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164 Pattern Formation the Dependence of DNA and Protein 164 Pattern formation The dependence of DNA and protein biosynthesis on cytoplasmic pH during the cell cycle in Dictyostelium discoideum Rob Aerts, Tony Durston and Wouter Moolenaar, Hubrecht Laboratory, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands We measured cytoplasmic pH in Dictyostelium discoideum under various conditions, using the digitonin null-point method (Rink, T. J., Tsien, R. Y., Pozzan, T. (1982) /. Cell Bioi, 95,189. In synchronised cell populations the cytoplasmic pH clearly fluctuates during the cell cycle. These fluctuations coincide with fluctuations in the rates of biosynthetic processes; the rates of DNA synthesis and protein synthesis show a sharp optimum in the cytoplasmic pH range 7-40 to 7-45. The cytoplasmic pH of Dictyostelium discoideum can be changed by certain weak acids and bases. The effect of ammonia is concentration dependent; a low concentration is able to increase cytoplasmic pH, high concentrations have the opposite effect. Artificial manipulation of cytoplasmic pH influences the rate of DNA and protein synthesis and shows the same correlation between these processes and cytoplasmic pH as mentioned above. On the other hand, if DNA synthesis is inhibited in synchronous populations of cells then the cell cycle fluctuations in cytoplasmic pH continue. The interrelation between cytoplasmic pH and DNA synthesis in Dictyostelium discoideum as revealed by the experiments summed up above corresponds strikingly to the relation between intracellular pH and DNA synthesis established in a taxonomically quite different system, namely lymphocytes of the mouse (Gerson, D. F., Kiefer, H., Eufe, W. (1982) Science 216,1009). That in Dictyostelium discoideum, on the one hand, artificial manipulation of cytoplasmic pH influences the rates of biosynthetic processes, and on the other hand, fluctuations in cytoplasmic pH continue after DNA synthesis inhibition suggests that the cytoplasmic pH may be a general regulator of the cell cycle. The differentiation direction of a Dictyostelium discoideum cell depends on the cell cycle phase in which it enters development (McDonald, S., Durston, A. J., /. Cell ScL, in press) and as we have shown, the cytoplasmic pH (and the concomitant rates of biosynthetic processes) changes with the cell cycle phase. Furthermore a recent model proposes that a low intracellular pH favours stalk cell differentiation and a high intracellular pH favours spore differentiation (Gross, J. D. etal. (1983). Nature, 303, 244). With regard to this it is particularly interesting to note that we find higher rates of 3H-uridine incorporation in the prespore region of the slug than in the prestalk region. Morphogenesis in the chick limb Karen Bell and J. C. McLachlan, Oxford University, Dept. Zoology, South Parks Road, Oxford Two regions of the chick wing bud have been shown to influence pattern and growth during development. The apical ectodermal ridge (AER) is essential for the outgrowth of the limb and the formation of the proximo-distal axis (Summerbell, 1974). The AER appears to influence the positional value of cells in the distal mesoderm by controlling their density and division rates (Summerbell & Lewis, 1975). The zone of polarizing activity (ZPA) is an area of cells at the base of the limb bud which, when transplanted anteriorly, induces the formation of supernumerary limb structures along the antero-posterior axis (Saunders & Gasseling, 1968). Considerable growth is observed in the limb bud following ZPA grafting, which results in the host tissue doubling within 36 hours (Cooke & Summerbell, 1980). To investigate cell division and pattern formation in vitro we cultured ZPA and AER cells with an embryonic cell line (mouse 3T3 cells) and measured changes in the initiation of DNA synthesis by the uptake of labelled thymidine. At the same time distal and posterior tissue were grafted anteriorly into host wing buds. These were either labelled with tritiated thymidine 16-19 hours after operating, and sectioned, or allowed to develop for 7 days to assess gross cartilage patterns. Anterior wing tissues was used as a control in all the experiments. So far we have shown that chick tissue can induce 3T3 cells to initiate DNA synthesis, while conditioned media and non-contact experiments suggest that a diffusible substance is involved. COOKE, J. & SUMMERBELL, D. (1980). Cell cycle and experimental pattern duplication in the chick wing during embryonic development. Nature, 287, 697-701. SAUNDERS, J. W. & GASSELING, M. T. (1968). Ectodermal-mesenchymal interactions in the origin of limb symmetry. In Epithelial-Mesenchymal Interactions (ed. R. Fleischmajer & R. E. Billingham), pp. 78-97. Baltimore, Williams & Wilkins. SUMMERBELL, D. (1974). A quantitative analysis of the effect of excision of the AER from the chick limb bud. /. EmbryoL exp. Morph. 32, 651-660. SUMMERBELL, D. & LEWIS, J. H. (1975). Time place and positional value in the chick limb bud. /. Embryol. exp. Morph. 33, 621-643. Pattern formation 165 Metamorphosis and pattern formation in Hydractinia echinata S. Berking, Zoologisches Institut, INF'230, D-6900 Heidelberg, West Germany Hydractinia echinata is a marine, colony forming Coelenterate. Fertilized eggs develop into freely swimming planula larvae, which undergo metamorphosis to a sessile (primary) polyp. Metamorphosis can be triggered by means of certain marine bacteria and by Cs+ ions. Half a day after such a treatment a larva will have developed into a polyp. The induction of metamorphosis can be prevented by addition of inhibitor I, a substance partially purified from tissue of Hydra. The larvae of Hydractinia echinata also appear to contain this substance. Inhibitor I applied after the onset of metamorphosis blocks its continuation as long as it remains in the culture medium. Cs+ ions applied within the same time period also block the continuation of metamorphosis. However, these two agents have opposite effects on the body pattern of the resultant polyps. The experiments indicate that the application of Cs+ ions trigger the generation of the prepattern. Inhibitor I appears to be an element of this prepattern. Anterior-like cells may play a role in cell type proportioning in Dictyostelium Angela Blaschke, Cornells Weijer, Harry MacWilliams*, Zoologisches Institut, Luisenstrasse 14, 8000Munchen2, West Germany Most discussions of cell type proportioning in Dictyostelium consider only the prestalk and prespore cells; the 'anterior-like' cells (resembling prestalk cells but dispersed in the prespore zone) are ignored. We have investigated the role of anterior-like cells by studying these cells in chimeric slugs; the chimeras were mixtures of prestalk-prespore proportioning mutants with their parental type or of axenic cells grown with (G+) and without (G—) glucose. The cells were mixed in the vegetative phase; in each case one of the cell types had been labelled with tetramethylrhodamine isothiocyanate. Chimeric slugs were dissected into prestalk and prespore zones and disaggregated into single cells. MUD-1 (a monoclonal antibody against prespore cells, donated by M. Krefft) was used to distinguish prespore from anterior-like cells in the disaggregated prespore tissue. We could thus determine the ratio of mutant to wild type (or G+ to G-) among prestalk, prespore, and anterior-like cells. The short-prestalk mutant HS2 shows a reduced fraction of prestalk cells but a normal complement of anterior-like cells. In mixtures of HS2; with its parent HS1 the prespore cells are enriched in HS2; the ratio of mutant to parent among anterior-like cells is essentially the same as that among prespore cells. The long-prestalk mutant HS3 shows an increased fraction of both prestalk and anterior-like cells. In mixtures of HS3 and HS1 the prestalk zone is enriched in HS3; the HS3: HS1 ratio in the anterior-like cells is essentially the same as in the prestalk cells. Similar behaviour is seen in mixtures of G- and G+ cells. These findings cannot be explained with a 'two-threshold model' in which different levels of the same parameter are necessary for prestalk and anterior-like differentiation. The findings support a model in which two. independent feedback loops regulate cell type proportions in slime molds. One loop controls the proportion of anterior-like cells to prespore cells; the other controls the ratio of anterior-like cells to prestalk cells. HS3 and G— cells are modified in sensitivity to the feedback signal in the first loop (but not in the second); HS2 is modified in sensitivity to the feedback signal of the second loop (but not in the first). It is conceivable that there is no direct interaction between prestalk and prespore cells. 166 Pattern formation Proportion regulation in hydra is complex Hans R. Bode* and Patricia M. Bode, Developmental Biology Center, University of California, Irvine, California, CA 92717, U.S.A. Excision of a piece of the body column of hydra results in regeneration of a diminutive version of the animal in a morphallactic manner. As part of the patterning that occurs, tissue is allocated to each of the structures: hypostome, tentacle zone, tentacles, body column, and basal disk. An approach to the processes underlying this allocation is to examine the extent to which proportion regulation occurs. If a high degree proportion regulation exists, a constant fraction of the tissue will be allotted to each structure independent of the size of piece and the regenerate would be a quite precise miniature of a normal adult. Excised pieces of the body column varying 40-fold in size were analyzed to determine quantitatively how the pattern changes as it is constrained to form in less and less tissue. Alterations were found in tissue allocation, body shape and tentacle number. Only the total number of cells in the basal disk and in the tentacles remained proportional to the whole animal. With decreasing size, the hypostome and tentacle zone increased allometrically at the expense of body tissue. The body column circumference followed the circumference of the head, which was proportionately larger in smaller animals, resulting in increasingly wider and squatter columns.
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