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Protozoa


by
Diana Lipscomb
George Washington University
The diverse assemblage of organisms that carry out all of their life functions within the confines of a single, complex eukaryotic (see glossary) cell is called protozoa. Paramecium, Euglena, and Amoeba are well-known examples of these major groups of organisms. Some protozoa are more closely related to animals, others to plants, and still others are relatively unique. Although it is not appropriate to group them together into a single taxonomic category, the research tools used to study any unicellular organism are usually the same, and the field of protozoology has been created to carry out this research. The unicellular photosynthetic protozoa are sometimes also called algae and are addressed elsewhere. This report considers the status of our knowledge of heterotrophic protozoa (protozoa that cannot produce their own food).

Free-living Protozoa

Protozoans are found in all moist habitats within the United States, but we know little about their specific geographic distribution. Because of their small size, production of resistant cysts, and ease of distribution from one place to another, many species appear to be cosmopolitan and may be collected in similar microhabitats worldwide (Cairns and Ruthven 1972). Other species may have relatively narrow limits to their distribution.
Marine ciliates inhabit interstices of sediment and beach sands, surfaces, deep sea and cold Antarctic environments, planktonic habitats, and the algal mats and detritus of estuaries and wetlands. Our actual knowledge of salinity, temperature, and oxygen requirements of marine protozoa is poor (although some groups, such as the foraminifera, are better studied than others), and even the broadest outlines of their biogeographic ranges are usually a mystery. In general, freshwater protozoan communities are similar to marine communities except the specialized interstitial fauna of the sand is largely missing. In freshwater habitats, the foraminifera and radiolaria common in marine environments are absent or low in numbers while testate amoebae exist in greater numbers. Relative abundance of species in the marine versus freshwater habitat is unknown.
Soil-dwelling protozoa have been documented from almost every type of soil and in every kind of environment from the peat-rich soil of bogs to the dry sands of deserts. In general, protozoa are found in greatest abundance near the soil surface, especially in the upper 15 cm (6 in), but occasional isolates can be obtained at depths of a meter (yard) or more. Protozoa do not constitute a major part of soil biomass, but in some highly productive regions such as forest litter, the protozoa are a significant food source for the microinvertebrates, with a biomass that may reach 20 g/m2 of soil surface area there.

Environmental Quality Indicators

Polluted waters often have a rich and characteristic protozoan fauna. The relative abundance and diversity of protozoa are used as indicators of organic and toxic pollution (Cairns et al. 1972; Foissner 1987; Niederlehner et al. 1990; Curds 1992). Bick (1972), for example, provided a guide to ciliates that are useful as indicators of environmental quality of European freshwater systems, along with their ecological distribution with respect to parameters such as amount of organic material and oxygen levels. Foissner (1988) clarified the taxonomy of European ciliates as part of a system for classifying the state of aquatic habitats according to their faunas.

Symbiotic Protozoa

Parasites
Protozoa are infamous for their role in causing disease, and parasitic species are among the best-known protozoa. Nevertheless, our knowledge has large gaps, especially of normally free-living protozoa that may become pathogenic in immuno-compromised individuals. For example, microsporidia comprise a unique group of obligate, intracellular parasitic protozoa. Microsporidia are amazingly diverse organisms with more than 700 species and 80 genera that are capable of infecting a variety of plant, animal, and even other protist hosts. They are found worldwide and have the ability to thrive in many ecological conditions. Until the past few years, their ubiquity did not cause a threat to human health, and few systematists worked to describe and classify the species. Since 1985, however, physicians have documented an unusual rise in worldwide infections in AIDS patients caused by four different genera of microsporidia (Encephalitozoon, Nosema, Pleistophora, and Enterocytozoon). According to the Centers for Disease Control in the United States, difficulties in identifying microsporidian species are impeding diagnosis and effective treatment of AIDS patients.
Protozoan Reservoirs of Disease
The presence of bacteria in the cytoplasm of protozoa is well known whereas that of viruses is less frequently reported. Most of these reports simply record the presence of bacteria or viruses and assume some sort of symbiotic relationship between them and the protozoa. Recently, however, certain human pathogens were shown to not only survive but also to multiply in the cytoplasm of free-living, nonpathogenic protozoa. Indeed, it is now believed that protozoa are the natural habitat for certain pathogenic bacteria. To date, the main focus of attention has been on the bacterium Legionella pneumophila, the causative organism of Legionnaires' disease; these bacteria live and reproduce in the cytoplasm of some free-living amoebae (Curds 1992).
Symbionts
Some protozoa are harmless or even beneficial symbionts. A bewildering array of ciliates, for example, inhabit the rumen and reticulum of ruminates and the cecum and colon of equids. Little is known about the relationship of the ciliates to their host, but a few may aid the animal in digesting cellulose.

Data on Protozoa

Bibliography
While our knowledge of recent and fossil foraminifera in the U.S. coastal waterways is systematically growing, other free-living protozoa are poorly known. There are some regional guides and, while some are excellent, many are limited in scope, vague on specifics, or difficult to use. Largely because of these problems, most ecologists who include protozoa in their studies of aquatic habitats do not identify them, even if they do count and measure them for biomass estimates (Taylor and Sanders 1991).
Parasitic protozoa of humans, domestic animals, and wildlife are better known although no attempt has been made to compile this information into a single source. Large gaps in our knowledge exist, especially for haemogregarines, microsporidians, and myxosporidians (see Kreier and Baker 1987).
Museum Specimens
For many plant and animal taxa, museums represent a massive information resource. This is not true for protozoa. In the United States, only the National Natural History Museum (Smithsonian Institution) has a reference collection preserved on microscope slides, but it does not have a protozoologist curator and cannot provide species' identification or verification services. The American Type Culture Collection has some protozoa in culture, but its collection includes relatively few kinds of protozoa.

Ecological Role of Protozoa

Although protozoa are frequently overlooked, they play an important role in many communities where they occupy a range of trophic levels. As predators upon unicellular or filamentous algae, bacteria, and microfungi, protozoa play a role both as herbivores and as consumers in the decomposer link of the food chain. As components of the micro- and meiofauna, protozoa are an important food source for microinvertebrates. Thus, the ecological role of protozoa in the transfer of bacterial and algal production to successive trophic levels is important.
Factors Affecting Growth and Distribution
Most free-living protozoa reproduce by cell division (exchange of genetic material is a separate process and is not involved in reproduction in protozoa). The relative importance for population growth of biotic versus chemical-physical components of the environment is difficult to ascertain from the existing survey data. Protozoa are found living actively in nutrient-poor to organically rich waters and in fresh water varying between 0°C (32°F) and 50°C (122°F). Nonetheless, it appears that rates of population growth increase when food is not constrained and temperature is increased (Lee and Fenchel 1972; Fenchel 1974; Montagnes et al. 1988).
Comparisons of oxygen consumption in various taxonomic groups show wide variation (Laybourn and Finlay 1976), with some aerobic forms able to function at extremely low oxygen tensions and to thereby avoid competition and predation. Many parasitic and a few free-living species are obligatory anaerobes (grow without atmospheric oxygen). Of the free-living forms, the best known are the plagiopylid ciliates that live in the anaerobic sulfide-rich sediments of marine wetlands (Fenchel et al. 1977). The importance of plagiopylids in recycling nutrients to aerobic zones of wetlands is potentially great.
Ecological Interactions
Because of the small size of protozoa, their short generation time, and (for some species) ease of maintaining them in the laboratory, ecologists have used protozoan populations and communities to investigate competition and predation. The result has been an extensive literature on a few species studied primarily under laboratory conditions. Few studies have been extended to natural habitats with the result that we know relatively little about most protozoa and their roles in natural communities. Intraspecific competition for common resources often results in cannibalism, sometimes with dramatic changes in morphology of the cannibals (Giese 1973). Field studies of interspecific competition are few and most evidence for such species interactions is indirect (Cairns and Yongue 1977).
For further information:
Diana Lipscomb
George Washington University
Department of Biological Sciences
Washington, DC 20052

References
Bick, H. 1972. Ciliated protozoa. An illustrated guide to the species used as biological indicators in freshwater biology. World Health Organization, Geneva. 198 pp.

Cairns, J., G.R. Lanza, and B.C. Parker. 1972. Pollution related structural and functional changes in aquatic communities with emphasis on freshwater algae and protozoa. Proceedings of the National Academy of Sciences 124:79-127.

Cairns, J., and J.A. Ruthven. 1972. A test of the cosmopolitan distribution of fresh-water protozoans. Hydrobiologia 39:405-427.

Cairns, J., and W.H. Yongue. 1977. Factors affecting the number of species of freshwater protozoan communities. Pages 257-303 in J. Cairns, ed. Aquatic microbial communities. Garland, New York.

Curds, C.R. 1992. Protozoa and the water industry. Cambridge University Press, MA. 122 pp.

Fenchel, T. 1974. Intrinsic rate increase: the relationship with body size. Oecologia 14:317-326.

Fenchel, T., T. Perry, and A. Thane. 1977. Anaerobiosis and symbiosis with bacteria in free-living ciliates. Journal of Protozoology 24:154-163.

Foissner, W. 1987. Soil protozoa: fundamental problems, ecological significance, adaptations in ciliates and testaceans, bioindicators, and guide to the literature. Progress in Protistology 2:69-212.

Foissner, W. 1988. Taxonomic and nomenclatural revision of Stádecek's list of ciliates (Protozoa: Ciliophora) as indicators of water quality. Hydrobiologia 166:1-64.

Giese, A.C. 1973. Blepharisma. Stanford University Press, CA. 366 pp.

Kreier, J.P., and J.R. Baker. 1987. Parasitic protozoa. Allen and Unwin, Boston, MA. 241 pp.

Laybourn, J., and B.J. Finlay. 1976. Respiratory energy losses related to cell weight and temperature in ciliated protozoa. Oecologia 44:165-174.

Lee, C.C., and T. Fenchel. 1972. Studies on ciliates associated with sea ice from Antarctica. II. Temperature responses and tolerances in ciliates from Antarctica, temperate and tropical habitats. Archive für Protistenkunde 114:237-244.

Montagnes, D.J.S., D.H. Lynn, J.C. Roff, and W.D. Taylor. 1988. The annual cycle of heterotrophic planktonic ciliates in the waters surrounding the Isles of Shoals, Gulf of Maine: an assessment of their trophic role. Marine Biology 99:21-30.

Niederlehner, B.R., K.W. Pontasch, J.R. Pratt, and J. Cairns. 1990. Field evaluation of predictions of environmental effects from multispecies microcosm toxicity test. Archives of Environmental Contamination and Toxicology 19:62-71.

Taylor, W., and R. Sanders. 1991. Protozoa. Pages 37-93 in J.H. Thorp and A.P. Covich, eds. Ecology and classification of North American freshwater invertebrates. Academic Press, New York.



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