphological sciences. Most techniques for numerically describing shape abstract the shape into the most logical ideal Euclidean dimension. The fractional, or fractal, dimension is a simple computation that expresses shape in real, rather than ideal, space. The structured walk technique developed for the fractal analysis of rugged boundaries is applied here to the contour of the human sagittal suture in order to discriminate the separate morphological patterns of interfingering and interlocking. These attributes contribute differentially to the suture's “complexity,” a concept often used in biomechanical hypotheses. Previous techniques for estimating sutural complexity do not isolate small-scale from large-scale morphological patterns. Results indicate that despite the visual appearance of great variation, human sagittal sutures are remarkably consistent in the degree of complexity expressed separately by large-scale interfingering lateral excursions and small-scale interlocking ruggedness. There is no significant correlation between the absolute or bregma-lambda chord length of the human sagittal suture and its degree of complexity as determined by the structured walk technique.     

Genomic divergence across ecological gradients in the Central African rainforest songbird (Andropadus virens)

The little greenbul, a common rainforest passerine from sub‐Saharan Africa, has been the subject of long‐term evolutionary studies to understand the mechanisms leading to rainforest speciation. Previous research found morphological and behavioural divergence across rainforest–savannah transition zones (ecotones), and a pattern of divergence with gene flow suggesting divergent natural selection has contributed to adaptive divergence and ecotones could be important areas for rainforests speciation. Recent advances in genomics and environmental modelling make it possible to examine patterns of genetic divergence in a more comprehensive fashion. To assess the extent to which natural selection may drive patterns of differentiation, here we investigate patterns of genomic differentiation among populations across environmental gradients and regions. We find compelling evidence that individuals form discrete genetic clusters corresponding to distinctive environmental characteristics and habitat types. Pairwise F_ST between populations in different habitats is significantly higher than within habitats, and this differentiation is greater than what is expected from geographic distance alone. Moreover, we identified 140 SNPs that showed extreme differentiation among populations through a genomewide selection scan. These outliers were significantly enriched in exonic and coding regions, suggesting their functional importance. Environmental association analysis of SNP variation indicates that several environmental variables, including temperature and elevation, play important roles in driving the pattern of genomic diversification. Results lend important new genomic evidence for environmental gradients being important in population differentiation.