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Protista

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Protista

Protist
Temporal range: Neoproterozoic – Recent
Scientific classification
Domain: Eukarya
Excluded groups

Many others;
classification varies

Protists /ˈprtɨst/ are a diverse group of eukaryotic microorganisms. Historically, protists were treated as a biological kingdom formally called the Protista and included mostly unicellular organisms that did not fit into the other kingdoms. Molecular information has been used to redefine this group in modern taxonomy as diverse and often distantly related phyla. The group of protists is now considered to mean diverse phyla that are not closely related through evolution and have different life cycles, trophic levels, modes of locomotion and cellular structures.[1][2] Besides their relatively simple levels of organization, the protists do not have much in common.[3] They are unicellular, or they are multicellular without specialized tissues, and this simple cellular organization distinguishes the protists from other eukaryotes, such as fungi, animals and plants.

The term protista was first used by Ernst Haeckel in 1866. Protists were traditionally subdivided into several groups based on similarities to the "higher" kingdoms: the unicellular "animal-like" protozoa, the "plant-like" protophyta (mostly unicellular algae), and the "fungus-like" slime molds and water molds. These traditional subdivisions, largely based on superficial commonalities, have been replaced by classifications based on phylogenetics (evolutionary relatedness among organisms). However, the older terms are still used as informal names to describe the morphology and ecology of various protists.

Protists live in almost any environment that contains liquid water. Many protists, such as the algae, are photosynthetic and are vital primary producers in ecosystems, particularly in the ocean as part of the plankton. Other protists, such as the kinetoplastids and apicomplexans, are responsible for a range of serious human diseases, such as malaria and sleeping sickness.

Classification

Historical classifications

The first division of the protists from other organisms came in the 1830s, when the German biologist Georg August Goldfuss introduced the word protozoa to refer to organisms such as ciliates and corals.[4] This group was expanded in 1845 to include all unicellular animals, such as foraminifera and amoebae. The formal taxonomic category Protoctista was first proposed in the early 1860s by John Hogg, who argued that the protists should include what he saw as primitive unicellular forms of both plants and animals. He defined the Protoctista as a "fourth kingdom of nature", in addition to the then-traditional kingdoms of plants, animals and minerals.[4] The kingdom of minerals was later removed from taxonomy by Ernst Haeckel, leaving plants, animals, and the protists as a “kingdom of primitive forms”.[5]

Herbert Copeland resurrected Hogg's label almost a century later, arguing that "Protoctista" literally meant "first established beings", Copeland complained that Haeckel's term protista included anucleated microbes such as bacteria. Copeland's use of the term protoctista did not. In contrast, Copeland's term included nucleated eukaryotes such as diatoms, green algae and fungi.[6] This classification was the basis for Whittaker's later definition of Fungi, Animalia, Plantae and Protista as the four kingdoms of life.[7] The kingdom Protista was later modified to separate prokaryotes into the separate kingdom of Monera, leaving the protists as a group of eukaryotic microorganisms.[8] These five kingdoms remained the accepted classification until the development of molecular phylogenetics in the late 20th century, when it became apparent that neither protists nor monera were single groups of related organisms (they were not monophyletic groups).[9]

Modern classifications

Although systematists today do not treat protists as a formal taxon, the term protist is currently used under two ways. The most popular contemporary definition is a phylogenetic one that identifies a paraphyletic group: protists are anything that is a eukaryote but is not an animal, (land) plant, or (true) fungus. The other definition describe protists primarily by functional or biological criteria: protists are essentially those eukaryotes that are never multicellular,[10] that either exist as independent cells, or if they occur in colonies, do not show differentiation into tissues.[11] The term protozoa is used to refer to heterotrophic species of protists that do not form filaments. These terms are not used in current taxonomy, and are retained only as convenient ways to refer to these organisms.

The taxonomy of protists is still changing. Newer classifications attempt to present monophyletic groups based on ultrastructure, biochemistry, and genetics. Because the protists as a whole are paraphyletic, such systems often split up or abandon the kingdom, instead treating the protist groups as separate lines of eukaryotes. The recent scheme by Adl et al. (2005)[11] is an example that does not bother with formal ranks (phylum, class, etc.) and instead lists organisms in hierarchical lists. This is intended to make the classification more stable in the long term and easier to update. Some of the main groups of protists, which may be treated as phyla, are listed in the taxobox at right.[12] Many are thought to be monophyletic, though there is still uncertainty. For instance, the excavates are probably not monophyletic and the chromalveolates are probably only monophyletic if the haptophytes and cryptomonads are excluded.[13]

Metabolism

Nutrition in some different types of protists is variable. In flagellates, for example, filter feeding may sometimes occur where the flagella find the prey. Other protists can engulf bacteria and digest them internally, by extending their cell membrane around the food material to form a food vacuole. This is then taken into the cell via endocytosis (usually phagocytosis; sometimes pinocytosis).

Nutritional types in protist metabolism
Nutritional type Source of energy Source of carbon Examples
 Phototrophs   Sunlight   Organic compounds or carbon fixation  Algae, Dinoflagellates or Euglena 
 Organotrophs  Organic compounds   Organic compounds   Apicomplexa, Trypanosomes or Amoebae 

Reproduction

Some protists reproduce sexually (gametes), while others reproduce asexually (binary fission).

Some species, for example Plasmodium falciparum, have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually.[14] However, it is unclear how frequently sexual reproduction causes genetic exchange between different strains of Plasmodium in nature and most populations of parasitic protists may be clonal lines that rarely exchange genes with other members of their species.[15]

Sexual reproduction

Eukaryotes emerged in evolution more than 1.5 billion years ago.[16] The earliest eukaryotes were likely protists. Although sexual reproduction is widespread among extant eukaryotes, it seemed unlikely until recently, that sex could be a primordial and fundamental characteristic of eukaryotes. A principal reason for this view was that sex appeared to be lacking in certain pathogenic protists whose ancestors branched off early from the eukaryotic family tree. However, several of these protists are now known to be capable of, or to recently have had the capability for, meiosis and hence sexual reproduction. For example, the common intestinal parasite Giardia lamblia was once considered to be a descendant of a protist lineage that predated the emergence of meiosis and sex. However, G. lamblia was recently found to have a core set of genes that function in meiosis and that are widely present among sexual eukaryotes.[17] These results suggested that G. lamblia is capable of meiosis and thus sexual reproduction. Furthermore, direct evidence for meiotic recombination, indicative of sex, was also found in G. lamblia.[18]

Parasitic protozoa of the genus Leishmania cause human diseases. These protists have recently been shown to have a sexual cycle as well.[19]

Trichomonas vaginalis, a parasitic protist, is not known to undergo meiosis, but when Malik et al.[20] tested for 29 genes that function in meiosis, they found 27 to be present, including 8 of 9 genes specific to meiosis in model eukaryotes. These findings suggest that T. vaginalis may be capable of meiosis. Since 21 of the 29 meiotic genes were also present in G. lamblia, it appears that most of these meiotic genes were likely present in a common ancestor of T. vaginalis and G. lamblia. These two species are descendants of protist lineages that are highly divergent among eukaryotes, leading Malik et al.[20] to suggest that these meiotic genes were likely present in a common ancestor of all eukaryotes.

Based on a phylogenetic analysis, Dacks and Roger proposed that facultative sex was present in the common ancestor of all eukaryotes.[21]

This view was further supported by a study of amoebae by Lahr et al.[22] Amoeba have generally been regarded as asexual protists. However these authors describe evidence that most amoeboid lineages are anciently sexual, and that the majority of asexual groups likely arose recently and independently.

Protists generally reproduce asexually under favorable environmental conditions, but tend to reproduce sexually under stressful conditions, such as starvation or heat shock.[23] Oxidative stress, which is associated with the production of reactive oxygen species leading to DNA damage, also appears to be an important factor in the induction of sex in protists.[23]

Role as pathogens

There are some protists that are significant pathogens of animals and others that are pathogens of plants; for example there are five species of the parasitic genus Plasmodium, which cause malaria in humans; and the oomycete Phytophthora infestans, which causes late blight in potatoes.[24] A more thorough understanding of protist biology may allow these diseases to be treated more efficiently.

Recent papers have proposed the use of viruses to treat infections caused by protozoa.[25][26]

Researchers from the Agricultural Research Service are taking advantage of protists as pathogens in an effort to control red imported fire ant (Solenopsis invicta) populations in Argentina. With the help of spore-producing protists such as Kneallhazia solenopsae (this is more widely recognized as belonging to the fungus kingdom now) the red fire ant populations can be reduced by 53-100%.[27] Researchers have also found a way to infect phorid flies with the protist without harming the flies. This is important because the flies act as a vector to infect the red fire ant population with the pathogenic protist.[28]

See also

References

Bibliography

General

  • Haeckel, E. Das Protistenreich. Leipzig, 1878.
  • Hausmann, K., N. Hulsmann, R. Radek. Protistology. Schweizerbart'sche Verlagsbuchshandlung, Stuttgart, 2003.
  • Margulis, L., J.O. Corliss, M. Melkonian, D.J. Chapman. Handbook of Protoctista. Jones and Bartlett Publishers, Boston, 1990.
  • Margulis, L., K.V. Schwartz. Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth, 3rd ed. New York: W.H. Freeman, 1998.
  • Margulis, L., L. Olendzenski, H.I. McKhann. Illustrated Glossary of the Protoctista, 1993.
  • Margulis, L., M.J. Chapman. Kingdoms and Domains: An Illustrated Guide to the Phyla of Life on Earth. Amsterdam: Academic Press/Elsevier, 2009.
  • Schaechter, M. Eukaryotic microbes. Amsterdam, Academic Press, 2012.

Physiology and ecology

  • Foissner, W.; D.L. Hawksworth. Protist Diversity and Geographical Distribution. Dordrecht: Springer, 2009
  • Fontaneto, D. Biogeography of Microscopic Organisms. Is Everything Small Everywhere? Cambridge University Press, Cambridge, 2011.
  • Levandowsky, M. Physiological Adaptations of Protists. In: Cell physiology sourcebook : essentials of membrane biophysics. Amsterdam; Boston: Elsevier/AP, 2012.

External links

  • Tree of Life: Eukaryotes
  • A java applet for exploring the new higher level classification of eukaryotes
  • Plankton Chronicles - Protists - Cells in the Sea - video
  • Database of protist images
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