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Anatomy                                                                      91

                 these fibrils would act as channels for the extruded slime to flow along. Therefore, if the fibrils are
                 helically arranged, the cell will rotate as it glides; if the fibrils are aligned radially, the cell will
                 not rotate. In all species studied to date, this correlation is consistent, and provides a structural
                 explanation for why some species rotate as they glide while others do not.
                     In diatoms, motility is restricted to pennate species possessing a raphe. These diatoms display a
                 characteristic jerky movement forward or backward, with specie-specific path patterns. The general
                 velocity of their movement is 1–25 mm sec 21 , but they can accelerate up to 100–200 mm sec 21 .
                 Raphid diatoms possess an actin-based cytoskeletal system located just beneath the plasma
                 membrane at the raphe. Transmembrane components with an adhesive extracellular domain are
                 connected to these actin bundles, and their interaction is somehow involved in both adhesion
                 and motility mechanisms. Microtubules are also present in this region; in addition secretory ves-
                 icles containing polysaccharides often appear near the actin filaments at the raphe, providing the
                 mucilage strands that project from the raphe and adhere to the substratum during the gliding
                 process.
                     At least two models exist which provide reasonable explanation for diatom locomotion. In the
                 first model, a force applied to the transmembrane protein-actin connectors, parallel to the actin
                 bundles, would result in the movement of trasmembrane proteins through the cell and subsequent
                 movement of the cell in the opposite direction to the force. In the second model, the energy required
                 for motility would be generated by a conformational change of the adhesive mucilage on hydration
                 that occurs when it is secreted from the raphe. In this model, the actin bundles restrict the secretion
                 of mucilage to one end of the raphe, which generates a net force moving the cell over the site of
                 secretion. In both models, the secreted mucilage plays a central role either by providing traction
                 to translate the force into cell movement or by generating the energy through conformational
                 changes on hydration.
                     A slow gliding movement over solid substrata has been observed in Porphyridium sp.
                 (Rhodophyta) and in some desmids (Chlorophyta). In Porphyridium, the mucilage produced in
                 mucilage sacs located inside the cell is excreted through the membrane. In desmids mucilage is
                 excreted through the cell wall by flask-shaped pores. As they move, these gliding cells leave
                 behind a fibrillar mucilaginous trail, whose swelling by water pushes the cells forward.
                 Table 2.1 presents swimming and gliding speeds of some planktonic algae.





                                 TABLE 2.1
                                 Swimming and Gliding Speeds of Some Planktonic Algae
                                 Name                                  Mean Speed
                                 Gymnodinium gracilentum                500 mm sec 21
                                 Symbiodinium sp.                       250 mm sec 21
                                 Tetraselmis suecica                    180 mm sec 21
                                 Chattonella sp.                        120 mm sec 21
                                 Cryptomonad                            110 mm sec 21
                                 Chlorarachnion reptans                 110 mm sec 21
                                 Euglena gracilis                       100 mm sec 21
                                 Dunaliella salina                      95 mm sec 21
                                 Ochromonas danica                      80 mm sec 21
                                 Bigelowiella natans                    70 mm sec 21
                                 Pavlova salina                         50 mm sec 21
                                 Oscillatoria spp.                      10 mm sec 21
                                 Leptolyngbya spp.                    0.004 mm sec 21
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