Protozoa: Protozoa (Singular Protozoon or Protozoan
Protozoa: Protozoa (Singular Protozoon or Protozoan
Protozoa: Protozoa (Singular Protozoon or Protozoan
Despite awareness that the traditional taxonomic concept of "Protozoa" did not meet contemporary
taxonomic standards, some authors have continued to use the name, while applying it to differing scopes of
organisms. In a series of classifications by Thomas Cavalier-Smith and collaborators since 1981, the taxon
Protozoa was applied to a restricted circumscription of organisms, and ranked as a kingdom.[10][11][12] A
scheme presented by Ruggiero et al. in 2015, places eight not closely related phyla within Kingdom
Protozoa: Euglenozoa, Amoebozoa, Metamonada, Choanozoa sensu Cavalier-Smith, Loukozoa,
Percolozoa, Microsporidia and Sulcozoa.[9] Notably, this approach excludes several major groups of
organisms traditionally placed among the protozoa, including the ciliates, dinoflagellates, foraminifera, and
the parasitic apicomplexans, which were located in other groups such as Alveolata and Stramenopiles,
under the polyphyletic Chromista. The Protozoa in this scheme do not form a monophyletic and
holophyletic group (clade), but a paraphyletic group or evolutionary grade, because it excludes some
descendants of Protozoa, as used in this sense.[9]
Contents
History
Characteristics
Reproduction
Size
Habitat
Feeding
Motility
Walls, pellicles, scales, and skeletons
Life cycle
Classification
Ecology
Disease
References
Bibliography
External links
History
The word "protozoa" (singular protozoon) was coined
in 1818 by zoologist Georg August Goldfuss
(=Goldfuß), as the Greek equivalent of the German
Urthiere, meaning "primitive, or original animals" (ur-
‘proto-’ + Thier ‘animal’).[13] Goldfuss created
Protozoa as a class containing what he believed to be
the simplest animals.[5] 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.[14]
The term Protozoa is formed from the Greek words
πρῶτος (prôtos), meaning "first", and ζῶα (zôa),
Class Protozoa, order Infusoria, family Monades plural of ζῶον (zôon), meaning "animal".[15][16] The
by Georg August Goldfuss, c. 1844 use of Protozoa as a formal taxon has been
discouraged by some researchers, mainly because the
term implies kinship with animals (Metazoa)[17][18]
and promotes an arbitrary separation of "animal-like" from "plant-like" organisms.[19]
In 1848, as a result of advancements in the design and construction of microscopes and the emergence of a
cell theory pioneered by Theodor Schwann and Matthias Schleiden, the anatomist and zoologist C. T. von
Siebold proposed that the bodies of protozoa 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).[20] At the same
time, he raised the group to the level of a phylum containing two broad classes of microorganisms:
Infusoria (mostly ciliates) and flagellates (flagellated protists) and amoebae (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".[21] With its
increasing visibility, the term 'protozoa' and the discipline of 'protozoology' came into wide use.
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, 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.[22] A variety of multi-kingdom
systems were proposed, and the Kingdoms Protista and Protoctista became established in biology texts and
curricula.[23][24][25]
While most taxonomists have abandoned Protozoa as a high-level group, Cavalier-Smith used the term with
a different circumscription. In 2015, Protozoa sensu Cavalier-Smith excluded several major groups of
organisms traditionally placed among the protozoa (such as ciliates, dinoflagellates and foraminifera). This
and similar concepts of Protozoa are of a paraphyletic group which does not include all organisms that
descended from Protozoa. In this case, the most significant absences were of the animals and fungi.[9] The
continued use by some of the 'Protozoa' in its old sense[26] highlights the uncertainty as to what is meant by
the word 'Protozoa', the need for disambiguating statements (here, the term 'Protozoa' is used in the sense
intended by Goldfuß), and the problems that arise when new meanings are given to familiar taxonomic
terms.
Some authors classify Protozoa as a subgroup of mostly motile Protists.[27] Others class any unicellular
eukaryotic microorganism as a Protist, and make no reference to 'Protozoa'.[28]
In 2005, members of the Society of Protozoologists voted to change its name to the International Society of
Protistologists.[29]
Characteristics
Reproduction
Reproduction in Protozoa can be sexual or asexual.[30] Most Protozoa reproduce asexually through binary
fission.[31]
Many parasitic Protozoa reproduce both asexually and sexually.[30] However, sexual reproduction is rare
among free-living protozoa and it usually occurs when food is scarce or the environment changes
drastically.[32] Both isogamy and anisogamy occur in Protozoa with anisogamy being the more common
form of sexual reproduction.[33]
Size
Protozoa, as traditionally defined, range in size from as little as 1 micrometre to several millimetres, or
more.[34] Among the largest are the deep-sea–dwelling xenophyophores, single-celled foraminifera whose
shells can reach 20 cm in diameter.[35]
Size in
Species Cell type
micrometres
Paramecium
free-living ciliate[43] 120–330
caudatum
Noctiluca
free-living dinoflagellate[45] 700–2000
scintillans
Syringammina up to
foraminifera amoeba[35]
fragilissima 200 000
Habitat
Free-living protozoa 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
springs[46] and hypersaline lakes and lagoons.[47] All protozoa require a moist habitat; however, some can
survive for long periods of time in dry environments, by forming resting cysts that 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.
Feeding
All protozoa are heterotrophic, deriving nutrients from other organisms, either by ingesting them whole by
phagocytosis or taking up dissolved organic matter or micro-particles (osmotrophy). Phagocytosis may
involve engulfing organic particles with pseudopodia (as amoebae do), taking in food through a specialized
mouth-like aperture called a cytostome, or using stiffened ingestion organelles[51]
Parasitic protozoa 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.[52]
Protozoa may also live as mixotrophs, combining a heterotrophic diet with some form of autotrophy. Some
protozoa form close associations with symbiotic photosynthetic algae (zoochlorellae), which live and grow
within the membranes of the larger cell and provide nutrients to the host. The algae are not digested, but
reproduce and are distributed between division products. The organism may benefit at times by deriving
some of its nutrients from the algal endosymbionts or by surviving anoxic conditions because of the oxygen
produced by algal photosynthesis. Some protozoans 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. The symbionts 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.[53]
Motility
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
undulating and beating flagella). Ciliates (which move by using
hair-like structures called cilia) and amoebae (which move by the
use of temporary extensions of cytoplasm called pseudopodia).
Many protozoa, such as the agents of amoebic meningitis, use both
pseudopodia and flagella. Some protozoa attach to the substrate or
form cysts so they do not move around (sessile). Most sessile
protozoa are able to move around at some stage in the life cycle,
such as after cell division. The term 'theront' has been used for
actively motile phases, as opposed to 'trophont' or 'trophozoite' that
refers to feeding stages.
Life cycle
Protozoa mostly reproduce asexually by binary fission or multiple fission. Many protozoa 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.[55] Thus, sexuality can be
optional.
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. Owing to recent advances in gene detection
and other techniques, evidence has been found for some form of meiotic sex in an increasing number of
protozoa of lineages that diverged early in eukaryotic evolution.[56]
(See eukaryote reproduction.) 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.
Life cycle of parasitic protozoan,
Classification Toxoplasma gondii
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, once a few extraneous members (such as
Stephanopogon or protociliates and opalinids were removed. The Mastigophora, Sarcodina, and Sporozoa
were polyphyletic groups. The similarities of appearance and ways of life by which these groups were
defined had emerged independently in their members by convergent evolution.
In most systems of eukaryote classification, such as one published by the International Society of
Protistologists, members of the old phylum Protozoa have been distributed among a variety of
supergroups.[59]
Ecology
Free-living protozoa are found in almost all ecosystems that contain, at least some of the time, free water.
They have a critical role in the mobilization of nutrients in natural ecosystems. Their role is best conceived
within the context of the microbial food web in which they include the most important bacterivores.[51] In
part, they facilitate the transfer of bacterial and algal production to successive trophic levels, but also they
solubilize the nutrients within microbial biomass, allowing stimulation of microbial growth. As consumers,
protozoa prey upon unicellular or filamentous algae, bacteria, microfungi, and micro-carrion. In the context
of older ecological models of the micro- and meiofauna, protozoa may be a food source for
microinvertebrates.
That most species of free-living protozoa have been found in similar habitats in all parts of the globe is an
observation that dates back to the 19th Century (e.g. Schewiakoff). In the 1930s, Lourens Baas Becking
asserted "Everything is everywhere, but the environment selects". This has been restated and explained,
especially by Tom Fenchel and Bland Findlay[60] and methodically explored and affirmed at least in
respect of morphospecies of free-living flagellates.[61][62] The widespread distribution of microbial is
explained by the ready dispersal of physically small organisms. While Baas Becking's hypothesis is not
universally accepted,[63] the natural microbial world is undersampled, and this will favour conclusions of
endemism.
Disease
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Chisholm, Hugh, ed. (1911). "Protozoa" (https://en.wikisource.org/wiki/1911_Encyclop%C
3%A6dia_Britannica/Protozoa). Encyclopædia Britannica (11th ed.). Cambridge University
Press.