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Reciprocal functions of the Drosophila yellow and ebony proteins in the development and evolution of pigment patterns.

Body coloration affects how animals interact with the environment. In insects, the rapid evolution of black and brown melanin patterns suggests that these are adaptive traits. The developmental and molecular mechanisms that generate these pigment patterns are largely unknown. We demonstrate that the regulation and function of the yellow and ebony genes in Drosophila melanogaster play crucial roles in this process. The Yellow protein is required to produce black melanin, and is expressed in a pattern that correlates with the distribution of this pigment. Conversely, Ebony is required to suppress some melanin formation, and is expressed in cells that will produce both melanized and non-melanized cuticle. Ectopic expression of Ebony inhibits melanin formation, but increasing Yellow expression can overcome this effect. In addition, ectopic expression of Yellow is sufficient to induce melanin formation, but only in the absence of Ebony. These results suggest that the patterns and levels of Yellow and Ebony expression together determine the pattern and intensity of melanization. Based on their functions in Drosophila melanogaster, we propose that changes in the expression of Yellow and/or Ebony may have evolved with melanin patterns. Consistent with our hypothesis, we find that Yellow and Ebony are expressed in complementary spatial patterns that correlate with the formation of an evolutionary novel, male-specific pigment pattern in Drosophila biarmipes wings. These findings provide a developmental and genetic framework for understanding the evolution of melanin patterns.

Animals↗

Pattern formation in janus-mutant zebrafish embryos.

Mechanisms that underlie the formation of the vertebrate body appear to be highly conserved between amphibia and teleosts. For teleosts, however, mesoderm induction and the establishment of dorsoventral polarity are poorly understood. In this study, we present an analysis of early pattern formation in the zebrafish maternal-effect mutation janus. This mutation frequently results in a separation of the cleavage stage blastoderm into two halves that undergo separate development until fusion occurs at the end of gastrulation. Here, we employ janus-mutant embryos to analyze the mechanisms of mesoderm formation and ventral specification in a teleost. Analysis of the expression of the panmesodermal marker no tail in janus-mutant embryos indicates that mesoderm induction depends on a marginal position. In an analysis of ventral specification, we show that the early expression of the ventral marker GATA-2 is confined to the area on both blastodermal halves opposite the dorsal shield region. Since, in janus-mutant embryos, the dorsal position is random with respect to the division plane bisecting the two blastodermal halves, a variety of dorsoventral asymmetries arise within individual embryos. In one constellation, the dorsal position is localized to the plane of bisection and two ventral positions develop at opposite ends of the blastodermal halves. Hence, ventral fates can be specified at any position around the blastodermal margins and are excluded from the dorsal position. The diblastodermic system of the janus-mutant embryo allows for the study of the interactions of dorsal and ventral determinants in varying spatial arrangements. We have studied pattern formation in dorsal half-blastoderms that contain the entire shield region but only a reduced ventrolateral marginal zone. As assessed by the presence of the most ventral cell type, blood, ventral specification within a dorsal half-blastoderm is not suppressed.

Animals↗

Physical model of anode glow patterns in elevated-pressure gas discharges.

A physical self-consistent model is developed to explain single spots or complex current structures at the anode of elevated-pressure parallel-plate dc discharges. The model is based on a fluid description of electron and ion transport coupled with Poisson's equation and involves a pair of coupled reaction-diffusion equations of an activator-inhibitor-type. This system of one-dimensional equations containing no phenomenological (adjustable) parameters allows one to find the current-density (activator) and anode potential drop (inhibitor) distributions on the anode surface. In a certain range of supply voltage, an anode glow stratification, resulting in the formation of separate glowing regions, takes place. However, the growth of perturbations and formation of a spatially periodic current pattern are complicated by competition between the current stripes, leading to suppressing of the neighboring current stripes. The bifurcation behavior of the model with respect to the characteristic electron energy, recombination coefficient, and discharge gap has been analyzed. The properties of a single anode current structure, including the normal current density effect, have been investigated. The application of these results to available findings in experiments and two-dimensional numerical simulations is discussed.

Journal Article↗

Ripening of surface phases coupled with oscillatory dynamics and self-induced spatial chaos through surface roughening.

Some pattern formation processes on single-crystal catalytic surfaces involve transitions between alternative surface phases coupled with oscillatory reaction dynamics. We describe a two-tier symmetry-breaking model of this process, based on nanoscale boundary dynamics interacting with oscillations of adsorbate coverage on microscale. The surface phase distribution oscillates together with adsorbate coverage, and, in addition, undergoes a slow coarsening process due to the curvature dependence of the drift velocity of interphase boundaries. The coarsening is studied both statistically, assuming a circular shape of islands of the minority phase, and through detailed Lagrangian modeling of boundary dynamics. Direct simulation of boundary dynamics allows us to take into account processes of surface reconstruction, leading to self-induced surface roughening. As a result, the surface becomes inhomogeneous, and the coarsening process is arrested way before the thermodynamic limit is reached, leaving a chaotic distribution of surface phases. (c) 1999 American Institute of Physics.

Journal Article↗

Symmetry breaking and coarsening in spatially distributed evolutionary processes including sexual reproduction and disruptive selection.

Sexual reproduction presents significant challenges to formal treatment of evolutionary processes. A starting point for systematic treatments of ecological and evolutionary phenomena has been provided by the gene-centered view of evolution which assigns effective fitness to each allele instead of each organism. The gene-centered view can be formalized as a dynamic mean-field approximation applied to genes in reproduction and selection dynamics. We show that the gene-centered view breaks down for symmetry breaking and pattern formation within a population and show that spatial distributions of organisms with local mating neighborhoods in the presence of disruptive selection give rise to such symmetry breaking and pattern formation in the genetic composition of local populations. Global dynamics follows conventional coarsening of systems with nonconserved order parameters. The results have significant implications for the ecology of genetic diversity and species formation.

Alleles↗

Spatio-temporal VEGF and PDGF delivery patterns blood vessel formation and maturation.

PURPOSE: Biological mechanisms of tissue regeneration are often complex, involving the tightly coordinated spatial and temporal presentation of multiple factors. We investigated whether spatially compartmentalized and sequential delivery of factors can be used to pattern new blood vessel formation. MATERIALS AND METHODS: A porous bi-layered poly(lactide-co-glycolide) (PLG) scaffold system was used to locally present vascular endothelial growth factor (VEGF) alone in one spatial region, and sequentially deliver VEGF and platelet-derived growth factor (PDGF) in an adjacent region. Scaffolds were implanted in severely ischemic hindlimbs of SCID mice for 2 and 6 weeks, and new vessel formation was quantified within the scaffolds. RESULTS: In the compartment delivering a high dose of VEGF alone, a high density of small, immature blood vessels was observed at 2 weeks. Sequential delivery of VEGF and PDGF led to a slightly lower blood vessel density, but vessel size and maturity were significantly enhanced. Results were similar at 6 weeks, with continued remodeling of vessels in the VEGF and PDGF layer towards increased size and maturation. CONCLUSIONS: Spatially localizing and temporally controlling growth factor presentation for angiogenesis can create spatially organized tissues.

Animals↗

Temporal and spatial regulation of fibronectin in early Xenopus development.

Pattern formation and temporal control of gene expression in Xenopus development were investigated using fibronectin as a biochemical marker. We determined the spatial localization of fibronectin in the embryo by immunofluorescence and the temporal program of its expression by biosynthesis studies and Western blotting techniques. At the start of gastrulation, fibronectin is localized on the roof of the blastocoel which serves as the surface upon which mesodermal cells will migrate. However, since we find fibronectin secreted by all parts of the embryo, localization is probably achieved through spatially localized receptors that bind secreted fibronectin. Fibronectin levels and fibronectin synthesis rates increase following the midblastula stage. This increase is independent of transcription and therefore involves activation of maternal RNA for fibronectin. Since this message mobilization also occurs in activated but unfertilized eggs, this event must be regulated separately from the midblastula transition.

Animals↗

Dancing waves in reaction-diffusion systems.

The mechanism of pattern formation in reaction-diffusion systems is treated as an interesting subject, generally for understanding self-organization observed in living systems and natural phenomena. Several spatial patterns appear in the reaction-diffusion systems where an activator and an inhibitor coexist as an intermediate, as represented by a traveling wave, a stationary wave called a Turing structure, etc. Here, we show new kinds of waves in reaction-diffusion systems, which exhibit reciprocating motion without colliding into each other or blinking periodically. These patterns have never been observed in the conventional numerical models, although experimentally oscillating spots have been often observed. Our model demonstrates that other than the ratio of diffusion coefficients for both intermediates, the thickness of reaction media acts to generate inhibitory effect. The spatial factor of the medium contributes to new pattern formation in reaction-diffusion systems. For the design of new functional materials, the concept might be useful as a simple controlling method for pattern dynamics.

Journal Article↗

The role of a reaction-diffusion system in the initiation of primary hair follicles.

A mechanism based on a reaction-diffusion system is proposed for the initiation of hair follicles in the epidermis during fetal development. It is demonstrated that initiation of primary follicles in a series of waves, within the proposed mechanism, is a consequence of the size and shape dependent properties of the reaction-diffusion system without the need for the propagation of signals through the skin. The observed trio grouping of follicles and variation of primary follicle density per unit skin area during development are also correctly predicted. An explanation, based on the reaction-diffusion system and the variation of its characteristic spatial wavelength with time during development, is suggested for the termination of both primary and secondary follicle initiation as well as follicle neogenesis. The proposed initiation mechanism is basically the same as that used to explain various spatial patterns observed in hair fibre formation (Nagorcka & Mooney, 1982).

Animals↗

Cloning and characterization of zebrafish smad2, smad3 and smad4.

smad genes encode transcription factors involved in the signal transduction of members of the TGFbeta superfamily. We report here the cloning, characterization and genomic mapping of smad2, smad3 and smad4 from the zebrafish, Danio rerio. In Xenopus, smad2 overexpression has been shown to interfere with gastrulation and dorsal cell fate specification. However, full-length zebrafish smad2, although functionally active in Xenopus explants, has no effect when overexpressed in zebrafish embryos. In contrast, an N-terminally truncated, constitutively active version of Smad2 protein causes severe dorsalization or partial secondary axis formation, pointing to a role of Smad2 during mesoderm and axis formation. The temporal and spatial expression patterns of zebrafish smad2, 3 and 4 were investigated by developmental RT-PCR and whole mount in-situ hybridization. All three genes show strong and ubiquitous maternal expression. Zygotic expression is weak and ubiquitous in the case of smad2, and strong and ubiquitious in the case of smad4, while smad3 shows a spatially restricted zygotic expression pattern. It is expressed in migrating neural crest cells of the trunk and a subset of cells in the diencephalon in close proximity to the expression domain of the Nodal-related protein Cyclops/Ndr2/Znr1, a potential signal upstream of Smad2/3 required for eye-field separation and floor plate specification. Overexpression of truncated smad2 in cyclops mutant embryos leads to a rescue of the eye and floorplate defects. These data suggest that Smad2 acts as a mediator of Nodal signals during zebrafish midline signaling, while Smad3 might be involved in later steps of eye field separation.

Amino Acid Sequence↗

Using mathematical models to help understand biological pattern formation.

One of the characteristics of biological systems is their ability to produce and sustain spatial and spatio-temporal pattern. Elucidating the underlying mechanisms responsible for this phenomenon has been the goal of much experimental and theoretical research. This paper illustrates this area of research by presenting some of the mathematical models that have been proposed to account for pattern formation in biology and considering their implications.

Animals↗

Extracellular matrix during laminar pattern formation of neocortex in normal and reeler mutant mice.

The spatial and temporal distribution of extracellular matrix, which occupied the large extracellular spaces in the developing cerebral cortex, was studied during pre- and perinatal ontogenesis of normal and reeler mutant mice. Colloidal iron-staining material was localized principally in the marginal zone and subplate of normal mice, whereas in reeler mutants, most of the material was found in the outer layers of the cortex. Patterns of extracellular matrix localization in both genotypes followed the laminar pattern formation of cerebral cortex architecture. Histochemical ultrastructural visualization of this extracellular matrix and its susceptibility to enzymatic treatment suggested that the major components are glycosaminoglycans. Their possible role in relation to afferent axon targeting is discussed.

Animals↗

Some physical, mathematical and evolutionary aspects of biological pattern formation.

An important mechanism in the generation of biological structures is the production of defined spatial patterns within initially near-uniform cells and tissues. This process can be modelled on the basis of conventional molecular kinetics if there is a short-range activating effect in conjunction with depletion or inhibition extending over a wider range ('lateral inhibition'). Such pattern-generating systems exhibit simple self-regulatory properties empirically observed in developmental biology such as polarity effects, proportion regulation and the inducibility of secondary centres. Autocatalysis and lateral inhibition have been shown to be mathematically necessary for the simplest two-factor case. Certain generalizations of these conditions to multi-component systems are possible; they are suitable for modelling intercalary regeneration. The evolution of higher organisms seems to be determined to a considerable extent by many small changes of patterns and proportions. While evolution proceeds at varying rates in the course of time, the rate-limiting steps may be due to mutations of low selection pressure. A semi-quantitative argument suggests that there might be an upper limit of evolutionarily effective genetic complexity.

Animals↗

Temporal and spatial patterning of an organ by a single transcription factor.

During the formation of animal organs, a single regulatory factor can control the majority of cell-fate decisions, but the mechanisms by which this occurs are poorly understood. One such regulator, the nematode transcription factor PHA-4, functions together with various cis-regulatory elements in target genes to regulate spatial and temporal patterning during development of the pharynx.

Animals↗

Genes, lineages and the neural crest: a speculative review.

Sensory and sympathetic neurons are generated from the trunk neural crest. The prevailing view has been that these two classes of neurons are derived from a common neural crest-derived progenitor that chooses between neuronal fates only after migrating to sites of peripheral ganglion formation. Here I reconsider this view in the light of new molecular and genetic data on the differentiation of sensory and autonomic neurons. These data raise several paradoxes when taken in the context of classical studies of the timing and spatial patterning of sensory and autonomic ganglion formation. These paradoxes can be most easily resolved by assuming that the restriction of neural crest cells to either sensory or autonomic lineages occurs at a very early stage, either before and/or shortly after they exit the neural tube.

Animals↗

Self-organization of chaos synchronization and pattern formation in coupled chaotic oscillators.

Pattern formation in spatiotemporal chaotic systems is investigated. Temporally chaotic and spatially ordered patterns are observed by varying the coupling strength. Spatial orderings emerge spontaneously due to self-organization of partial and nonlocal chaos synchronization, governed by various types of spatial symmetries. The first and secondary bifurcations from spatially disordered chaos to chaos with different levels of spatial orderings are observed and the scaling behaviors associated with these bifurcations are statistically analyzed.

Journal Article↗

Control of formation of the two types of polyps in Thecocodium quadratum (Hydrozoa, Cnidaria).

Thecocodium quadratum (Werner, Jber. Biol. Anst. Helgoland, 1965) is a colonial hydroid which produces 2 different types of polyps: gastrogonozooids and dactylozooids. The mouthless dactylozooids bear tentacles and catch the prey, which is then taken over and swallowed by the gastrogonozooids which have no tentacles. It is obvious that for a colony to survive both polyps must exist simultaneously arranged in a certain spatial pattern. Our experiments indicate that the formation of polyps in a growing culture is governed by at least 3 principles: (1) short range inhibition between polyps irrespective of their differentiation; (2) long range specific inhibition between gastrogonozooids; and (3) long range supporting influence (lateral help, Meinhardt, H., Models of Biological Pattern Formation, 1982) between gastrogonozooids and dactylozooids.

Animals↗

Spatial pattern of smooth muscle differentiation is specified by the epithelium in the stomach of mouse embryo.

Smooth muscle induction in the stomach of mouse embryo was analyzed experimentally by tissue recombination and organ culture technique. When observed immunohistochemically with anti-alpha-smooth muscle actin antibody, smooth muscle myoblasts first differentiated and formed the circular muscle layer at 13 days of gestation. The longitudinal muscle layer appeared at 15 days in the forestomach and at neonatal period in the glandular stomach. The lamina muscularis mucosae was observed by the neonatal period. When the undifferentiated 11-day stomach was cultivated in vitro for 3 days, an alpha-smooth muscle actin-positive cell layer was formed and the explant exhibited rhythmical contraction. The mesenchyme alone remained as an undifferentiated cell mass. When the mesenchyme was recombined with the epithelium, a distinct smooth muscle layer was formed. These demonstrated that the alpha-smooth muscle actin expression was induced by the epithelium. Next, we examined by heterologous recombination the roles of the epithelium in formation of the organ-specific spatial pattern of smooth muscle differentiation. The stomach mesenchyme recombined with the tracheal epithelium exhibited a partially tracheal type of alpha-smooth muscle actin expression. The uterine epithelium induced only a small number of smooth muscle myoblasts and the spatial organization was disordered. Finally, we assayed if the mode of smooth muscle induction by the stomach epithelium changes during development. Heterochronic recombination proved that the 14-day forestomach epithelium induced a muscularis mucosae-like smooth muscle layer that was contiguous to the epithelium, whereas the 11-day stomach and the 13-day forestomach epithelium induced a circular muscle-like smooth muscle layer that was distant from the epithelium. As a whole, our data suggest that the spatial pattern of smooth muscle differentiation is regulated by organ-specific and stage-specific influences from the epithelium.

Actins↗