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288 Miroslav Ovečka, Milan Bobák, Jozef Šamaj Shoot regeneration in callus culture requires a combination of auxins and cytokinins (Christianson and Warnick 1983, De Klerk et al. 1997). The main change in cell division and cell expansion was observed during competence acquisition. Meristemoids were produced from rarely dividing callus cells. Cell division was more patterned within meristemoids. Their cells were small, micro-vacuolated, and able to stick together. This indicates that the fate of the callus culture is altered through cell morphogenesis of the activated competent cells as a result of the action of growth regulators. The second switching point in cell morphogenesis of poppy callus culture occurred in the multicellular meristemoids as a defined step of cell determination within the shootforming tissue. A critical event was the localisation of cell division to the meristemoid periphery. The cell determination has been expressed when modified polar periclinal cell division occurred in meristemoids. Apical daughter cells generated dividing peripheral layers with morphogenetic competence while the distal daughter cells became more differentiated and generated storage cells possessing metabolic sources. The previously observed differences in cell size and shape showed that the peripheral cells were smaller, having an oblong shape as a result of frequent cell division, while the central cells became larger due to globular expansion (Ovečka et al. 1997). Starch-containing cells were the largest ones. They had isodiametric shape, and any changes (cell division, plastid division, vacuolation) took place only after starch utilisation had been initiated (not shown). Central meristemoid cells thus can support organogenesis by the energy. It was clearly shown that some metabolic indications of shoot regeneration (protein inclusions, starch, lipids, high enzyme activities) were found only in meristemoids (Ross et al. 1973, Nessler and Mahlberg 1979, Patel and Berlyn 1983). Meristemoids possessed a high starch content, while high activities of starch-degrading enzymes were observed in shoot buds (Thorpe and Murashige 1970, Thorpe and Meier 1974, Patel and Berlyn 1983). A high N/C ratio of peripheral meristemoid cells of opium poppy, comparable with cells in shoot apical meristem (Fig. 5), together with comparable rate of chromatin de-condensation (Ovečka et al. 1997) are also important parameters of cell determination within the meristemoids. Variability in the size of nucleus and the correlation with DNA amount is closely related to shoot regeneration (Flinn et al. 1989, Fournier et al. 1991). A two-phase organogenesis, with meristemoid formation during the first phase and formation of globular structures from the surface during the second phase, was observed in sundew callus tissue (Bobák et al. 1993). This data indicates that “mitotic zonation” established a distinct morphological competence of meristemoid cells in the process of shoot primordia formation when the site of origin of shoot buds was located at the meristemoid periphery. Concluding our comparative analysis of direct and indirect shoot regeneration in P. somniferum L., important changes in cell morphogenesis, characterised by non-random cell divisions, took place during dermal layer and protomeristem formation in shoot primordia. We observed a random orientation of cell divisions in meristemoids, while protomeristem and especially tunica formation were accompanied by regulated periclinal and subsequently anticlinal cell divisions. In directly regenerated shoots, the dermal layer was formed by original epidermal cells through anticlinal cell divisions. This type of division maintains epidermal cell polarity. Peripheral meristemoid cells are polarised with non-homogenous cell surfaces due to the presence of adhered and outer non-adhered cell walls (Ovečka and Bobák 1999). This polarity and cell position are important morphogenetic factors in cell determination. Cell expansion was found to be minimal in cells involved in formative, morphogenetically important cell division. The cells expanded, but not over the cell volume necessary for a subsequent cell division. In addition, cell size and cell shape have to be negatively correlated. This relationship would indicate an independence of post-mitotic cell growth and an extensive cell vacuolation, originally observed in maize root apex (Baluška et al. 1990). A comparative analysis revealed that cell activation in shoot organogenesis of P. somniferum L. depends on the stage of cell differentiation before initiation, while cell morphogenesis of the determined cells is similarly regulated both spatially and temporally, during direct and indirect shoot regeneration. Acknowledgements. We gratefully acknowledge D. Jantová for technical assistance, and Dr. M. Čiamporová for the English correction. This work was supported by Grant Agency VEGA, Grant No.1/6030/99 and Grant No. 1/6107/99 from the Slovak Academy of Sciences. M.O. gratefully acknowledges the Alexander von Humboldt-Stiftung (Bonn, Germany) for donation of the Leica Q500MC image analysis system. References Altamura MM, Capitani F, Falasca G, Gallelli A, Scaramagli S, Bueno M, Torrigiany P, Bagni N (1995) Morphogenesis in cultured thin layers and pith explants of tobacco. I. Effect of putrescine on cell size, xylogenesis and meristemoid organization. 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