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Kinesin- and Dynein-Powered Movements

Within cells, proteins, organelles, and other membrane-limited vesicles, organelles, and proteins are frequently transported distances of many micrometers along well-defined routes in the cytosol and delivered to particular addresses. Diffusion alone cannot account for the rate, directionality, and destinations of such transport processes. Findings from early experiments with fish-scale pigment cells and nerve cells first demonstrated that microtubules function as tracks in the intracellular transport of various types of "cargo. " Eventually, two families of motor proteins—kinesins and dyneins— were found to mediate transport along microtubules.

A second type of movement that depends on microtubule motor proteins is the beating of cilia and flagella. Huge numbers of cilia (more than 107/mm2) cover the surfaces of mammalian respiratory passages where their beating dislodges and expels particulate matter that collects in the mucus secretions of these tissues. In the oviduct, cilia help transport eggs down the fallopian tube. In contrast, sperm cells and many unicel-

► EXPERIMENTAL FIGURE 20-17 The rate of axonal transport in vivo can be determined by radiolabeling and gel electrophoresis. The cell bodies of neurons in the sciatic nerve are located in dorsal-root ganglia. Radioactive amino acids injected into these ganglia in experimental animals are incorporated into newly synthesized proteins, which are then transported down the axon to the synapse. Animals are sacrificed at various times after injection and the dissected sciatic nerve is cut into small segments for analysis with the use of gel electrophoresis. The red, blue, and purple dots represent groups of proteins that are transported down the axon at different rates, red most rapidly, purple least rapidly.

lular organisms have a single flagellum, which propels the cells forward at velocities approaching 1 mm/s.

In this section, we first consider the transport of materials in axons. Studies of such axonal transport, a process first discovered more than 50 years ago, have contributed greatly to our understanding of microtubule-associated intracellular transport. We then consider the structure and function of the microtubule motor proteins. A description of the unique mi-crotubule-based structures and motor proteins responsible for the movement of cilia and flagella concludes this section.

Axonal Transport Along Microtubules Is in Both Directions

A neuron must constantly supply new materials—proteins and membranes—to an axon terminal to replenish those lost in the exocytosis of neurotransmitters at the junction (synapse) with another cell. Because proteins and membranes are synthesized only in the cell body, these materials must be transported down the axon, which can be as much as a meter in length, to the synaptic region. This movement of materials is accomplished on microtubules, which are all oriented with their (+) ends toward the terminal (see Figure 20-14c).

The results of classic pulse-chase experiments in which radioactive precursors are microinjected into the dorsal-root ganglia near the spinal cord and then tracked along their nerve axons showed that axonal transport is in both directions. Anterograde transport proceeds from the cell body to

Microtubule

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