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276 H. PLATTNER ET AL» (stretched and shot out) by exogenous Ca 2+ , once Ca 2+ can come in via the already formed exocytotic canal (B i 1 i n s k i et al. 1981a). Conclusions Mebrane-integrated and membrane-associated proteins may exert a modulating effect on membrane fusion in ciliates, other protozoa and even in metazoan cells. At least some of these systems display conspicuous ultrastructural details which could be interpreted along these lines. The actual membrane fusion appears to be a focally restricted event. In retrospect it appears now that research on membrane fusion in protozoa has provided important clues for a general understanding of membrane fusion processes. Such research appears also important with regard to the interaction of protozoan parasites and host cells. ACKNOWLEDGEMENTS We thank Prof. J. Beisson for providing us generously some of her mutations, for allowing us to reproduce the freeze-fracture micrographs of Fig. 3 and for giving us valuable suggestions and criticism to this article. We also gratefully acknowledge the skilful technical (assistance of Miss C. Braun, Mrs. R. Hildebramd and Mrs. R. Sturzl as well as the secretarial help of Mrs. U. Remensperger. Supported by SFB 138. b REFERENCES Allen R. D. 1978 a: Membranes of ciliates: ultrastructure, biochemistry and fusion. In: Membrane Fusion (eds. G. P o s t e and G. L. N i c o 1 s o n), North- Holland Publishing Company (Amsterdam, New York, Oxford), 657-763. Allen R. D. 1978 b: Particle arrays in the surface membrane of Paramecium: Junctional and possible sensory sites. J. Ultrastruct. Res., 63, 64^78. Bachmann L., Schmitt W. W. and Plattner H. 1972: Improved cryofixatian: Demonstrated on free.ze-etched solutions, cell fractions and unicellular organisms. In: Proc. Fifth Eur. Conf. Electron Microsc. (ed. V. E. C o s s 1 e 11), The Institute of Physics (London, Bristol), 244-245. Beisson J., Cohen J., Lefor t-T ran M., P o u p h i 1 e M. and R o s s i g n o 1 M. 1980: Control of membrane fusion in exocytosis: Physiological studies on a Paramecium mutant blocked in the final step of the trichocyst extrusion process. J. Cell Biol., 85, 213-227. Beisson J., Lefort-Tran M., P o u p hi 1 e M., Rossignol M. and Satir B. 1976: Genetic analysis of membrane differentiation in Paramecium. Freezefracture study of the trichocyst cycle in wild-type and mutant strains. J. Cell Biol., 69, 126-143. Bilinski M., Plattner H. and Matt H. 1981: Secretory protein decondensation as a distinct, Ca 2 +-mediated event during the final steps of exocytosis in Paramecium cells. J. Cell Biol., 88, 179-188. http://rcin.org.pl
277 B i 1 i n s k i M., Plattner H. and Tiggeraamn R. 1981: Isolation of surface membranes from normal and exocytotic mutant strains of Paramecium tetraurelia. Ultrastructural and biochemical characterization. Eur. J. Cell Biol., 24, 108-115. Davidson L. A. 1976: infrastructure of the membrane attachment sites of the extrusomes of Ciliophrys marina and Heterophrys marina (Actinopoda). Cell Tiss. Res., 170, 353-365. Dubremetz J. F. and Tor pier G. 1978: Freeze fracture study of the pellicle of an eimerian sporozoite (protozoa, coccidia). J. Ultrastruct. Res., 62, 94-109. Dubremetz J. F., Torpier G., Maurois P., Prensier G. and S i n d e n R. 1979: Structure de la pellicule du sporozoite de Plasmodium yoelii: etude par cryofracture. C. R. Acad. Sci. Paris, 288, 623-626. H ausmann K. and Allen R. D. 1976: Membrane behavior of exocytic vesicles. II. Fate of the trichocyst membranes in Paramecium after induced trichocyst discharge. J. Cell Biol., 69, 313-326. Henley G. L., Lee C. M. and Takeuchi A. 1976: Freeze-etching observations of trophozoites of pathogenic Entamoeba histolytica. Z. Parasitenk., 48, 181- 190. Hui S. W., Stewart T. P., Boni L. T. and Yeagle P. L. 1981: Membrane fusion through point defects in bilayers. Science, 212, 921-923. Janisch R. 1972: Pellicle of Paramecium caudatum as revealed by freeze etching. J. Protozool., 19, 470-472. Jensen J. B. and Edgar S. A. 1976: Possible secretory function of rhoptries of Eimeria magna during penetration of cultured cells. J. Parasitol., 62, 988-992. Lawson D. and Raff M. C. 1979: Some membrane events occurring during fusion and exocytosis on rat peritoneal mast cells. In: Secretory Mechanisms (eds. C. R. Hopkins and C. J. Duncan), Cambridge University Press (Cambridge), 337-347. Lef ort-Tran M., Aufderheide K., Pouphile M., Rossignol M. and Beisson J. 1981: Control of exocytotic processes: Cytological and physiological studies of trichocyst mutants in Paramecium tetraurelia. J. Cell Biol., 88, 301-311. Matt H., Plattner H., Reichel K., Lef ort-Tran M. and Beisson J. 1980: Genetic dissection of the final exocytosis steps in Paramecium tetraurelia cells: Trigger analyses. J. Cell Sea., 46, 41-60. Mel dole si J., Borgese N., DeCamilli P. and Ceccarelli B. 1978: Cytoplasmic membranes and the secretory process. In: Membrane Fusion (eds. G. Poste and G. L. Ni cols on), North-Holland Publishing Company (Amsterdam, New York, Oxford), 509-627. Nichols B. A. and O'Connor G. R. 1981: Penetration of mouse peritoneal macrophages by the protozoan Toxoplasma gondii: New evidence for active invasion and phagocytosis. Lab. Invest., 44, 324-335. Orci L. and Perrelet A. 1978: Ultrastructural aspects of exocytotic membrane fusion. In: Membrane Fusion (eds. G. Poste and G. L. Nicolson), North- Holland Publishing Company (Amsterdam, New York, Oxford), 629-656. Papahadjopoulos D. 1978: Calcium-induced phase changes and fusion in natural and model membranes. In: Membrane Fusion (eds. G. Poste and G. L. Nicolson), North-Holland Publishing Company (Amsterdam. New York, Oxford), 765-790. Plattner H. 1974: Intramembraneous changes on oationophoire-triggered exocytosis in Paramecium. Nature, 252, 722-724. Plattner H. 1981: Membrane behaviour during exocytosis. Cell Biol. Int. Rep., 5, 435-459. Plattner H.. Miller F. and Bachmann L. 1973: Membrane specializations in the form of regular membrane-to-membrane attachment sites in Paramecium. A correlated freeze-etching and ultrathin-sectioning analysis. J. Cell Sci., 13, 687-719. Plattner H., Wolfram D., Bachmann L. and Wachter E. 1975: Tracer and freeze-etching analysis of intra-cellular membrane-junctions in Paramecium. With a note on a new heme-nonapeptide tracer. Histochemistry, 45, 1-21. http://rcin.org.pl
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277<br />
B i 1 i n s k i M., Plattner H. and Tiggeraamn R. 1981: Isolation of surface<br />
membranes from normal and exocytotic mutant strains of Paramecium tetraurelia.<br />
Ultrastructural and biochemical characterization. Eur. J. Cell Biol.,<br />
24, 108-115.<br />
Davidson L. A. 1976: infrastructure of the membrane attachment sites of the<br />
extrusomes of Ciliophrys marina and Heterophrys marina (Actinopoda). Cell<br />
Tiss. Res., 170, 353-365.<br />
Dubremetz J. F. and Tor pier G. 1978: Freeze fracture study of the pellicle of<br />
an eimerian sporozoite (protozoa, coccidia). J. Ultrastruct. Res., 62, 94-109.<br />
Dubremetz J. F., Torpier G., Maurois P., Prensier G. and S i n d e n R.<br />
1979: Structure de la pellicule du sporozoite de Plasmodium yoelii: etude par<br />
cryofracture. C. R. Acad. Sci. Paris, 288, 623-626.<br />
H ausmann K. and Allen R. D. 1976: Membrane behavior of exocytic vesicles.<br />
II. Fate of the trichocyst membranes in Paramecium after induced trichocyst<br />
discharge. J. Cell Biol., 69, 313-326.<br />
Henley G. L., Lee C. M. and Takeuchi A. 1976: Freeze-etching observations<br />
of trophozoites of pathogenic Entamoeba histolytica. Z. Parasitenk., 48, 181-<br />
190.<br />
Hui S. W., Stewart T. P., Boni L. T. and Yeagle P. L. 1981: Membrane<br />
fusion through point defects in bilayers. Science, 212, 921-923.<br />
Janisch R. 1972: Pellicle of Paramecium caudatum as revealed by freeze etching.<br />
J. Protozool., 19, 470-472.<br />
Jensen J. B. and Edgar S. A. 1976: Possible secretory function of rhoptries<br />
of Eimeria magna during penetration of cultured cells. J. Parasitol., 62,<br />
988-992.<br />
Lawson D. and Raff M. C. 1979: Some membrane events occurring during<br />
fusion and exocytosis on rat peritoneal mast cells. In: Secretory Mechanisms<br />
(eds. C. R. Hopkins and C. J. Duncan), Cambridge University Press<br />
(Cambridge), 337-347.<br />
Lef ort-Tran M., Aufderheide K., Pouphile M., Rossignol M. and<br />
Beisson J. 1981: Control of exocytotic processes: Cytological and physiological<br />
studies of trichocyst mutants in Paramecium tetraurelia. J. Cell Biol.,<br />
88, 301-311.<br />
Matt H., Plattner H., Reichel K., Lef ort-Tran M. and Beisson J.<br />
1980: Genetic dissection of the final exocytosis steps in Paramecium tetraurelia<br />
cells: Trigger analyses. J. Cell Sea., 46, 41-60.<br />
Mel dole si J., Borgese N., DeCamilli P. and Ceccarelli B. 1978: Cytoplasmic<br />
membranes and the secretory process. In: Membrane Fusion (eds.<br />
G. Poste and G. L. Ni cols on), North-Holland Publishing Company (Amsterdam,<br />
New York, Oxford), 509-627.<br />
Nichols B. A. and O'Connor G. R. 1981: Penetration of mouse peritoneal<br />
macrophages by the protozoan Toxoplasma gondii: New evidence for active<br />
invasion and phagocytosis. Lab. Invest., 44, 324-335.<br />
Orci L. and Perrelet A. 1978: Ultrastructural aspects of exocytotic membrane<br />
fusion. In: Membrane Fusion (eds. G. Poste and G. L. Nicolson), North-<br />
Holland Publishing Company (Amsterdam, New York, Oxford), 629-656.<br />
Papahadjopoulos D. 1978: Calcium-induced phase changes and fusion in<br />
natural and model membranes. In: Membrane Fusion (eds. G. Poste and<br />
G. L. Nicolson), North-Holland Publishing Company (Amsterdam. New<br />
York, Oxford), 765-790.<br />
Plattner H. 1974: Intramembraneous changes on oationophoire-triggered exocytosis<br />
in Paramecium. Nature, 252, 722-724.<br />
Plattner H. 1981: Membrane behaviour during exocytosis. Cell Biol. Int. Rep.,<br />
5, 435-459.<br />
Plattner H.. Miller F. and Bachmann L. 1973: Membrane specializations<br />
in the form of regular membrane-to-membrane attachment sites in Paramecium.<br />
A correlated freeze-etching and ultrathin-sectioning analysis. J. Cell<br />
Sci., 13, 687-719.<br />
Plattner H., Wolfram D., Bachmann L. and Wachter E. 1975: Tracer<br />
and freeze-etching analysis of intra-cellular membrane-junctions in Paramecium.<br />
With a note on a new heme-nonapeptide tracer. Histochemistry, 45,<br />
1-21.<br />
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