Organelle division

Recent studies have demonstrated that peroxisome division requires at least one dynamin-like protein, Vpslp, in the yeast Saccharomyces cerevi-siae and DLPl (DRPl) in mammalian cells. Although the requirement for these proteins in peroxisome division is supported by many lines of evidence, their roles in peroxisome division have yet to be identified. Given the independence of peroxisomes from other organelle systems, the peroxisome system appears to have unique attributes for studying the function of dynamin-like proteins in organelle division. Here, we present methods that have been used for studying the role of DLPl in peroxisome biogenesis and division. 

Dynamin-related GTPases (DRPs) have evolved in eukaryotic cells to function in such diverse processes as membrane trafficking, organelle division, and resistance to viral infection (Danino and Hinshaw, 2001 Osteryoung and Nunnari, 2003 Praefcke and McMahon, 2004 Song and... 

Certain proteins endow cells with unique capabilities for movement. Cell division, muscle contraction, and cell motility represent some of the ways in which cells execute motion. The contractile and motile proteins underlying these motions share a common property they are filamentous or polymerize to form filaments. Examples include actin and myosin, the filamentous proteins forming the contractile systems of cells, and tubulin, the major component of microtubules (the filaments involved in the mitotic spindle of cell division as well as in flagella and cilia). Another class of proteins involved in movement includes dynein and kinesin, so-called motor proteins that drive the movement of vesicles, granules, and organelles along microtubules serving as established cytoskeletal tracks. ... 

The cytoplasm of all eukaryotic cells contains a cytoskeletal framework that serves a multitude of dynamic functions exemplified by the control of cell shape, the internal positioning and movement of organelles, and the capacity of the cell to move and undergo division. 

Intracellular motility is also of vital importance in the lives of cells and the organisms they form. Material and organelles are transported within cells along microtubules and microfilaments an extreme example of this are the axons of nerve cells which transport materials to the synapses where they are secreted—another motile event. Other examples of intracellular motility include phagocytosis, pino-cytosis, the separating of chromosomes and cells in cell division, and maintenance of cell polarity. 

A three-dimensional meshwork of proteinaceous filaments of various sizes fills the space between the organelles of all eukaryotic cell types. This material is known collectively as the cytoskeleton, but despite the static property implied by this name, the cytoskeleton is plastic and dynamic. Not only must the cytoplasm move and modify its shape when a cell changes its position or shape, but the cytoskeleton itself causes these movements. In addition to motility, the cytoskeleton plays a role in metabolism. Several glycolytic enzymes are known to be associated with actin filaments, possibly to concentrate substrate and enzymes locally. Many mRNA species appear to be bound by filaments, especially in egg cells where they may be immobilized in distinct regions thereby becoming concentrated in defined tissues upon subsequent cell divisions. 

Division and segregation of organelles. Edited by 5.A. Boffey and D. Lloyd... 

Until the past decade, the cytoplasm was widely considered to be structurally unorganized with the main division of labor at the organellar level. Certainly, relatively little was known about the nature of the cyto-skeleton (with the notable exception of the mitotic apparatus and striated muscle), and the dynamics of cytoplasmic behavior were conceptualized vaguely in terms of sol-gel transitions without a sound molecular foundation. Substantial improvements in electron, light, and fluorescence microscopy, as well as the isolation of discrete protein components of the cytoskeleton, have led the way to a much better appreciation of the structural organization of the cytoplasm. Indeed, the lacelike network of thin filaments, intermediate filaments, and microtubules in nonmuscle cells is as familiar today as the organelles identified... 

One of the major features in the sequence of cell division is the formation of the mitotic spindle and the subsequent separation of chromosomes into their respective daughter cells. An important element of the spindle is the highly conserved, helical molecule tubulin. In addition to spindle formation and the segregation of chromosomes in cell division, alternating helices of a- and -tubulin form the microtubules that form part of the cytoskeleton and have active roles in cell organelle organisation. 

Mitochondrial DNA codes for the mitochondrial tRNAs and rRNAs and for a few mitochondrial proteins. More than 95% of mitochondrial proteins are encoded by nuclear DNA. Mitochondria and chloroplasts divide when the cell divides. Their DNA is replicated before and during division, and the daughter DNA molecules pass into the daughter organelles. 

As we will see, the evolutionary tree is bisected into a lower prokaryotic domain and an upper eukaryotic domain. The terms prokaryote and eukaryote refer to the most basic division between cell types. The fundamental difference is that eukaryotic cells contain a membrane-bounded nucleus, whereas prokaryotes do not. The cells of prokaryotes usually lack most of the other membrane-bounded organelles as well. Plants, fungi, and animals are eukaryotes, and bacteria are prokaryotes. The biochemical functions associated with organelles are frequently present in bacteria, but they are usually located on the inner plasma membrane. 

Hirokawa, N., Noda, Y., and Okada, Y. (1998). Kinesin and dynein superfamily proteins in organelle transport and cell division. Curr. Opin. Cell Biol. 10, 67-73. 

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