Cell polarity

Cell polarity refers to spatial differences in shape, structure, and function within a cell. Almost all cell types exhibit some form of polarity, which enables them to carry out specialized functions. Classical examples of polarized cells are described below, including epithelial cells with apical-basal polarity, neurons in which signals propagate in one direction from dendrites to axons, and migrating cells. Furthermore, cell polarity is important during many types of asymmetric cell division to set up functional asymmetries between daughter cells. Cell polarity refers to spatial differences in shape, structure, and function within a cell. Almost all cell types exhibit some form of polarity, which enables them to carry out specialized functions. Classical examples of polarized cells are described below, including epithelial cells with apical-basal polarity, neurons in which signals propagate in one direction from dendrites to axons, and migrating cells. Furthermore, cell polarity is important during many types of asymmetric cell division to set up functional asymmetries between daughter cells. Many of the key molecular players implicated in cell polarity are well conserved. For example, in metazoan cells, the PAR-3/PAR-6/aPKC complex plays a fundamental role in cell polarity. While the biochemical details may vary, some of the core principles such as negative and/or positive feedback between different molecules are common and essential to many known polarity systems. Epithelial cells adhere to one another through tight junctions, desmosomes and adherens junctions, forming sheets of cells that line the surface of the animal body and internal cavities (e.g., digestive tract and circulatory system). These cells have an apical-basal polarity defined by the apical membrane facing the outside surface of the body, or the lumen of internal cavities, and the basolateral membrane oriented away from the lumen. The basolateral membrane refers to both the lateral membrane where cell-cell junctions connect neighboring cells and to the basal membrane where cells are attached to the basement membrane, a thin sheet of extracellular matrix proteins that separates the epithelial sheet from underlying cells and connective tissue. Epithelial cells also exhibit planar cell polarity, in which specialized structures are orientated within the plane of the epithelial sheet. Some examples of planar cell polarity include the scales of fish being oriented in the same direction and similarly the feathers of birds, the fur of mammals, and the cuticular projections (sensory hairs, etc.) on the bodies and appendages of flies and other insects. A neuron receives signals from neighboring cells through branched, cellular extensions called dendrites. The neuron then propagates an electrical signal down a specialized axon extension to the synapse, where neurotransmitters are released to propagate the signal to another neuron or effector cell (e.g., muscle or gland). The polarity of the neuron thus facilitates the directional flow of information, which is required for communication between neurons and effector cells. Many cell types are capable of migration, such as leukocytes and fibroblasts, and in order for these cells to move in one direction, they must have a defined front and rear. At the front of the cell is the leading edge, which is often defined by a flat ruffling of the cell membrane called the lamellipodium or thin protrusions called filopodia. Here, actin polymerization in the direction of migration allows cells to extend the leading edge of the cell and to attach to the surface. At the rear of the cell, adhesions are disassembled and bundles of actin microfilaments, called stress fibers, contract and pull the trailing edge forward to keep up with the rest of the cell. Without this front-rear polarity, cells would be unable to coordinate directed migration. Generation of cellular polarity is critically important for the function of practically every cell type and underlies processes like cell division, differentiation, cell migration, cell–cell signalling and fertilization. Cell polarity is an example of the self-organization property that all living organism share. All the cells within a multicellular organism, or any single cell species i.e. yeast, displays a polarized organization necessary for its proliferation, differentiation or physiological function. Budding yeast is a highly accessible experimental system, which serves as a paradigm for deciphering the molecular mechanism underlying the generation of polarity. Yeast cells share many features about cell polarity with other organism like: regulation by intrinsic and extrinsic cues, conserved regulatory molecules such as Cdc42GTPase, and asymmetry of the cytoskeleton. Cell polarization is associated with a type of symmetry breaking, that occurs in the cytoskeleton and guides the direction of growth of the future daughter cell. This symmetry breaking facilitates the polarized flux and localization of several protein in the polarized patch. When cells can perform symmetry breaking in absence of any spatial cue (landmarks), is called spontaneous polarization or spontaneous symmetry breaking. In short, polarity establishment or symmetry breaking, in this context, is the first step for cell polarity and consequently cell division. We are interested in the spontaneous symmetry breaking, as an example of self-organization phenomena in living cells. One of the way, and one of the most popular ones, to study spontaneous symmetry breaking is by correlating it with a type of reaction- diffusion phenomena. The molecular identity of most important proteins involved in leading spontaneous symmetry breaking have been extensively studied. Those proteins have been categorized in general modules that represent the main functional cores for yeast life cycle. However, the molecular mechanisms responsible for this regulatory network are still poorly understood. Extensive work done in many organisms ranging from prokaryotes to high complex ones has revealed a central role of small GTPases in the cell polarization process. Particularly in yeast, this protein is Cdc42, which is a member of the eukaryotic Ras-homologous Rho-family of GTPases, which is part of the wider super-family of small GTPases, including Rop GTPases in plants and small GTPases in prokaryotes. A recent study to elucidate the connection between cell cycle timing and Cdc42 accumulation in the bud site uses optogenetics to control protein localization using light.