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Asignatura: Biología, Profesor: Javier Regadera, Carrera: Medicina, Universidad: UAM
Tipo: Apuntes
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University of Exeter, Exeter, UK
Movement is a major characteristic of living organisms, and can take the form either of movements of cells or of movements within cells themselves.
Movement is a major characteristic of living organisms, and can take the form either of movements of cells or of movements within cells themselves. Lower prokaryotic cells, such as some bacteria, are able to swim within their environment with the aid of fine appendages, but the motile
Introductory article
Article Contents
repertoire exhibited by higher eukaryotic cells is much greater. Higher cells may also have beating appendages in the form of numerous short cilia or fewer longer flagella, and in unicellular protozoa these function in a coordinated way, not only to propel the organism but also to assist feeding. Cilia are also present on cells of certain differentiated tissues of metazoans, where their beating serves to move the environment relative to the stationary cells. Not all protozoa move by means of cilia and flagella. Some, such as freshwater and soil amoebae, crawl over their substrates, and a variety of cells, such as leucocytes, in multicellular organisms show similar crawling movements. Higher cells, as well as being much larger than bacteria, have separate nuclear and cytoplasmic compartments and often show considerable asymmetry. This has necessitated the development of mechanisms for moving components intracellularly, a property that is well illustrated by plant cells, many of which show different forms of exaggerated cytoplasmic streaming. Intracellular movement is also an important feature of animal cells in which organelles, vesicles and genetic
messages are transported to and from specific sites within the cell. Replicated genetic material within chromosomes is also separated into two prospective daughter cells by similar mechanisms. Many cells are capable of shape changes as a result of reversible or irreversible contraction. The most obvious examples of cellular contraction are seen in muscle tissue. Similar contractions, however, also occur in cells of other tissues, and the division of a cell’s cytoplasm following the separation of its genetic material at mitosis is the result of a drawstring-like contraction. A further type of movement is typified by fibroblasts, cells of mesodermal origin, which migrate individually through connective tissue and synthesize collagen in the extracellular matrix. Their movement differs from that of amoebae in that they do not show continual deformation during migration, but glide over their substrate as a result of extension, attachment and contraction of a broad, flattened, leading edge. Indeed, in a similar fashion the growth cone at the tip of axons of differentiating nerve cells
is thought to extend, interact with its substrate, and contract to extend the axon and thereby innervate tissues. As well as moving independently, cells change shape and move as aggregates during development. This is seen most obviously and dramatically as early as gastrulation, following extensive cleavage of the fertilized egg, when interrelated movements of sheets of cells lay down the different cell layers of the embryo. The movement of aggregates of cells also occurs extensively at later stages of development in numerous well-documented examples of organogen esis. The cytoskeleton forms the basis for most of the active movements exhibited by higher cells. In some instances motility is generated simply by its regulated assembly and disassembly. In others cases motility results from the activities of ‘motor’ proteins which interact with the different cytoskeletal elements, and both phenomena are discussed elsewhere.
Swimming Cells: Bacteria, Ciliates and Flagellates, Sperm
Many bacteria can sense various aspects of their surround- ings and respond by moving towards or away from stimuli, for example by migrating up and down chemical gradients of attractants and repellents. Some motile bacteria glide over surfaces. Other bacteria, such as Escherichia coli, swim by means of a number of fine helical filamentous flagella approximately 20 nm in diameter and several micrometres in length. These act as propellers, rotating rapidly at about 300 Hz, driven by a rotary motor at their base. The energy for driving the motor derives directly from the transmembrane proton gradient. The rotatory motors on any one cell are capable of turning in both directions, and their collective direction determines the swimming path. Observations have shown that when all the flagellar motors are rotating in the same direction the flagella form a bundle and the bacterium is propelled
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net 1
Cell Motility
As well as propelling unicells, cilia propel multicellular organisms and embryos. Indeed, most organisms possess cilia or flagella at some stage in their life cycle. They also occur extensively on the epithelia of many organisms, where they serve to move fluid over the cell surfaces. The activities of ciliated epithelia serve a wide range of functions and in humans, for example, occur in the lungs and airways, the eustachian tubes, the middle ear, the pharynx, the lining of the brain, as well as in the female reproductive tract. Cilia and flagella of higher organisms are structurally and functionally different from bacterial flagella. Both possess an internal axoneme comprised of a characteristic arrangement of nine peripheral doublet microtubules with a central pair of single microtubules (Figure 1^ ), and the basis of their movement is the relative sliding of the outer doublets driven by a pair of projecting arms consisting of the motor protein, dynein. Sliding is then converted into bending by additional links and spokes comprised of the large numbers of polypeptides identified in the organelle.
Crawling Cells: Amoebae, Leucocytes
A wide variety of cells move by means of crawling over substrates rather than swimming through their environ- ment. Since microscopes began to be used to look at cells, observers have been fascinated by the movements of free- living protozoans, such as the freshwater Amoeba proteus. Observations of A. proteus have shown that it advances over its substrate by extending large processes known as pseudopodia and that their direction of extension from the
Outer doublet microtubules
Nexin link
Outer dynein arm Radial spoke
Inner dynein arm
Central singlet microtubule Plasma membrane
Figure 1 Diagram of a transverse section of a cilium or a flagellum. The major elements of the internal axoneme are nine peripheral microtubule doublets surrounding a central pair of single microtubules. Pairs of dynein arms attached to one doublet interact with the adjacent doublet to produce relative sliding. This is then converted into ciliary or flagellar bending by other structures including nexin links and radial spokes.
2 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net