ECOLOGICAL CONSIDERATIONS OF DIATOM CELL MOTILITY. I. CHARACTERIZATION OF MOTILITY AND ADHESION IN FOUR DIATOM SPECIES1

To better determine the ecological role of motility in pennate diatoms, we quantitatively characterized several motility and adhesion properties of four species of motile pennate diatoms (Craticula sp., Pinnularia sp., Nitzschia sp., and Stauroneis sp.) isolated from the same freshwater pond. Using computer-assisted video microscopy, we measured speed, size/shape, functional adhesion, path curvature, and light sensitivity for these species, each of which shows a distinctive set of motile behaviors. The average speeds of Stauroneis, Pinnularia, Nitzschia, and Craticula cells are 4.6, 5.3, 10.4, and 10.0 μm · s^?1, respectively. Craticula and Nitzschia cells move in a relatively straight path (<4 degrees rotation per 100 μm movement), Stauroneis exhibits minor rotation (about 7 degrees per 100 μm movement), and Pinnularia rotates considerably during movement (about 22 degrees per 100 μm moved). Functional adhesion (as measured by the release rate of attached cells from the underside of an inverted coverslip) shows a half time for cell release of approximately 50 min for Craticula, 192 min for Pinnularia, and >1 day for Nitzschia and Stauroneis. Direction reversal at light/dark boundaries, which appears to be the main contributor to diatom Phototaxis, is most responsive for Craticula, Pinnularia, and Nitzschia at wavelengths around 500 nm. Craticula and Nitzschia cells are the most sensitive in the photophobic response, with over 60% of these cells responding to a 30-1x light/dark boundary at 500 nm, whereas Pinnularia cells are only moderately responsive at this irradiance, showing a maximal response of approximately 30% of cells at 450 nm. Stauroneis cells, in contrast, had a maximal photosensitive response at 700 nm, suggesting that this cell type may use a different response mechanism than the other three cell types. In addition, Craticula and Pinnularia show a net movement out of the light spot when illuminated at 650 nm, whereas Stauroneis shows a net movement out of the light spot when illuminated at 450 nm. Such quantitative characterizations of species-specific responses to environmental stimuli should give us a firm foundation for future studies analyzing the behavior of interspecies diatom competition for limited light or nutrient resources.     

ACTIVE GLIDING MOTILITY IN AN ARAPHID MARINE DIATOM,ARDISSONEA (FORMERLY SYNEDRA) CRYSTALLINA1

Active gliding movement over long distances was observed and filmed in the marine pennate diatom Ardissonea (Synedra) crystallina (Agardh) Kütz. Typical speeds measured ca. 1–2 μm-s^?1. Motion wax often smooth and steady; however, discontinuous jerky motions and rolling movements were common. Motion, was associated with secretion of twin or, less commonly, single straight trails of mucilage from one end of the cell. In a few instances, reversal in direction was related to cessation of mucilage secretion at one end and commencement at the other. Temporary cessation of movement due to an obstruction was accompanied by a build-up of mucilage at one end of the cell. Mucilage was apparently secreted at two specific sites at each end of the cell and was stained by alcian blue. Persistent trails were visible under scanning electron microscopy (SEM). SEM confirmed that cells had no raphes or labiate processes. The apparent site of secretion was a deep groove formed at the junction of the valve and valvocopula (first girdle band) at each end of the cell. Transmission electron microscopy confirmed the presence of mucilage vesicles in the cytoplasm, but these were not in any manner obviously related to secretion nor was any morphological structure associated with secretion. Cells often become epiphytic through secretion of a terminal stipe. Both stipe secretion and movement may involve the same structural differentiation of the frustule. These results demonstrate a previously unrecorded type of diatom motility. The mechanism, involves mucilage secretion and appears similar to that seen, for example, in some other algae such as the desmids (green algae).     

RELATIONSHIPS OF SEDIMENTED DIATOM SPECIES (BACILLARIOPHYCEAE) TO ENVIRONMENTAL GRADIENTS IN DILUTE NORTHERN NEW ENGLAND LAKES1

Limnological gradients of small, oligotrophic, and low conductance lakes in northern New England were defined by principal components analysis; relationships of sedimented diatom species to the gradients were investigated by correlation analysis. Diatom distributions were most strongly related to the gradient of pH and alkalinity and the covarying variables, conductance, Mg, Ca, total Al, and exchangeable Al. Weaker relationships to lake morphology, dissolved organic carbon and water color, altitude and marine aerosol inputs, and the distinctive water chemistry of some New Hampshire lakes were also present. Results for 16 taxa of importance in our studies of lake acidity are given in detail and are compared to results from other regions of eastern North America. Planktonic taxa were absent below pH 5.5, with the exception of the long form of Asterionella ralfsii var. americana Korn. The two forms of this taxon differed ecologically: the long form (>45μm) had an abundance weighted mean (AWM) pH 4.90 and occurred mostly in lakes that were deep relative to transparency; the short form (<45μm)had an AWM pH and occurred on lakes that were shallow relative to transparency. The ecological advantage of a “splitter” approach to diatom taxonomy was demonstrated by examination of other taxa as well, including Tabellaria flocculosa (Roth) Kütz. These results have important implications for paleolimnological interpretations.     

INFLUENCE OF TRICHOME LENGTH ON LINEAR TRANSLATION IN OSCILLATORIA PRINCEPS (CYANOPHYCEAE)1

Motile trichomes of Oscillatoria princeps Voucher were examined to determine the relationship between trichome length and the ability to glide through hardened or viscous media. A minimum length for movement in solid media was found to be 0.1 mm in 0.5% agar (w/v).Viscous media tests revealed that coordinated movements through methyl cellulose media were not seen in trichomes 15–40 μ long at the viscosity employed. The minimum length motility increases with the viscosity of the medium and the gliding rates observed are greater in longer trihomes. These findings are discussed in relation to trichome surface area and the hypothesis that the motility system is restricted to lateral surfaces of the trichome.     

CHANGES IN CELL COMPOSITION AND LIPID METABOLISM MEDIATED BY SODIUM AND NITROGEN AVAILABILITY IN THE MARINE DIATOM PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

The effects of nitrogen starvation in the presence or absence of sodium in the culture medium were monitored in batch cultures of the marine diatom Phaeodactylum tricornutum Bohlin. During nitrogen starvation in the presence of sodium, cell nitrogen and chlorophyll a decreased, mainly as a consequence of continued cell division. These decreases were accompanied by decreases in the rates of photosynthesis and respiration. There was no change in either cell volume or carbohydrate, but both carbon and lipid increased. During nitrogen starvation in the absence of sodium, cell division ceased. Cell nitrogen and chlorophyll a remained constant, and respiration did not decrease, but the changes in the photosynthetic rate and the lipid content per cell were similar to cultures that were nitrogen-starved in the presence of sodium. The carbon-to-nitrogen ratio increased in both cultures. Nitrogen, in the form of nitrate, and sodium were resupplied to cultures that had been preconditioned in nitrogen- and sodium-deficient medium for 5 d. Control cultures to which neither nitrate or sodium were added remained in a static state with respect to cell number, volume, and carbohydrate but showed slight increases in lipid. Cells in cultures to which 10 mM nitrate alone was added showed a similar response to cultures where no additions were made. Cells in cultures to which 50 mM sodium alone was added divided for 2 d, with concomitant small decreases in all measured constituents. Cell division resumed in cultures to which both sodium and nitrate were added. The lipid content fell dramatically in these cells and was correlated to metabolic oxidation via measured increases in the activity of the glyoxylate cycle enzyme, isocitrate lyase. We conclude that lipids are stored as a function of decreased growth rate and are metabolized to a small extent when cell division resumes. However, much higher rates of metabolism occur if cell division resumes in the presence of a nitrogen source.     

INFECTION,GROWTH, AND COMMUNITY-LEVEL CONSEQUENCES OF A DIATOM PATHOGEN IN A SONORAN DESERT STREAM1

We describe effect of a pathogen that spread through a dense, rapidly growing, benthic diatom community during two infection periods (February and mid-April 1991) in Sycamore Creek, Arizona. Infected areas appeared as gray rings within a matrix of healthy diatom growth and spread rapidly, eventually covering all benthic substrata and causing algal sloughing (within 2 wk in February and 1 wk in April). Examination of algal material with transmission electron microscopy revealed the presence of invasive bacteria within diatom cells from infected areas, suggesting a pathogenic bacterium as the most probable cause of this phenomenon. Infected area supported lower chlorophyll a concentrations and contained higher percentages of diatom cells with fragmented or reduced chloroplasts than uninfected areas. Spread of the pathogen appeared to be linked most strongly with diatom densities. The infection spread most rapidly in April, when cell densities were highest, and decimated all diatom species populations. The February infection was more species-specific in its action, affecting large motile and rosetteforming taxa more strongly than small, adnate diatoms. This latter group likely resided at the base of communities and may have been buffered from pathogen transfer by mucilage and/or dentrital particles. Consequently, relative abundance of small, adnate diatom taxa increased in algal communities as a result of the February infection. Pathogen-induced alteration of diatom species composition and abundance should influence primary production in this ecosystem and affect the dynamics of organisms that exploit algae as a resource.     

BRACKISH WATER AND FRESHWATER SPECIES OF THE DIATOM GENUS SKELETONEMA. II. SKELETONEMA POTAMOS COMB. NOV.1

Electron microscope investigations of the siliceous frustule show that the diatom described by Hustedt as Stephanodiscus subsalsus (A. Cleve) Hust. is not Skeletonema subsalsum (A. Cleve) Bethge (Melosira subsalsa A. Cleve) but is Microsiphona potamos Weber. This species is so similar to Skeletonema costatum (Grev.) Cleve and Skeletonema subsalsum that the combination Skeletonema potamos (Weber) Hasle is suggested. Present records classify Skeletonema potamos as a freshwater species of lakes and rivers. In Sandusky Bay, Lake Erie (U.S.A.) and in River Wümme, a tributary of the River Weser (Germany) it grows with Skeletonema subsalsum. In nature, and when grown in cultures at a salinity of 0%, the processes are extremely short; when grown at salinities of 2% or more, the processes are much longer.