Protozoa (also protozoan, plural protozoans) is an informal term for a group of single-celled eukaryotes, either free-living or parasitic, which feed on organic matter such as other microorganisms or organic tissues and debris.Historically, protozoans were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae. Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to describe single-celled protists (that is, eukaryotes that aren't animals, plants, or fungi) that feed by heterotrophy.
In some systems of biological classification, Protozoa remains a high-level taxonomic group. When first introduced by Georg Goldfuss in 1818, Protozoa was erected as a class within the animals,and its etymology is literally "first animals". In later classification schemes it was elevated to a variety of higher ranks, including phylum, subkingdom and kingdom, and sometimes included within Protoctista or Protista. With the advent of techniques such as molecular phylogenetics, it was realized that Protozoa did not represent a natural group; but while it is not an accepted taxon in cladistic analyses, some systematists continue to use it as a formal taxon.
In a series of classifications proposed by Thomas Cavalier-Smith and his collaborators since 1981, Protozoa has been ranked as a kingdom.The seven-kingdom scheme presented by Ruggiero et al. in 2015, places eight phyla under Kingdom Protozoa: Euglenozoa, Amoebozoa, Metamonada, Choanozoa sensu Cavalier-Smith, Loukozoa, Percolozoa, Microsporidia and Sulcozoa. Notably, this kingdom excludes several major groups of organisms traditionally placed among the protozoa, including the ciliates, dinoflagellates, foraminifera, and the parasitic apicomplexans, all of which are classified under Kingdom Chromista. Kingdom Protozoa, as defined in this scheme, does not form a natural group or clade, but a paraphyletic group or evolutionary grade, within which the members of Fungi, Animalia and Chromista are thought to have evolved.
The word "protozoa" (singular protozoon or protozoan) was coined in 1818 by zoologist Georg August Goldfuss, as the Greek equivalent of the German Urthiere, meaning "primitive, or original animals" ( ur- ‘proto-’ + Thier ‘animal’). πρῶτος (prôtos), meaning "first", and ζῶα (zôa), plural of ζῶον (zôon), meaning "animal". The use of Protozoa as a formal taxon has been discouraged by some researchers, mainly because the term implies kinship with animals (Metazoa) and promotes an arbitrary separation of "animal-like" from "plant-like" organisms.Goldfuss created Protozoa as a class containing what he believed to be the simplest animals. Originally, the group included not only single-celled microorganisms but also some "lower" multicellular animals, such as rotifers, corals, sponges, jellyfish, bryozoa and polychaete worms. The term Protozoa is formed from the Greek words
In 1848, as a result of advancements in cell theory pioneered by Theodor Schwann and Matthias Schleiden, the anatomist and zoologist C. T. von Siebold proposed that the bodies of protozoans such as ciliates and amoebae consisted of single cells, similar to those from which the multicellular tissues of plants and animals were constructed. Von Siebold redefined Protozoa to include only such unicellular forms, to the exclusion of all metazoa (animals).At the same time, he raised the group to the level of a phylum containing two broad classes of microorganisms: Infusoria (mostly ciliates and flagellated algae), and Rhizopoda (amoeboid organisms). The definition of Protozoa as a phylum or sub-kingdom composed of "unicellular animals" was adopted by the zoologist Otto Bütschli—celebrated at his centenary as the "architect of protozoology" —and the term came into wide use.
As a phylum under Animalia, the Protozoa were firmly rooted in the old "two-kingdom" classification of life, according to which all living beings were classified as either animals or plants. As long as this scheme remained dominant, the protozoa were understood to be animals and studied in departments of Zoology, while photosynthetic microorganisms and microscopic fungi—the so-called Protophyta—were assigned to the Plants, and studied in departments of Botany.
Criticism of this system began in the latter half of the 19th century, with the realization that many organisms met the criteria for inclusion among both plants and animals. For example, the algae Euglena and Dinobryon have chloroplasts for photosynthesis, but can also feed on organic matter and are motile. In 1860, John Hogg argued against the use of "protozoa", on the grounds that "naturalists are divided in opinion—and probably some will ever continue so—whether many of these organisms, or living beings, are animals or plants."As an alternative, he proposed a new kingdom called Primigenum, consisting of both the protozoa and unicellular algae (protophyta), which he combined together under the name "Protoctista". In Hoggs's conception, the animal and plant kingdoms were likened to two great "pyramids" blending at their bases in the Kingdom Primigenum.
Six years later, Ernst Haeckel also proposed a third kingdom of life, which he named Protista. At first, Haeckel included a few multicellular organisms in this kingdom, but in later work he restricted the Protista to single-celled organisms, or simple colonies whose individual cells are not differentiated into different kinds of tissues.
Despite these proposals, Protozoa emerged as the preferred taxonomic placement for heterotrophic microorganisms such as amoebae and ciliates, and remained so for more than a century. In the course of the 20th century, however, the old "two kingdom" system began to weaken, with the growing awareness that fungi did not belong among the plants, and that most of the unicellular protozoa were no more closely related to the animals than they were to the plants. By mid-century, some biologists, such as Herbert Copeland, Robert H. Whittaker and Lynn Margulis, advocated the revival of Haeckel's Protista or Hogg's Protoctista as a kingdom-level eukaryotic group, alongside Plants, Animals and Fungi.A variety of multi-kingdom systems were proposed, and Kingdoms Protista and Protoctista became well established in biology texts and curricula.
While many taxonomists have abandoned Protozoa as a high-level group, Thomas Cavalier-Smith has retained it as a kingdom in the various classifications he has proposed. As of 2015, Cavalier-Smith's Protozoa excludes several major groups of organisms traditionally placed among the protozoa, including the ciliates, dinoflagellates and foraminifera (all members of the SAR supergroup). In its current form, his kingdom Protozoa is a paraphyletic group which includes a common ancestor and most of its descendants, but excludes two important clades that branch within it: the animals and fungi.
Since the protozoa, as traditionally defined, can no longer be regarded as "primitive animals" the terms "protists", "Protista" or "Protoctista" are sometimes preferred. In 2005, members of the Society of Protozoologists voted to change its name to the International Society of Protistologists.
Protozoa, as traditionally defined, range in size from as little as 1 micrometre to several millimetres, or more. cm in diameter.Among the largest are the deep-sea–dwelling xenophyophores, single-celled foraminifera whose shells can reach 20
|Species||Cell type||Size in micrometres|
|Plasmodium falciparum||malaria parasite, trophozoite phase||1-2|
|Massisteria voersi||free-living cercozoan amoeboid||2.3–3|
|Bodo saltans||free living kinetoplastid flagellate||5-8|
|Plasmodium falciparum||malaria parasite, gametocyte phase||7-14|
|Trypanosoma cruzi||parasitic kinetoplastid, Chagas disease||14-24|
|Entamoeba histolytica||parasitic amoebozoan||15–60|
|Balantidium coli||parasitic ciliate||50-100|
|Paramecium caudatum||free-living ciliate||120-330|
|Amoeba proteus||free-living amoebozoan||220–760|
|Noctiluca scintillans||free-living dinoflagellate||700–2000|
|Syringammina fragilissima||foraminiferan amoeboid||up to 200000|
Free-living protozoans are common and often abundant in fresh, brackish and salt water, as well as other moist environments, such as soils and mosses. Some species thrive in extreme environments such as hot springsand hypersaline lakes and lagoons. All protozoa require a moist habitat; however, some can survive for long periods of time in dry environments, by forming resting cysts which enable them to remain dormant until conditions improve.
Parasitic and symbiotic protozoa live on or within other organisms, including vertebrates and invertebrates, as well as plants and other single-celled organisms. Some are harmless or beneficial to their host organisms; others may be significant causes of diseases, such as babesia, malaria and toxoplasmosis.
Association between protozoan symbionts and their host organisms can be mutually beneficial. Flagellated protozoans such as Trichonympha and Pyrsonympha inhabit the guts of termites, where they enable their insect host to digest wood by helping to break down complex sugars into smaller, more easily digested molecules.A wide range of protozoans live commensally in the rumens of ruminant animals, such as cattle and sheep. These include flagellates, such as Trichomonas , and ciliated protozoa, such as Isotricha and Entodinium . The ciliate subclass Astomatia is composed entirely of mouthless symbionts adapted for life in the guts of annelid worms.
All protozoans are heterotrophic, deriving nutrients from other organisms, either by ingesting them whole or consuming their organic remains and waste-products. Some protozoans take in food by phagocytosis, engulfing organic particles with pseudopodia (as amoebae do), or taking in food through a specialized mouth-like aperture called a cytostome. Others take in food by osmotrophy, absorbing dissolved nutrients through their cell membranes.[ citation needed ]
Parasitic protozoans use a wide variety of feeding strategies, and some may change methods of feeding in different phases of their life cycle. For instance, the malaria parasite Plasmodium feeds by pinocytosis during its immature trophozoite stage of life (ring phase), but develops a dedicated feeding organelle (cytostome) as it matures within a host's red blood cell.
Protozoa may also live as mixotrophs, supplementing a heterotrophic diet with some form of autotrophy. Some protozoa form close associations with symbiotic photosynthetic algae, which live and grow within the membranes of the larger cell and provide nutrients to the host. Others practice kleptoplasty, stealing chloroplasts from prey organisms and maintaining them within their own cell bodies as they continue to produce nutrients through photosynthesis. The ciliate Mesodinium rubrum retains functioning plastids from the cryptophyte algae on which it feeds, using them to nourish themselves by autotrophy. These, in turn, may be passed along to dinoflagellates of the genus Dinophysis, which prey on Mesodinium rubrum but keep the enslaved plastids for themselves. Within Dinophysis, these plastids can continue to function for months.
Organisms traditionally classified as protozoa are abundant in aqueous environments and soil, occupying a range of trophic levels. The group includes flagellates (which move with the help of whip-like structures called flagella), ciliates (which move by using hair-like structures called cilia) and amoebae (which move by the use of foot-like structures called pseudopodia). Some protozoa are sessile, and do not move at all.
Unlike plants, fungi and most types of algae, protozoans do not typically have a rigid cell wall, but are usually enveloped by elastic structures of membranes that permit movement of the cell. In some protozoans, such as the ciliates and euglenozoans, the cell is supported by a composite membranous envelope called the "pellicle". The pellicle gives some shape to the cell, especially during locomotion. Pellicles of protozoan organisms vary from flexible and elastic to fairly rigid. In ciliates and Apicomplexa, the pellicle is supported by closely packed vesicles called alveoli. In euglenids, it is formed from protein strips arranged spirally along the length of the body. Familiar examples of protists with a pellicle are the euglenoids and the ciliate Paramecium . In some protozoa, the pellicle hosts epibiotic bacteria that adhere to the surface by their fimbriae (attachment pili).
Some protozoa have two-phase life cycles, alternating between proliferative stages (e.g., trophozoites) and dormant cysts. As cysts, protozoa can survive harsh conditions, such as exposure to extreme temperatures or harmful chemicals, or long periods without access to nutrients, water, or oxygen for periods of time. Being a cyst enables parasitic species to survive outside of a host, and allows their transmission from one host to another. When protozoa are in the form of trophozoites (Greek tropho = to nourish), they actively feed. The conversion of a trophozoite to cyst form is known as encystation, while the process of transforming back into a trophozoite is known as excystation.
Protozoans reproduce asexually by binary fission or multiple fission. Many protozoan species also exchange genetic material by sexual means (typically, through conjugation), but this is generally decoupled from the process of reproduction, and does not immediately result in increased population.
Although meiotic sex is widespread among present day eukaryotes, it has, until recently, been unclear whether or not eukaryotes were sexual early in their evolution. Due to recent advances in gene detection and other techniques, evidence has been found for some form of meiotic sex in an increasing number of protozoans of ancient lineage that diverged early in eukaryotic evolution.(See eukaryote reproduction.) Thus, such findings suggest that meiotic sex arose early in eukaryotic evolution. Examples of protozoan meiotic sexuality are described in the articles Amoebozoa , Giardia lamblia , Leishmania , Plasmodium falciparum biology , Paramecium , Toxoplasma gondii , Trichomonas vaginalis and Trypanosoma brucei .
Historically, the Protozoa were classified as "unicellular animals", as distinct from the Protophyta, single-celled photosynthetic organisms (algae) which were considered primitive plants. Both groups were commonly given the rank of phylum, under the kingdom Protista.In older systems of classification, the phylum Protozoa was commonly divided into several sub-groups, reflecting the means of locomotion. Classification schemes differed, but throughout much of the 20th century the major groups of Protozoa included:
With the emergence of molecular phylogenetics and tools enabling researchers to directly compare the DNA of different organisms, it became evident that, of the main sub-groups of Protozoa, only the ciliates (Ciliophora) formed a natural group, or monophyletic clade (that is, a distinct lineage of organisms sharing common ancestry). The other classes or subphyla of Protozoa were all polyphyletic groups composed of organisms that, despite similarities of appearance or way of life, were not necessarily closely related to one another. In the system of eukaryote classification currently endorsed by the International Society of Protistologists, members of the old phylum Protozoa have been distributed among a variety of supergroups.
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. As predators, they prey upon unicellular or filamentous algae, bacteria, and microfungi. Protozoan species include both herbivores and consumers in the decomposer link of the food chain. They also control bacteria populations and biomass to some extent.
A number of protozoan pathogens are human parasites, causing diseases such as malaria (by Plasmodium ), amoebiasis, giardiasis, toxoplasmosis, cryptosporidiosis, trichomoniasis, Chagas disease, leishmaniasis, African trypanosomiasis (sleeping sickness), Acanthamoeba keratitis, and primary amoebic meningoencephalitis (naegleriasis).
The protozoan Ophryocystis elektroscirrha is a parasite of butterfly larvae, passed from female to caterpillar. Severely infected individuals are weak, unable to expand their wings, or unable to eclose, and have shortened lifespans, but parasite levels vary in populations. Infection creates a culling effect, whereby infected migrating animals are less likely to complete the migration. This results in populations with lower parasite loads at the end of the migration.This is not the case in laboratory or commercial rearing, where after a few generations, all individuals can be infected.
List of protozoan diseases in humans:
|Disease||Causative agent||Source of Transmission|
|Amoebiasis||Entamoeba histolytica (Amoebozoa)||Water, food|
|Acanthamoeba keratitis||Acanthamoeba (Amoebozoa)||Water, contaminated contact lens solution|
|Giardiasis||Giardia lamblia (Metamonada)||Water, Contact|
|Trichomoniasis||Trichomonas vaginalis (Metamonada)||Sexual contact|
|Dientamoebiasis||Dientamoeba fragilis (Metamonada)||Uncertain|
|African sleeping sickness (African trypanosomiasis)||Trypanosoma brucei (Kinetoplastida)||Tsetse fly (Glossina)|
|Chagas disease (American sleeping sickness)||Trypanosoma cruzi (Kinetoplastida)||Triatomine bug (Triatominae)|
|Leishmaniasis||Leishmania spp. (Kinetoplastida)||Phlebotomine Sandfly (Phlebotominae)|
|Balantidiasis||Balantidium coli (Ciliate)||Food, water|
|Malaria||Plasmodium spp. (Apicomplexa)||Mosquito (Anopheles)|
|Toxoplasmosis||Toxoplasma gondii (Apicomplexa)||Undercooked meat, cat feces, fetal infection in pregnancy|
|Babesiosis||Babesia spp. (Apicomplexa)||Deer tick ( Ixodes scapularis )|
|Cryptosporidiosis||Cryptosporidium spp. (Apicomplexa)||Fecal contamination of food or water|
|Cyclosporiasis||Cyclospora cayetanensis (Apicomplexa)||Fecal contamination of food or water|
A microorganism, or microbe, is a microscopic organism, which may exist in its single-celled form or in a colony of cells.
In biology, kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.
The alveolates are a group of protists, considered a major clade and superphylum within Eukarya, and are also called Alveolata.
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free-living and symbiotic forms, and also includes some important parasites of humans, including Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They are classified based on their flagellar structures, and they are considered to be the most basal Flagellate lineage.. Phylogenomic analyses split the members of the Excavates into three different and not all closely related groups: Discobids, Metamonads and Malawimonads. Except for Euglenozoa, they are all non-photosynthetic.
Chromista is a biological kingdom consisting of some single-celled and multicellular eukaryotic organisms, which share similar features in their photosynthetic organelles (plastids). It includes all protists such as some algae, diatoms, oomycetes, and protozoans whose plastids contain chlorophyll c. It is probably a polyphyletic group whose members independently arose as separate evolutionary group from the common ancestor of all eukaryotes. As it is assumed the last common ancestor already possessed chloroplasts of red algal origin, the non-photosynthetic forms evolved from ancestors able to perform photosynthesis. Their plastids are surrounded by four membranes, and are believed to have been acquired from some red algae.
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Unicellular organisms fall into two general categories: prokaryotic organisms and eukaryotic organisms. All prokaryotes are unicellular and are classified into bacteria and archaea. Many eukaryotes are multicellular, but many are unicellular such as protozoa, unicellular algae, and unicellular fungi. Unicellular organisms are thought to be the oldest form of life, with early protocells possibly emerging 3.8–4 billion years ago.
Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow, is a Professor of Evolutionary Biology in the Department of Zoology, at the University of Oxford. His research has led to discovery of a number of unicellular organisms (protists) and definition of taxonomic positions, such as introduction of the kingdom Chromista, and other groups including Chromalveolata, Opisthokonta, Rhizaria, and Excavata. He is well known for his system of classification of all organisms.
The Rhizaria are a species-rich supergroup of mostly unicellular eukaryotes. Except from the Chlorarachniophyte and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has also been described. This supergroup was proposed by Cavalier-Smith in 2002. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. Many produce shells or skeletons, which may be quite complex in structure, and these make up the vast majority of protozoan fossils. Nearly all have mitochondria with tubular cristae.
Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamentous forms. Xanthophyte chloroplasts contain the photosynthetic pigments chlorophyll a, chlorophyll c, β-carotene, and the carotenoid diadinoxanthin. Unlike other heterokonts, their chloroplasts do not contain fucoxanthin, which accounts for their lighter colour. Their storage polysaccharide is chrysolaminarin. Xanthophyte cell walls are produced of cellulose and hemicellulose. They appear to be the closest relatives of the brown algae.
Stephanopogon is a genus of flagellate marine protozoan that superficially resembles a ciliate.
Corticata, in the classification of eukaryotes, is a clade suggested by Thomas Cavalier-Smith to encompass the eukaryote supergroups of the following two groups:
Protozoan infections are parasitic diseases caused by organisms formerly classified in the Kingdom Protozoa. They are usually contracted by either an insect vector or by contact with an infected substance or surface and include organisms that are now classified in the supergroups Excavata, Amoebozoa, SAR, and Archaeplastida.
Monera is a kingdom that contains unicellular organisms with a prokaryotic cell organization, such as bacteria. They are single-celled organisms with no true nuclear membrane.
A protist is any eukaryotic organism that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor, the exclusion of other eukaryotes means that protists do not form a natural group, or clade. So some protists may be more closely related to animals, plants, or fungi than they are to other protists; however, like algae, invertebrates, or protozoans, the grouping is used for convenience. The study of protists is termed protistology.
In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia or Metazoa contains approximately 35 phyla; the plant kingdom Plantae contains about 14, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.
The ciliates are a group of protozoans characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different undulating pattern than flagella. Cilia occur in all members of the group and are variously used in swimming, crawling, attachment, feeding, and sensation.
Eukaryotes are organisms whose cells have a nucleus enclosed within a nuclear envelope. Eukaryotes belong to the domain Eukaryota or Eukarya; their name comes from the Greek εὖ and κάρυον. The domain Eukaryota makes up one of the domains of life in the three-domain system; the two other domains are Bacteria and Archaea. Eukaryotes represent a tiny minority of the number of living organisms; however, due to their generally much larger size, their collective worldwide biomass is estimated to be about equal to that of prokaryotes. Eukaryotes evolved approximately 1.6–2.1 billion years ago, during the Proterozoic eon.
The biological classification system of life introduced by British zoologist Thomas Cavalier-Smith involves systematic arrangements of all life forms on earth. Following and improving the classification systems introduced by Carl Linnaeus, Ernst Haeckel, Robert Whittaker, and Carl Woese, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy. His classification has been a major foundation in modern taxonomy, particularly with revisions and reorganisations of kingdoms and phyla.
The Scotokaryotes (Cavalier-Smith) or Opimoda is a proposed basal Neokaryote clade as sister of the Diaphoratickes. Basal Scotokaryote groupings are the Metamonads, the Malawimonas and the Podiata. The Discoba are sometimes seen as paraphyletic and basal Eukaryotes.
Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are usually single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics, because they are paraphyletic.
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