Eukaryotic cells, movement

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. 

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. 

Microtubules, an integral component of the cellular cy-toskeleton, consist of cytoplasmic tubes 25 nm in diameter and often of extreme length. Microtubules are necessary for the formation and function of the mitotic spindle and thus are present in all eukaryotic cells. They are also involved in the intracellular movement of endocytic and exocytic vesicles and form the major structural components of cilia and flagella. Microtubules are a major component of axons and dendrites, in which they maintain structure and participate in the axoplasmic flow of material along these neuronal processes. 

Active transport of a solute against a concentration gradient also can be driven by a flow of an ion down its concentration gradient. Table 17.6 lists some of the active-transport systems that operate in this way. In some cases, the ion moves across the membrane in the opposite direction to the primary substrate (antiport) in others, the two species move in the same direction (symport). Many eukaryotic cells take up neutral amino acids by coupling this uptake to the inward movement of Na+ (see fig. 17.26c). As we discussed previously, Na+ influx is downhill thermodynamically because the Na+-K+ pump keeps the intracellular concentration of Na+ lower than the extracellular concentration and sets up a favorable electric potential difference across the membrane. Another example is the /3-galactosidc transport system of E. coli, which couples uptake of lactose to the inward flow of protons (see fig. 17.26Proton influx is downhill because electron-transfer reactions (or,... 

Eukaryotic cells have an internal scaffold, the cytoskeleton, that controls the shape and movement of the cell. The cytoskeleton is made up of actin microfilaments, intermediate filaments and microtubules. 

The size of the genomic DNA in eukaryotic cells (such as the cells of yeast, plants, or mammals) is much larger (up to 10+11 base pairs) than in E. coli (ca. 10+6 base pairs). The rate of the eukaryotic replication fork movement is about fifty nucleotides per second, which is about ten times slower than in E. coli. To complete replication in the relatively short time periods observed, multiple origins of replication are used. In yeast cells, these multiple origins of replication are called autonomous replication sequences (ARSs). As with prokaryotic cells, eukaryotic cells have multiple DNA polymerases. DNA polymerase S, complexed with a protein called proliferating... 

Eukaryotic cells have an internal scaffolding called the cytoskeleton or cytomatrix that maintains their cellular morphology and enables them to migrate, undergo shape changes, and transport vesicles. Microfilaments, made of actin, intermediate filaments, which are composed of laminin and other proteins, and microtubules, formed from the protein tubulin, along with many different accessory proteins, comprise the cytoskeleton. Both the microfilaments and the microtubules can assemble and disassemble rapidly in the cell, whereas disassembly of intermediate filaments may require their destruction. Although much is known about the molecular composition of the cytoskeleton, the molecular events involved in most cell movements are still unknown. 

The mitotic spindle is part of a larger intracellular skeleton (cytoskele-ton) that is essential for the internal movements occurring in the cytoplasm of all eukaryotic cells. The mitotic spindle consists of chromatin, and a system of microtubules composed of the protein tubulin. The mitotic spindle is essential for the equal partitioning of DNA into the two daughter cells formed when a eukaryotic cell divides. Several plant-derived substances used as anticancer drugs disrupt this process by affecting the equilibrium between the polymerized and depolymerized forms of the microtubules, thereby causing cytotoxicity. 

Cilia and flagella are stable microtubule-based structures which project from the plasma membranes of particular eukaryotic cells. The energy-dependent oscillations of these structures can drive material over the surface of a cell or propel the cell along. For example, the whip-like motions of cilia on the cells at the head of the fallopian tube draw newly released ova from the ovaries into and along the oviduct. The snake-like movements of the flagellum on a sperm provide these cells with movement. 

Cytoskeleton is defined as the sum of the various filamentous proteins of eukaryotic cells that remain after the cells are extracted with a mild detergent. The cytoskeleton includes actin filaments, two-stranded helical polymers, which form the microfilaments and the actin-binding proteins. Other components are microtubules and intermediate filaments. The cytoskeleton has not only a role in maintaining the shape of cells, it is also actively engaged in cell division, in the organisation and the dynamic movement of ceD organelles and in the movement of cells in chemotaxis. 

Several important groups of plant compounds, including cytokinins, chlorophylls and the quinone-based electron carriers (the plastoquinones and ubiquinones), have terpenoid side chains attached to a non-terpenoid nucleus. These side chains facilitate anchoring to or movement within membranes. In the past decade, proteins have also been found to have terpenoid side chains attached. In fact, all eukaryotic cells appear to contain proteins that have been post-translationally modified by the attachment of C15 and C20 terpenoid side chains via a thioether linkage. 

The nucleus is a large membrane-bound cell organelle which houses the chromosomes and which occupies roughly 10% of the volume of all eukaryotic cells. The nucleus is separated from the rest of the cell and the cytoplasm by a double membraue known as the nuclear envelope. The outer layer of the nuclear membrane is studded with small openings called nuclear pores, which allow for the controlled movement of selected molecules in and out of the nucleus. Most of a eukaryotic cell s DNA is found in the chromosomes of the nucleus, while a very small amount of DNA is present in the mitochondria. All plant and animal cells with a nucleus and known as eukaryotic cells, (meaning true nucleus) while bacterial cells which lack a nucleus are known as prokaryotic cells. 

Actin is present in all eukaryotic cells where it has structural and mobility functions. Most movement associated with microfilaments requires myosin. The myosin-to-actin ratio is much lower in nonmuscle cells, and myosin bundles are much smaller (10-20 molecules rather than about 500), but the interaction between myosin and actin in nonmuscle cells is generally similar to that in muscle. As in smooth muscle, myosin aggregation and activation of the actin-myosin interaction are regulated primarily by light chain phosphorylation. Myosins involved in transporting organelles along actin filaments are often activated by Ca-CaM. 

Actin. This is one of the most important components in the cytoskeletal architecture, and in the movement of cytoplasm in eukaryotic cells. In addition to muscle contraction, actin is involved in cellular processes such as phagocytosis, secretion, cell migration, and the maintenance of cell shape. A recent report shows that a mammalian cell arginine-specific ADP-ribosyl transferase can ADP-ribosylate actin molecules, suggesting that this could be a regulatory mechanism in the above cellular functions (Terashima et al., 1992). 

All biomembranes form closed structures, separating the lumen on the inside from the outside, and are based on a similar bllayer structure. They control the movement of molecules between the inside and the outside of a cell and into and out of the organelles of eukaryotic cells. In accord with the Importance of Internal membranes to cell function, the total surface area of these membranes is roughly tenfold as great as that of the plasma membrane (Figure 5-1). 

A FIGURE 5-1 Schematic overview of the major components of eukaryotic cell architecture. The plasma membrane (red) defines the exterior of the cell and controls the movement of molecules between the cytosol and the extracellular medium. Different types of organelles and smaller vesicles enclosed within their own distinctive membranes (black) carry out special functions such as gene expression, energy production, membrane synthesis, and intracellular transport. 

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