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An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
or
unicellular organism 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. Organisms fall into two general categories: prokaryotic organisms and ...
with the ability to alter its shape, primarily by extending and retracting
pseudopods A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filamen ...
. Amoebae do not form a single
taxonomic group In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
; instead, they are found in every major
lineage Lineage may refer to: Science * Lineage (anthropology), a group that can demonstrate its common descent from an apical ancestor or a direct line of descent from an ancestor * Lineage (evolution), a temporal sequence of individuals, populati ...
of
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
organisms. Amoeboid cells occur not only among the
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
, but also in
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, and
animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
.
Microbiologist A microbiologist (from Ancient Greek, Greek ) is a scientist who studies microscopic life forms and processes. This includes study of the growth, interactions and characteristics of Microorganism, microscopic organisms such as bacteria, algae, f ...
s often use the terms "amoeboid" and "amoeba" interchangeably for any organism that exhibits
amoeboid movement Amoeboid movement is the most typical mode of locomotion in adherent eukaryotic cells. It is a crawling-like type of movement accomplished by protrusion of cytoplasm of the cell involving the formation of pseudopodia ("false-feet") and posterior ...
. In older classification systems, most amoebae were placed in the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
or
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of "subdivision (rank), subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists ...
Sarcodina, a grouping of single-celled organisms that possess pseudopods or move by
protoplasm Protoplasm (; ) is the living part of a cell that is surrounded by a plasma membrane. It is a mixture of small molecules such as ions, monosaccharides, amino acid, and macromolecules such as proteins, polysaccharides, lipids, etc. In some defini ...
ic flow. However,
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have shown that Sarcodina is not a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group whose members share
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
. Consequently, amoeboid organisms are no longer classified together in one group.Jan Pawlowski: ''The twilight of Sarcodina: a molecular perspective on the polyphyletic origin of amoeboid protists''. Protistology, Band 5, 2008, S. 281–302
(pdf, 570 kB)
The best known amoeboid
protists A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
are ''
Chaos carolinense ''Chaos'' is a genus of single-celled amoeboid organisms in the family Amoebidae. The largest and best-known species, the so-called "giant amoeba" ''Chaos carolinensis'', can reach lengths of 5 mm, although most specimens fall between 1 an ...
'' and ''
Amoeba proteus ''Amoeba proteus'' is a large species of amoeba closely related to another genus of giant amoebae, ''Chaos''. As such, the species is sometimes given the alternative scientific name ''Chaos diffluens''. This protozoan uses extensions called pseu ...
'', both of which have been widely cultivated and studied in classrooms and laboratories. Other well known species include the so-called "brain-eating amoeba" ''
Naegleria fowleri ''Naegleria fowleri'', colloquially known as a "brain-eating amoeba", is a species of the genus ''Naegleria'', belonging to the phylum Percolozoa, which is technically not classified as true amoeba, but a shapeshifting amoeboflagellate excava ...
'', the intestinal parasite ''
Entamoeba histolytica ''Entamoeba histolytica'' is an anaerobic parasitic amoebozoan, part of the genus ''Entamoeba''. Predominantly infecting humans and other primates causing amoebiasis, ''E. histolytica'' is estimated to infect about 35-50 million people worldwid ...
'', which causes
amoebic dysentery Amoebiasis, or amoebic dysentery, is an infection of the intestines caused by a parasitic amoeba ''Entamoeba histolytica''. Amoebiasis can be present with no, mild, or severe symptoms. Symptoms may include lethargy, loss of weight, colonic ulc ...
, and the multicellular "social amoeba" or
slime mould Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
''
Dictyostelium discoideum ''Dictyostelium discoideum'' is a species of soil-dwelling amoeba belonging to the phylum Amoebozoa, infraphylum Mycetozoa. Commonly referred to as slime mold, ''D. discoideum'' is a eukaryote that transitions from a collection of unicellular ...
''.


Shape, movement and nutrition

Amoeba do not have cell walls, which allows for free movement. Amoeba move and feed by using pseudopods, which are bulges of
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
formed by the coordinated action of
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
microfilaments Microfilaments, also called actin filaments, are protein filaments in the cytoplasm of eukaryotic cells that form part of the cytoskeleton. They are primarily composed of polymers of actin, but are modified by and interact with numerous other pr ...
pushing out the
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
that surrounds the cell. The appearance and internal structure of pseudopods are used to distinguish groups of amoebae from one another.
Amoebozoan Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported classi ...
species, such as those in the genus ''
Amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of Cell (biology), cell or unicellular organism with the ability to alter its shape, primarily by extending and ret ...
'', typically have bulbous (lobose) pseudopods, rounded at the ends and roughly tubular in cross-section.
Cercozoan Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major e ...
amoeboids, such as ''
Euglypha ''Euglypha'' is a genus of cercozoa Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or pol ...
'' and ''
Gromia ''Gromia'' is a genus of protists, closely related to foraminifera, which inhabit marine and freshwater environments. It is the only genus of the family Gromiidae. ''Gromia'' are ameboid, producing filose pseudopodia that extend out from the cel ...
'', have slender, thread-like (filose) pseudopods.
Foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
emit fine, branching pseudopods that merge with one another to form net-like (reticulose) structures. Some groups, such as the
Radiolaria The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell (biology), cell into the inner and outer portions of endoplasm and Ecto ...
and
Heliozoa Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms ( axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections fro ...
, have stiff, needle-like, radiating axopodia (actinopoda) supported from within by bundles of
microtubules Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
. Free-living amoebae may be "
testate Intestacy is the condition of the estate of a person who dies without having in force a valid will or other binding declaration. Alternatively this may also apply where a will or declaration has been made, but only applies to part of the estat ...
" (enclosed within a hard shell), or "naked" (also known as gymnamoebae, lacking any hard covering). The shells of testate amoebae may be composed of various substances, including
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
,
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
,
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, or agglutinations of found materials like small grains of sand and the
frustules A frustule is the hard and porous cell wall or external layer of diatoms. The frustule is composed almost purely of silica, made from silicic acid, and is coated with a layer of organic substance, which was referred to in the early literature on ...
of
diatoms A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
. To regulate
osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in a pure ...
, most freshwater amoebae have a contractile
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
which expels excess water from the cell. This
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
is necessary because freshwater has a lower concentration of
solutes In chemistry, a solution is a special type of homogeneous mixture composed of two or more substances. In such a mixture, a solute is a substance dissolved in another substance, known as a solvent. If the attractive forces between the solvent ...
(such as salt) than the amoeba's own internal fluids (
cytosol The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells (intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...
). Because the surrounding water is
hypotonic In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane-imp ...
with respect to the contents of the cell, water is transferred across the amoeba's cell membrane by
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
. Without a contractile vacuole, the cell would fill with excess water and, eventually, burst. Marine amoebae do not usually possess a contractile vacuole because the concentration of solutes within the cell are in balance with the
tonicity In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane-im ...
of the surrounding water.


Diet

The food sources of amoebae vary. Some amoebae are predatory and live by consuming bacteria and other
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s. Some are
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s and eat dead organic material. Amoebae typically ingest their food by
phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
, extending pseudopods to encircle and engulf live prey or particles of scavenged material. Amoeboid cells do not have a mouth or
cytostome A cytostome (from ''cyto-'', cell and ''stome-'', mouth) or cell mouth is a part of a cell specialized for phagocytosis, usually in the form of a microtubule-supported funnel or groove. Food is directed into the cytostome, and sealed into vacuole ...
, and there is no fixed place on the cell at which phagocytosis normally occurs. Some amoebae also feed by
pinocytosis In cellular biology, pinocytosis, otherwise known as fluid endocytosis and bulk-phase pinocytosis, is a mode of endocytosis in which small molecules dissolved in extracellular fluid are brought into the cell through an invagination of the cell me ...
, imbibing dissolved nutrients through
vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
formed within the cell membrane.


Size range

The size of amoeboid cells and species is extremely variable. The marine amoeboid '' Massisteria voersi'' is just 2.3 to 3
micrometre The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
s in diameter, within the size range of many bacteria. At the other extreme, the shells of deep-sea xenophyophores can attain 20 cm in diameter. Most of the free-living freshwater amoebae commonly found in pond water, ditches, and lakes are
microscopic The microscopic scale () is the scale of objects and events smaller than those that can easily be seen by the naked eye, requiring a lens (optics), lens or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded a ...
, but some species, such as the so-called "giant amoebae" ''Pelomyxa palustris'' and ''Chaos carolinense'', can be large enough to see with the naked eye.


Amoebae as specialized cells and life cycle stages

Some
multicellular organisms A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
have amoeboid cells only in certain phases of life, or use amoeboid movements for specialized functions. In the immune system of humans and other animals, amoeboid
white blood cells White blood cells, also called leukocytes or leucocytes, are the cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and derived from mult ...
pursue invading organisms, such as bacteria and pathogenic protists, and engulf them by
phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
. Amoeboid stages also occur in the multicellular fungus-like protists, the so-called
slime mould Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
s. Both the plasmodial slime moulds, currently classified in the class
Myxogastria Myxogastria/Myxogastrea (myxogastrids, International Code of Zoological Nomenclature, ICZN) or Myxomycetes (International Code of Nomenclature for algae, fungi, and plants, ICN), is a Class (biology), class of slime molds that contains 5 o ...
, and the cellular slime moulds of the groups
Acrasida The family Acrasidae ( ICZN, or Acrasiomycota, ICBN) is a family of slime molds which belongs to the excavate group Percolozoa. The name element - comes from the Greek ''akrasia'', meaning "acting against one's judgement". This group consists ...
and Dictyosteliida, live as amoebae during their feeding stage. The amoeboid cells of the former combine to form a giant
multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinat ...
organism, while the cells of the latter live separately until food runs out, at which time the amoebae aggregate to form a multicellular migrating "slug" which functions as a single organism. Other organisms may also present amoeboid cells during certain life-cycle stages, e.g., the gametes of some green algae (
Zygnematophyceae Zygnematophyceae (or Conjugatophyceae) is a class of green algae in the paraphylum streptophyte algae, also referred to as Charophyta, consisting of more than 4000 described species. It contains five orders: the Spirogloeales, the Serritaeniales, ...
) and pennate
diatoms A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
, the spores (or dispersal phases) of some
Mesomycetozoea The Mesomycetozoea (or DRIP clade, or Ichthyosporea) are a small group of Opisthokonta in Eukaryota (formerly protists), mostly parasites of fish and other animals. Significance They are not particularly distinctive morphologically, appearing in ...
, and the
sporoplasm Sporoplasm is an infectious material present in the cytoplasm of various fungi-like organisms, such as members of class ''Microsporidia''. Sporoplasm is defined as a mass of protoplasm that gives rise to or forms a spore. The protoplasmic body t ...
stage of
Myxozoa Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. O ...
and of
Ascetosporea The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...
.


Amoebae as organisms


Early history and origins of Sarcodina

The earliest record of an amoeboid organism was produced in 1755 by
August Johann Rösel von Rosenhof August Johann Rösel von Rosenhof (March 30, 1705 in Augustenburg near Arnstadt – March 27, 1759 in Nuremberg) was a German miniature painter, naturalist and entomologist. With his accurate, heavily detailed images of insects he was reco ...
, who named his discovery "Der Kleine Proteus" ("the Little Proteus"). Rösel's illustrations show an unidentifiable freshwater amoeba, similar in appearance to the common species now known as
Amoeba proteus ''Amoeba proteus'' is a large species of amoeba closely related to another genus of giant amoebae, ''Chaos''. As such, the species is sometimes given the alternative scientific name ''Chaos diffluens''. This protozoan uses extensions called pseu ...
. The term "Proteus animalcule" remained in use throughout the 18th and 19th centuries, as an informal name for any large, free-living amoeboid. In 1822, the genus ''Amiba'' (from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
ἀμοιβή ''amoibe'', meaning "change") was erected by the French naturalist
Bory de Saint-Vincent Jean-Baptiste Geneviève Marcellin Bory de Saint-Vincent was a French natural history, naturalist, Officer (armed forces), officer and politician. He was born on 6 July 1778 in Agen (Lot-et-Garonne) and died on 22 December 1846 in Paris. Biologis ...
. Bory's contemporary, C. G. Ehrenberg, adopted the genus in his own classification of microscopic creatures, but changed the spelling to ''Amoeba''. In 1841,
Félix Dujardin Félix Dujardin (5 April 1801 – 8 April 1860) was a French biologist born in Tours. He is remembered for his research on protozoans and other invertebrates. Biography In 1840 he was appointed professor of geology and mineralogy at the Univer ...
coined the term "''sarcode''" (from Greek σάρξ ''sarx'', "flesh," and εἶδος ''eidos'', "form") for the "thick, glutinous, homogeneous substance" which fills protozoan cell bodies. Although the term originally referred to the protoplasm of any protozoan, it soon came to be used in a restricted sense to designate the gelatinous contents of amoeboid cells. Thirty years later, the Austrian zoologist
Ludwig Karl Schmarda Ludwig Karl Schmarda (23 August 1819 – 7 April 1908) was an Austrians, Austrian natural science, naturalist and traveler, born at Olomouc, Olmütz, Moravia. Early life and education Schmarda was born at Olomouc, Olmütz where he attended the ...
used "sarcode" as the conceptual basis for his division Sarcodea, a
phylum In biology, a phylum (; plural: phyla) 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 f ...
-level group made up of "unstable, changeable" organisms with bodies largely composed of "sarcode". Later workers, including the influential taxonomist
Otto Bütschli Johann Adam Otto Bütschli (3 May 1848 – 2 February 1920) was a German zoologist and professor at the University of Heidelberg. He specialized in invertebrates and insect development. Many of the groups of protists were first recognized by him. ...
, amended this group to create the class Sarcodina, a
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
that remained in wide use throughout most of the 20th century. Within the traditional Sarcodina, amoebae were generally divided into morphological categories, on the basis of the form and structure of their
pseudopod A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filamen ...
s. Amoebae with pseudopods supported by regular arrays of
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s (such as the freshwater
Heliozoa Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms ( axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections fro ...
and marine
Radiolaria The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell (biology), cell into the inner and outer portions of endoplasm and Ecto ...
) were classified as Actinopoda; whereas those with unsupported pseudopods were classified as
Rhizopoda An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of Cell (biology), cell or unicellular organism with the ability to alter its shape, primarily by extending and ret ...
. The Rhizopods were further subdivided into lobose, filose, and reticulose amoebae, according to the morphology of their pseudopods.


Dismantling of Sarcodina

In the final decade of the 20th century, a series of molecular phylogenetic analyses confirmed that Sarcodina was not a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group. In view of these findings, the old scheme was abandoned and the amoebae of Sarcodina were dispersed among many other high-level taxonomic groups. Today, the majority of traditional sarcodines are placed in two eukaryote supergroups:
Amoebozoa Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported classi ...
and
Rhizaria The Rhizaria are an ill-defined but species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foramini ...
. The rest have been distributed among the excavates,
opisthokonts The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
, and
stramenopiles Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
. Some, like the
Centrohelida The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth. Characteristics Individuals are unicellular and spher ...
, have yet to be placed in any supergroup.


Classification

Recent classification places the various amoeboid genera in the following groups: Some of the amoeboid groups cited (e.g., part of
chrysophyte The Chrysophyceae, usually called chrysophytes, chrysomonads, golden-brown algae or golden algae are a large group of algae, found mostly in freshwater. Golden algae is also commonly used to refer to a single species, ''Prymnesium parvum'', which ...
s, part of
xanthophyte 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 filament ...
s,
chlorarachniophyte The chlorarachniophytes are a small group of exclusively marine algae widely distributed in tropical and temperate waters. They are typically mixotrophic, ingesting bacteria and smaller protists as well as conducting photosynthesis. Normally the ...
s) were not traditionally included in Sarcodina, being classified as
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
or
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
d protozoa.


Pathogenic interactions with other organisms

Some amoebae can infect other organisms
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
ically, causing disease: *''
Entamoeba histolytica ''Entamoeba histolytica'' is an anaerobic parasitic amoebozoan, part of the genus ''Entamoeba''. Predominantly infecting humans and other primates causing amoebiasis, ''E. histolytica'' is estimated to infect about 35-50 million people worldwid ...
'' is the cause of
amoebiasis Amoebiasis, or amoebic dysentery, is an infection of the intestines caused by a parasitic amoeba ''Entamoeba histolytica''. Amoebiasis can be present with no, mild, or severe symptoms. Symptoms may include lethargy, loss of weight, colonic ulce ...
, or amoebic dysentery. *''
Naegleria fowleri ''Naegleria fowleri'', colloquially known as a "brain-eating amoeba", is a species of the genus ''Naegleria'', belonging to the phylum Percolozoa, which is technically not classified as true amoeba, but a shapeshifting amoeboflagellate excava ...
'' (the "brain-eating amoeba") is a fresh-water-native species that can be fatal to humans if introduced through the nose. *''
Acanthamoeba ''Acanthamoeba'' is a genus of amoeboid, amoebae that are commonly recovered from soil, fresh water, and other habitat (ecology), habitats. ''Acanthamoeba'' has two evolutive forms, the metabolically active trophozoite and a dormant, stress-resi ...
'' can cause amoebic
keratitis Keratitis is a condition in which the eye's cornea, the clear dome on the front surface of the eye, becomes inflamed. The condition is often marked by moderate to intense pain and usually involves any of the following symptoms: pain, impaired e ...
and
encephalitis Encephalitis is inflammation of the brain. The severity can be variable with symptoms including reduction or alteration in consciousness, headache, fever, confusion, a stiff neck, and vomiting. Complications may include seizures, hallucinations, ...
in humans. *''
Balamuthia mandrillaris ''Balamuthia mandrillaris'' is a free-living amoeba that causes the rare but deadly neurological condition granulomatous amoebic encephalitis (GAE). ''B. mandrillaris'' is a soil-dwelling amoeba and was first discovered in 1986 in the brain of ...
'' is the cause of (often fatal)
granulomatous amoebic meningoencephalitis Granulomatous amoebic encephalitis (GAE) is a rare, usually fatal, subacute-to-chronic central nervous system disease caused by certain species of free-living amoebae of the genera ''Acanthamoeba'', ''Balamuthia'' and ''Sappinia''. The term is m ...
. Amoeba have been found to harvest and grow the bacteria implicated in
plague Plague or The Plague may refer to: Agriculture, fauna, and medicine *Plague (disease), a disease caused by ''Yersinia pestis'' * An epidemic of infectious disease (medical or agricultural) * A pandemic caused by such a disease * A swarm of pes ...
. Amoebae can likewise play host to microscopic organisms that are pathogenic to people and help in spreading such microbes. Bacterial pathogens (for example, ''
Legionella ''Legionella'' is a genus of pathogenic gram-negative bacteria that includes the species '' L. pneumophila'', causing legionellosis (all illnesses caused by ''Legionella'') including a pneumonia-type illness called Legionnaires' disease and a mi ...
'') can oppose absorption of food when devoured by amoebae. The currently generally utilized and best-explored amoebae that host other organisms are Acanthamoeba castellanii and Dictyostelium discoideum. Microorganisms that can overcome the defenses of one-celled organisms can shelter and multiply inside them, where they are shielded from unfriendly outside conditions by their hosts.


Meiosis

Recent evidence indicates that several Amoebozoa lineages undergo
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
.
Orthologs Sequence homology is the biological homology between DNA, RNA, or protein sequences, defined in terms of shared ancestry in the evolutionary history of life. Two segments of DNA can have shared ancestry because of three phenomena: either a spec ...
of genes employed in
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
of sexual
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s have recently been identified in the ''
Acanthamoeba ''Acanthamoeba'' is a genus of amoeboid, amoebae that are commonly recovered from soil, fresh water, and other habitat (ecology), habitats. ''Acanthamoeba'' has two evolutive forms, the metabolically active trophozoite and a dormant, stress-resi ...
''
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
. These genes included ''
Spo11 Spo11 is a protein that in humans is encoded by the ''SPO11'' gene. Spo11, in a complex with mTopVIB, creates double strand breaks to initiate meiotic recombination. Its active site contains a tyrosine which ligates and dissociates with DNA to pr ...
,
Mre11 Double-strand break repair protein MRE11 is an enzyme that in humans is encoded by the ''MRE11'' gene. The gene has been designated ''MRE11A'' to distinguish it from the pseudogene ''MRE11B'' that is nowadays named ''MRE11P1''. Function This ge ...
,
Rad50 DNA repair protein RAD50, also known as RAD50, is a protein that in humans is encoded by the ''RAD50'' gene. Function The protein encoded by this gene is highly similar to ''Saccharomyces cerevisiae'' Rad50, a protein involved in DNA double- ...
,
Rad51 DNA repair protein RAD51 homolog 1 is a protein encoded by the gene ''RAD51''. The enzyme encoded by this gene is a member of the RAD51 protein family which assists in repair of DNA double strand breaks. RAD51 family members are homologous to th ...
,
Rad52 RAD52 homolog (S. cerevisiae), also known as RAD52, is a protein which in humans is encoded by the ''RAD52'' gene. Function The protein encoded by this gene shares similarity with ''Saccharomyces cerevisiae'' Rad52, a protein important for DNA ...
, Mnd1,
Dmc1 Meiotic recombination protein DMC1/LIM15 homolog is a protein that in humans is encoded by the ''DMC1'' gene. Meiotic recombination protein Dmc1 is a homolog of the bacterial strand exchange protein RecA. Dmc1 plays the central role in homologous ...
, Msh'' and '' Mlh''. This finding suggests that the ‘'Acanthamoeba'’ are capable of some form of meiosis and may be able to undergo sexual reproduction. The meiosis-specific
recombinase Recombinases are genetic recombination enzymes. Site specific recombinases DNA recombinases are widely used in multicellular organisms to manipulate the structure of genomes, and to control gene expression. These enzymes, derived from bacteria (b ...
,
Dmc1 Meiotic recombination protein DMC1/LIM15 homolog is a protein that in humans is encoded by the ''DMC1'' gene. Meiotic recombination protein Dmc1 is a homolog of the bacterial strand exchange protein RecA. Dmc1 plays the central role in homologous ...
, is required for efficient meiotic
homologous recombination Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may ...
, and ''Dmc1'' is expressed in ''
Entamoeba histolytica ''Entamoeba histolytica'' is an anaerobic parasitic amoebozoan, part of the genus ''Entamoeba''. Predominantly infecting humans and other primates causing amoebiasis, ''E. histolytica'' is estimated to infect about 35-50 million people worldwid ...
''. The purified Dmc1 from ''E. histolytica'' forms
presynaptic In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell. Synapses are essential to the transmission of nervous impulses from ...
filaments and catalyses ATP-dependent homologous DNA pairing and DNA strand exchange over at least several thousand
base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
. The DNA pairing and strand exchange reactions are enhanced by the eukaryotic meiosis-specific recombination accessory factor (heterodimer) Hop2-Mnd1. These processes are central to meiotic recombination, suggesting that ''E. histolytica'' undergoes meiosis. Studies of '' Entamoeba invadens'' found that, during the conversion from the
tetraploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
uninucleate {{Short pages monitor