Control of metamorphosis and pattern formation in Hydratinia (hydrozoa,cnidaria)

Hydractinia echinata is a marine colonial hydroid, a relative of the more widely known Hydra. In contrast to Hydra, embryogenesis, metamorphosis and colony growth in Hydractinia are experimentally accessible and therefore, provide an ideal model system for investigating the biochemical basis of pattern formation. In particular, the processes involved in the transformation of the drop-shaped freely swimming larva into a sessile tube-shaped polyp are easily monitored, because this transfomation can be induced by application of various substances. Our results indicate that the internal level of S-adenosylmethionine (SAM), potentially the most important methyl donor in transmethylation processes, plays a key role in the onset of metamorphosis. It is also proposed that the internal level of SAM plays a pivotal role in the proportioning and spacing of polyps within the colony.     

Maternal control of pattern formation in Xenopus laevis

We review the essential role of maternal factors in pattern formation for Xenopus laevis, focusing on VegT, Vg1, and Wnt11. Results from loss of function experiments demonstrate a clear requirement for these genes in germ layer specification, dorsal-ventral axis formation, and convergence extension. We also discuss these genes in the broader context of metazoan development, exploring whether and how their functions in the X. laevis model organism may or may not be conserved in other species. Wnt11 signaling in particular provides a classic example where understanding context in development is crucial to understanding function. Genomic sequencing, gene expression, and functional screening data that are becoming available in more species are providing invaluable aid to decoding and modeling signaling pathways. More work is needed to develop a comprehensive catalog of the Wnt signaling, T-box, and TGF-beta genes in metazoans both near and far in evolutionary distance. We finally discuss some specific experimental and modeling efforts that will be needed to understand the behavior of these signaling networks in vivo so that we can interpret these critical pathways in an evolutionary framework.     

Strategies for control of pattern formation in Caenorhabditis elegans

In this paper, strategies for controlling pattern formation in Caenorhabditis elegans are reviewed. The somatic tissues of this small nematode develop, in large part, by invariant cell lineages, whereas the germ-line tissue arises primarily by a variable pattern of divisions. The spatial organization of the germ-line tissue depends on special regulatory cells, the distal tip cells, which appear to influence nearby germ cells to remain in mitosis. In somatic tissues, the problem of specifying that a cell in a particular position assumes a particular fate seems to be controlled by a number of different strategies. These include the production of non-equivalent cells in particular positions of the lineage tree, local interactions between apparently equivalent cells in close contact, and the influence of another special regulatory cell, the anchor cell, over certain neighbouring cells.     

SIF, a novel morphogenetic inducer in hydrozoa.

By analogy to processes in angiogenesis (blood vessel formation), the development of the stolonal network in colonial hydrozoa involves stimulation of branching and mutual chemotropic attraction of the growing branches by means of soluble morphogenetic factors. We have identified a glycoconjugate of about 20 kDa, termed SIF (Stolon-Inducing Factor), which induces the formation of stolon branches when applied locally. Micropipettes ejecting SIF mimic the inducing action of stolon tips, the putative sources of SIF. When whole animals are exposed to SIF, stolons sprout not only from the base of the polyps but also from abnormal sites along the entire body, even from the head. In addition, the polyp (hydranth) secretes a chitinous periderm which, in the species under investigation, normally envelops stolons but not hydranths. At high SIF doses the whole hydranth is transformed into stolon tissue. The factor has been isolated from conditioned medium and from butanol extracts of Hydractinia echinata and Podocoryne carnea.     

Transmethylation of Sterols in Aerobically Adapting Saccharomyces cerevisiae

The transmethylation of methyl-(14)C-methionine and methyl-(14)C-adenosylmethionine into the nonsaponifiable lipids of anaerobically grown yeast during adaptation to aerobic conditions was investigated. The rate and extent of methyl transfer increased with aeration time and was dependent upon the presence of a fermentable carbon source and O(2). Methionine and adenosylmethionine uptake rates increased in adaptation buffer but did not seem to be the rate-limiting factor for transmethylation under the conditions studied. Thinlayer chromatography of the nonsaponifiable fraction after exposure to label showed the labeled product to be ergosterol. Samples taken after short-term exposure to label were composed of two labeled steroidal products, one with kinetics of an ergosterol precursor.     

A pattern formation mechanism to control spatial organization in the embryo of Drosophila melanogaster

It is known that cells are already committed to a particular segment at the cellular blastoderm stage during embryogenesis of Drosophila melanogaster. Recently, several segmentation genes have been observed to be expressed in a sequence of banded spatial patterns in the syncytial blastoderm, prior to the formation of the cellular blastoderm. It is demonstrated in this paper that a two component reaction-diffusion (RD) system with net production functions which are antisymmetric with respect to the uniform steady-state values, is capable of producing a sequence of seven spatial patterns in the syncytial blastoderm. The sequence of patterns obtained exhibit a strong preference for banded or striped patterns. The first pattern is a simple anteroposterior gradient while the second is a gradient in the dorsoventral direction. The next five patterns are a sequence of banded patterns which exhibit frequency doubling, i.e. the number of bands in each pattern tend to be double the number in the previous pattern. The predicted pattern sequence is comparable to that observed in the expression of some segmentation genes. It is suggested that a pattern formation mechanism based on such an RD system may exist in the embryo where it produces a sequence of prepatterns to regulate the expression of various segmentation genes leading ultimately to a segmented embryo. There is sufficient spatial information in the sequence of banded prepatterns for the segments to be unique.     

Temporal control of self‐organized pattern formation without morphogen gradients in bacteria

Diverse mechanisms have been proposed to explain biological pattern formation. Regardless of their specific molecular interactions, the majority of these mechanisms require morphogen gradients as the spatial cue, which are either predefined or generated as a part of the patterning process. However, using Escherichia coli programmed by a synthetic gene circuit, we demonstrate here the generation of robust, self‐organized ring patterns of gene expression in the absence of an apparent morphogen gradient. Instead of being a spatial cue, the morphogen serves as a timing cue to trigger the formation and maintenance of the ring patterns. The timing mechanism enables the system to sense the domain size of the environment and generate patterns that scale accordingly. Our work defines a novel mechanism of pattern formation that has implications for understanding natural developmental processes.     

hyp loci control cell pattern formation in the vegetative mycelium of Aspergillus nidulans.

Aspergillus nidulans grows by apical extension of multinucleate cells called hyphae that are subdivided by the insertion of crosswalls called septa. Apical cells vary in length and number of nuclei, whereas subapical cells are typically 40 microm long with three to four nuclei. Apical cells have active mitotic cycles, whereas subapical cells are arrested for growth and mitosis until branch formation reinitiates tip growth and nuclear divisions. This multicellular growth pattern requires coordination between localized growth, nuclear division, and septation. We searched a temperature-sensitive mutant collection for strains with conditional defects in growth patterning and identified six mutants (designated hyp for hypercellular). The identified hyp mutations are nonlethal, recessive defects in five unlinked genes (hypA-hypE). Phenotypic analyses showed that these hyp mutants have aberrant patterns of septation and show defects in polarity establishment and tip growth, but they have normal nuclear division cycles and can complete the asexual growth cycle at restrictive temperature. Temperature shift analysis revealed that hypD and hypE play general roles in hyphal morphogenesis, since inactivation of these genes resulted in a general widening of apical and subapical cells. Interestingly, loss of hypA or hypB function lead to a cessation of apical cell growth but activated isotropic growth and mitosis in subapical cells. The inferred functions of hypA and hypB suggest a mechanism for coordinating apical growth, subapical cell arrest, and mitosis in A. nidulans.     

Dynamics of pattern formation

Experiments were designed to test the validity ofTuring's suggested pattern-forming mechanisms, which are initially capable of giving rise to only five to seven uniform structures.TheOregon-R (wild-type), mass-cultured strain ofDrosophila melanogaster was employed. Selection for the regular arrangement of microchaetac on the margin of the fourth abdominal sternite was practiced for twenty generations. In the L line, individuals with six uniformly spaced bristles were selected as parents of every generation. Due to the absence of nine bristles dividing the sternal margin uniformly, the progeny was raised in each generation in the H line from males and females with nine as equidistant bristles as possible. The whole experiment was performed at 25±0.50°C.Selection was effective in increasing the frequency of six regular bristles in the L line. But no progress in the desired direction was obtained in the H line, although the proportion of sternites with nine irregular structures was found to increase. The experimental results supportTuring's diffusion-reaction scheme of pattern formation in morphogenesis.Supported by grants GB-1388 and GB-3219 from the National Science Foundation.     

Morphological chimeras of larvae and adults in a hydrozoan--insights into the control of pattern formation and morphogenesis

In the marine hydroid Hydractinia echinata, metamorphosis transforms the spindle-shaped larva into a primary polyp. It bears a hypostome with a ring of tentacles at its apical end, a gastric region in the middle and stolons at the base. In nature, metamorphosis is induced in response to external stimuli provided by bacteria. These stimuli can be replaced by artificial inducers, one of which is heat shock. Among heat shock treated stages are those undergoing complete metamorphosis but also specimens forming chimeras of different developmental stages. In the chimeric larvae, the posterior is transformed into the apical hypostome of the adult polyp while the anterior part of the larva persists as larval tissue. After transverse sectioning, these stage chimeras regenerate the missing body parts with respect to the nature of the tissue at the wound surface. This shows that the decision to make larva or polyp morphology depends not on the majority of the tissue in the original body section, but on stage specificity within the regenerating animal part. Single cells can escape from this general rule, since RFamide nerve cells which usually differentiate in polyp tissue appear in regenerated larval tails of sectioned stage chimeras. The results indicate that the pattern-forming system of the larva and of the adult have features in common. The primary signals controlling patterning along the anterior-posterior axis in larvae and the apical-basal axis in polyps arethus likelyto be the same while the interpretation of these primary signals by the individual cells changes during metamorphosis.     

Positional information and pattern formation

Spatial patterns of cellular differentiation may arise from cells first being assigned a position, as in a coordinate system, and then interpreting the positional value that they have acquired. This interpretation will depend on their genetic constitution and developmental history. Different patterns may thus arise from similar positional fields. The specification of positional value may involve a positional signal, such as the concentration of a diffusible morphogen, but can also depend on how long the cells remain in a particular region, such as a progress zone. Positional values may also be acquired by direct transfer from one cell layer to another, as in directed embryonic induction. Positional value, unlike a positional signal, involves long-term memory, and can be regarded as a type of cell determination. Cells of the same differentiation class may have different positional values and may thus be non-equivalent. Evidence is presented for a signal providing positional information along the antero-posterior axis during chick limb development. This signal has properties similar to those of a diffusible morphogen.     

Morphogen pattern formation and development in growth

The dependence of the spatial concentration profiles of morphogens on a characteristic dimension is obtained by continuation techniques for Gierer and Meinhardt's activator-inhibitor model of morphogenesis. The study of the behaviour of the system during growth, where the linear and exponential increase of the characteristic dimension is considered, revealed that more complex patterns of morphogen spatial concentrations appear regularly in a reproducible way.