- Key People:
- Herbert Spencer Jennings
- Related Topics:
- protozoan
- algae
- slime mold
- dinoflagellate
- ciliate
Cell division in protists, as in plant and animal cells, is not a simple process, although it may superficially appear to be so. The typical mode of reproduction in most of the major protistan taxa is asexual binary fission. The body of an individual protist is simply pinched into two parts or halves; the “parental” body disappears and is replaced by a pair of offspring or daughter nuclei, although the latter may need to mature somewhat to be recognizable as members of the parental species. The length of time for completion of the process of binary fission varies among groups of organisms and with environmental conditions; generally it ranges from just a few hours in an optimal situation to many days under other circumstances. In some unicellular algal protists, reproduction occurs by fragmentation. Mitotic replications of the nuclear material presumably accompany or precede all divisions of the cytoplasm (cytokinesis) in protists.
Multiple fission also occurs among protists and is common in some parasitic species. The nucleus divides repeatedly to produce a number of daughter nuclei, which eventually become the nuclei of the progeny after repeated cellular divisions. There are several kinds of multiple fission, often correlated with phases or stages in the full life cycle of a given species. The number of offspring or filial products resulting from a multiple division (or very rapid succession of binary fissions) may vary from four to dozens or even hundreds, generally in a short period of time. Modes of such multiple fission range from budding, in which a daughter nucleus is produced and split from the parent together with some of the surrounding cytoplasm, to sporogony (production of sporozoites by repeated divisions of a zygote) and schizogony (formation of multiple merozoites, as in malarial parasites). The latter two phenomena are characteristic of many protists that are obligate parasites of more advanced eukaryotes. Some multicellular algal protists reproduce via asexual spores, structures that are themselves often produced by a series of rapid fissions.
Even under a light microscope, differences can be seen in the modes of division among diverse groups of protists. The flagellates, for example, exhibit a longitudinal, or mirror-image, type of fission (symmetrogenic fission). The ciliates, on the other hand, basically divide in a point-by-point correspondence of parts (homothetogenic fission), often seen as essentially transverse or perkinetal (across the kineties, or ciliary rows). Many amoebas exhibit, in effect, no clear-cut body symmetry or polarity, and thus their fission is basically simpler and falls into neither of the categories described above.
Sexual phenomena are known among the protists. The erroneous view that practically all protists reproduce asexually is explained by the fact that certain well-known organisms, such as species belonging to the genus Euglena, do not demonstrate sexuality. Even many of the unicellular species can, under appropriate conditions, form gametes (sex cells), which fuse and give rise to a new, genetically unique generation. In fact, sexual reproduction—the union of two gametes (syngamy)—is the most common sexual phenomenon and occurs quite widely among the protists—for example, among various flagellated organisms and pseudopods and among many parasitic phyla (e.g., in Plasmodium, a malaria-causing organism).
Conjugation, the second major kind of sexual phenomenon and one occurring in the ciliated protists, has genetic and evolutionary results identical to those of syngamy. The process involves the fusion of gametic nuclei rather than independent gamete cells. A zygotic, or fusion, nucleus, not a true zygote, is produced and undergoes a series of meiotic divisions to produce a number of haploid pronuclei; all but one of these pronuclei in each organism will disintegrate. The remaining pronuclei divide mitotically; one pronucleus from each organism is exchanged, and the new micronuclei and macronuclei of the next generation are formed. Following the exchange of the pronuclei and the subsequent formation of new micronuclei and macronuclei in each organism, a series of asexual fissions, accompanied by mitotic divisions of the new diploid micronuclei, occurs in each exconjugant line. The new polyploid macronuclei are distributed passively in the first of these divisions; in subsequent fission, the macronuclei duplicate themselves through a form of mitosis. This last stage constitutes the only reproduction involved in the process.
Conjugation, as described here, is essentially limited to the ciliates, and there is considerable variation in the manner in which it is exhibited among them. For example, the two ciliates themselves may be of noticeably different size (called macroconjugants and microconjugants), or the number of predivisions of the micronuclei may vary, as may the number of nuclear divisions that take place after the zygotic nucleus is formed. Furthermore, chemical signals (gamones) are given or exchanged before a pair of protists unite in conjugation. It is not known if these gamones should be considered as sex pheromones, reminiscent of those known in many animals (for example, certain insects), but they seem to serve the similar purpose of attracting or bringing together different mating types.
While conjugation may be considered a process of reciprocal fertilization, a parallel sexual phenomenon in ciliates, which takes place in single, unpaired individuals, may be considered a process of self-fertilization. In this type of fertilization, called autogamy, complete homozygosity is obtained in the lines derived from the single parent.
Protist life cycles range from relatively simple ones that may involve only periodic binary fissions to very complex schemes that may contain asexual and sexual phases, encystment and excystment, and—in the case of many symbiotic and parasitic forms—an alternation of hosts. In the more complicated life cycles in particular, the morphology of the organism may be strikingly different (polymorphism) from phase to phase in the entire life cycle. Among certain ciliate groups in which a larval or migratory form (known as a swarmer) is produced by the parent, the offspring may demonstrate remarkably different morphology.
Dormant stages in a life cycle are probably more common in algal protists than in protozoan protists. Such stages, somewhat analogous to hibernation in mammals, serve to preserve the species during unfavourable conditions, as in times of inadequate food supply or extreme temperatures. The occurrence of resistant cysts in the vegetative stage depends, therefore, on such environmental factors as season, temperature, light, water, and nutrient supply. The fertilized egg, or zygote, in a number of algal groups may also pass into a dormant stage (a zygospore). Temporary or long-lasting cysts may occur among other protist species as well. Many sporozoa and members of other totally parasitic phyla form a highly resistant stage—for example, the oocyst of the coccidian parasites, which may survive for a long time in the fecal material of the host or in the soil. This cyst is the infective stage for the next host in the parasite’s life cycle.
Some life cycles involve not only multiple hosts but also a vector—a particular metazoan organism that can act as either an active or a passive carrier of the parasite to the next host. In malaria, for example, a mosquito is required to transfer the Plasmodium species to the next vertebrate host.
Ecology
The distribution of protists is worldwide; as a group, these organisms are both cosmopolitan and ubiquitous. Every individual species, however, has preferred niches and microhabitats, and all protists are to some degree sensitive to changes in their surroundings. The availability of sufficient nutrients and water, as well as sunlight for photosynthetic forms, is, however, the only major factor restraining successful and heavy protist colonization of practically any habitat on Earth.
Free-living forms are particularly abundant in natural aquatic systems, such as ponds, streams, rivers, lakes, bays, seas, and oceans. Certain of these forms may occur at specific levels in the water column, or they may be bottom-dwellers (benthic). More specialized, sometimes human-made, habitats are also often well populated by both pigmented and nonpigmented protists. Such sites include thermal springs, briny pools, cave waters, snow and ice, beach sands and intertidal mud flats, bogs and marshes, swimming pools, and sewage treatment plants. Many are commonly found in various terrestrial habitats, such as soils, forest litter, desert sands, and the bark and leaves of trees. Cysts and spores may be recovered from considerable heights in the atmosphere.
Fossilized forms are plentiful in the geologic record. Fossils of unicellular organisms have been found in strata dated to about 1.9 billion years ago, during the Precambrian. Many lineages of protists have left no record of their now extinct forms, however, making speculation about early phylogenetic and evolutionary relationships with other eukaryotes difficult to verify.
Symbiotic protists are as widespread as free-living forms, since they occur everywhere their hosts are to be found. Hundreds or even thousands of kinds of protists live as ectosymbionts or episymbionts, finding suitable niches with plants, fungi, vertebrate and invertebrate animals, or even other protists. Seldom are the hosts harmed; in fact, these often mobile substrates are actually used as a means of dispersal.
Endosymbionts include commensals, facultative parasites, and obligate parasites; the latter category embraces forms that have effects on their hosts ranging from mild discomfort to death. Protozoan and certainly nonphotosynthetic protists are implicated far more often in such associations than are algal forms. In a few protists, both cytoplasm and nuclei can be invaded by other protists, and intimate, mutually beneficial relationships between protistan hosts and protistan symbionts have been seen, such as foraminiferans or ciliates that nourish symbiotic algae in their cytoplasm. When higher eukaryotes are hosts to protists, all body cavities and organ systems are susceptible to invasion, although terrestrial plants bear relatively few such parasites. In animal hosts, the three principal areas serving as sites for endosymbiotic species are the coelom, the digestive tract and its associated organs, and the circulatory system.
The numbers of individuals in populations of many protists reach staggering figures. There are, on the average, tens of thousands of protists in a gram of arable soil, hundreds of thousands in the gut of a termite, millions in the rumen of a bovine mammal, billions in a tiny patch of floating plankton in the sea, and trillions in the bloodstream of a person infected with severe malaria. Some severe diseases of humans are caused by protists, primarily blood parasites. Malaria, trypanosomiasis (e.g., African sleeping sickness), leishmaniasis, toxoplasmosis, and amoebic dysentery are debilitating or fatal afflictions.
Protist parasites infecting domesticated livestock, poultry, hatchery fishes, and other such food sources deplete supplies or render them unpalatable. The economic losses can be considerable. Certain free-living marine dinoflagellates are the causative agents of the so-called red tide outbreaks that occur periodically along coasts throughout the world; a toxin released by the blooming protists kills fishes in the affected area. Other dinoflagellates produce a toxin that may be taken up by certain shellfish (bivalve mollusks) and that causes shellfish poisoning, characterized in severe cases by respiratory paralysis and death, when the mollusk is eaten by humans. Some of the “lower” fungal protists have had significant effects on human history. One species was responsible for the great Irish potato famine of the mid-19th century, and later, another nearly ruined the entire French wine industry before a fungicide was developed to destroy it.
Many protists provide humans with benefits, some more obvious than others. Because protists are located near the bottom of the food chain in nature (just above the bacteria), they serve a crucial role in sustaining the higher eukaryotes in fresh and marine waters. In addition to directly and indirectly supplying organic molecules (such as sugars) for other organisms, the pigmented (chlorophyll-containing) algal protists produce oxygen as a by-product of photosynthesis. Algae may supply up to half of the net global oxygen. Deposits of natural gas and crude oil are derived from fossilized populations of algal protists. Much of the nutrient turnover and mineral recycling in the oceans and seas comes from the activities of the heterotrophic (nonpigmented) flagellates and the ciliates living there, species that feed on the bacteria and other primary producers present in the same milieu. Seaweeds (e.g., brown algae) have long been used as fertilizers.
The calcareous test, or shell, of the foraminiferans is preservable and constitutes a major component of limestone rocks. Assemblages of certain of these protists, which are abundant and usually easily recognized, are known to have been deposited during various specific periods in Earth’s geologic history. Geologists in the petroleum industry study foraminiferan species present in samples of drilled cores in order to determine the age of different strata in Earth’s crust, thus making possible the identification of rich oil deposits. Before synthetic substitutes, blackboard chalk consisted mostly of calcium carbonate derived from the scales (coccoliths) of certain algal protists and from the tests of foraminiferans. Diatoms and some ciliate species are useful as indicators of water quality and therefore of the amount of pollution in natural aquatic systems and in sewage purification plants. Selected species of parasitic protozoans may play a significant role as biological control organisms against certain insect predators of food plants.
Protists have been used as model cells in laboratory research, some of which is directed against major human diseases. The combination of characteristics that has made them superior to both prokaryotic cells and other eukaryotic cells includes their easy availability and maintenance, convenient size for handling in large numbers, short generation time, broad physiological adaptability, basic structural and functional similarity to the eukaryotic cells of animal organisms, and, most importantly for sophisticated work requiring purity of material and rigidity of controls, culturability (i.e., their successful growth axenically—free of other living organisms—and on chemically definable media). The culturability of some unicellular free-living protists has made them invaluable as assay organisms and pharmacological tools. Among those that have proved to be useful this way, one of the most important is the ciliate Tetrahymena, which serves as a model cell in investigations in cell and molecular biology. The value of such work in areas such as biomedical and cancer research is potentially great.
