[Endocytosis and actin-dependent transport of plant cell wall components]. / Endocytoza i zalezny od cytoszkieletu transport skladników scian komórkowych roslin.

Endocytosis is the way of uptaking and transporting substances from the immediate surroundings, as well as the way of removal and/or recycling of the plasma membrane components. For many years, it was thought that this process was not operating in plant cells in which high turgor pressure of the protoplast prevented invagination of the plasma membrane. Recent years, however, brought the evidence for intensive endocytosis in plants. So far, four types of endocytosis, among 5 known, have been identified in plants. Use of protein markers of different compartments of the endomembrane system, specific dyes, and the cytoskeleton inhibitors, made possible demonstration that, as opposed to animal and fungal systems, endocytosis in plant cells was dependent on actin and plant-specific myosins of VIII and XI classes. Although cell divisions in plants and animals differ considerably, they both are dependent on the proper endocytosis. It has been shown that during cytokinesis secretion and endocytosis act in concert, delivering polysaccharides directionally to the growing cell plate. What is more important, new cell walls are built from e.g. pectins transported by endocytosis from existing cell walls.

[At the plant side of formins--organizers of the actin cytoskeleton]. / Roslinna twarz formin--organizatorów cytoszkieletu aktynowego.

Rearrangements of actin cytoskeleton enable proper functioning of the cells under normal conditions, and also cellular adaptations to changes in the direct surroundings. Formins are actin binding proteins, responsible for actin nucleation and further elongation of microfilaments. The distinguishing feature of formins is the presence of conserved FH2 (formin homology domain 2) domain, as well as other domains typical for distinct formin classes. In animal cells formins are involved in cytokinesis and determination and maintenance of the cell shape and polarity, but also in the formation of filopodia, endocytosis and many other processes. The presence of proteins from the formin family in plant cells, and their involvement in the tip growth and cytokinesis, has been determined only recently. As the functional organization of plant and animal cells is different, one can assume that the range of putative functions of plant formins might also be diverse. One of such proposed functions for formins in plants is the role of linker protein within WMC continuum (cell wall-plasma membrane-cytoskeleton). Unfortunately, for that moment the state of knowledge about plant formins in comparison with animal or fungal ones is much poorer.

Nitric oxide modulates dynamic actin cytoskeleton and vesicle trafficking in a cell type-specific manner in root apices.

NO is an important regulatory molecule in eukaryotes. Much of its effect is ascribed to the action of NO as a signalling molecule. However, NO can also directly modify proteins thus affecting their activities. Although the signalling functions of NO are relatively well recognized in plants, very little is known about its potential influence on the structural integrity of plant cells. In this study, the reorganization of the actin cytoskeleton, and the recycling of wall polysaccharides in plants via the endocytic pathway in the presence of NO or NO-modulating substances were analysed. The actin cytoskeleton and endocytosis in maize (Zea mays) root apices were visualized with fluorescence immunocytochemistry. The organization of the actin cytoskeleton is modulated via NO levels and the extent of such modulation is cell-type specific. In endodermis cells, actin cables change their orientation from longitudinal to oblique and cellular cross-wall domains become actin-depleted/depolymerized. The reaction is reversible and depends on the type of NO donor. Actin-dependent vesicle trafficking is also affected. This was demonstrated through the analysis of recycled wall material transported to newly-formed cell plates and BFA compartments. Therefore, it is concluded that, in plant cells, NO affects the functioning of the actin cytoskeleton and actin-dependent processes. Mechanisms for the reorganization of the actin cytoskeleton are cell-type specific, and such rearrangements might selectively impinge on the functioning of various cellular domains. Thus, the dynamic actin cytoskeleton could be considered as a downstream effector of NO signalling in cells of root apices.

Domain-specific mechanosensory transmission of osmotic and enzymatic cell wall disturbances to the actin cytoskeleton.

Plant protoplasts are embedded within surrounding cell walls and the cell wall-plasma membrane-cytoskeleton (WMC) structural continuum seems to be crucial for the proper functioning of plant cells. We have utilised the protoplast preparation methodology to study the organisation and the putative components of the WMC continuum. Application of an osmotic agent evoked plasmolysis of the Zea mays root apex cells which appeared to be cell type- and growth stage-specific. Simultaneous use of wall polysaccharide-digesting enzymes selectively severed linkages between the components of the WMC continuum which changed the plasmolytic patterns in various cell types. This was followed by a reorganisation of filamentous actin aimed to reinforce protoplast boundaries and maintain the functioning of intercellular contact sites, especially at the cross walls. Particularly strong effects were evoked by pectin-degrading enzymes. Such treatments demonstrated directly the differentiated composition of various wall domains surrounding individual cells with the pectin-enriched cross walls (synapses), and the cellulose-hemicellulose network dominating the side walls. The same wall-degrading enzymes were used for in vitro digestion of isolated Lupinus albus cell walls followed by the extraction of wall proteins. Selective release of proteins suggested the importance of wall polysaccharide-protein interactions in the maintenance of the functioning and mechanical stability of root cell walls.