# Pseudopodia

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False leg found on slime molds, archaea, protozoans, leukocytes and certain bacteria

This article is about eukaryotic cells. For the band, see [Pseudopod (band)](/source/Pseudopod_(band)). For the podcast, see [Pseudopod (podcast)](/source/Pseudopod_(podcast)). For the structure in insect anatomy, see [Proleg](/source/Proleg).

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*Amoeba proteus* extending lobose pseudopodia

A **pseudopod** or **pseudopodium** (pl.: **pseudopods** or **pseudopodia**) is a temporary arm-like projection of an [eukaryotic](/source/Eukaryotic) [cell membrane](/source/Cell_membrane) that is emerged in the direction of movement. Filled with [cytoplasm](/source/Cytoplasm), pseudopodia primarily consist of [actin filaments](/source/Actin_filaments) and may also contain [microtubules](/source/Microtubule) and [intermediate filaments](/source/Intermediate_filament).[1][2] Pseudopods are used for [motility](/source/Motility) and [ingestion](/source/Ingestion). They are often found in [amoebas](/source/Amoeba).

Different types of pseudopodia can be classified by their distinct appearances.[3] [Lamellipodia](/source/Lamellipodia) are broad and thin. [Filopodia](/source/Filopodia) are slender, thread-like, and are supported largely by microfilaments. Lobopodia are bulbous and amoebic. [Reticulopodia](/source/Reticulopodia) are complex structures bearing individual pseudopodia which form irregular nets. [Axopodia](#Axopodia) are the [phagocytosis](/source/Phagocytosis) type with long, thin pseudopods supported by complex microtubule arrays enveloped with cytoplasm; they respond rapidly to physical contact.[4]

Generally, several pseudopodia arise from the surface of the body, (*polypodial*, for example, *[Amoeba proteus](/source/Amoeba_proteus)*), or a single pseudopod may form on the surface of the body (*monopodial*, such as *[Entamoeba histolytica](/source/Entamoeba_histolytica)*).[5]

## Formation

Cells which make pseudopods are generally referred to as *[amoeboids](/source/Amoeboids)*.[6]

### Via extracellular cue

To move towards a target, the cell uses [chemotaxis](/source/Chemotaxis). It senses extracellular signalling molecules, chemoattractants (e.g. cAMP for *[Dictyostelium](/source/Dictyostelium)* cells),[7] to extend pseudopodia at the membrane area facing the source of these molecules.

The chemoattractants bind to [G protein-coupled receptors](/source/G_protein-coupled_receptor), which activate [GTPases of the Rho family](/source/Rho_family_of_GTPases) (e.g. Cdc42, Rac) via [G proteins](/source/G_protein).

Rho GTPases are able to activate [WASp](/source/Wiskott%E2%80%93Aldrich_syndrome_protein) which in turn activate [Arp2/3 complex](/source/Arp2%2F3_complex) which serve as nucleation sites for [actin polymerization](/source/Actin#Nucleation_and_polymerization).[8] The actin polymers then push the membrane as they grow, forming the pseudopod. The pseudopodium can then adhere to a surface via its [adhesion proteins](/source/Cell_adhesion) (e.g. [integrins](/source/Integrin)), and then pull the cell's body forward via contraction of an actin-myosin complex in the pseudopod.[9][10] This type of locomotion is called *[amoeboid movement](/source/Amoeboid_movement)*.

Rho GTPases can also activate [phosphatidylinositol 3-kinase](/source/Phosphatidylinositol_3-kinase) (PI3K) which recruit [PIP3](/source/Phosphatidylinositol_(3%2C4%2C5)-trisphosphate) to the membrane at the leading edge and detach the PIP3-degrading enzyme [PTEN](/source/PTEN_(gene)) from the same area of the membrane. PIP3 then activate GTPases back via [GEF](/source/Guanine_nucleotide_exchange_factor) stimulation. This serves as a feedback loop to amplify and maintain the presence of local GTPase at the leading edge.[8]

Otherwise, pseudopodia cannot grow on other sides of the membrane than the leading edge because myosin filaments prevent them to extend. These myosin filaments are induced by [cyclic GMP](/source/Cyclic_guanosine_monophosphate) in *[D. discoideum](/source/Dictyostelium_discoideum)* or [Rho kinase](/source/Rho_kinase) in [neutrophils](/source/Neutrophil) for example.[8]

Different physical parameters were shown to regulate the length and time-scale of pseudopodia formation. For example, an increase in membrane [tension](/source/Tension_(physics)) inhibits actin assembly and protrusion formation.[11] It was demonstrated that the lowered negative [surface charge](/source/Surface_charge) on the inner surface of the [plasma membrane](/source/Cell_membrane) generates protrusions via activation of the Ras-[PI3K/AKT/mTOR](/source/PI3K%2FAKT%2FmTOR_pathway) signalling pathway.[12]

### Without extracellular cue

In the case there is no extracellular cue, all moving cells navigate in random directions, but they can keep the same direction for some time before turning. This feature allows cells to explore large areas for colonization or searching for a new extracellular cue.

In *Dictyostelium* cells, a pseudopodium can form either *de novo* as normal, or from an existing pseudopod, forming a Y-shaped pseudopodium.

The Y-shaped pseudopods are used by *Dictyostelium* to advance relatively straight forward by alternating between retraction of the left or right branch of the pseudopod. The *de novo* pseudopodia form at different sides than pre-existing ones, they are used by the cells to turn.

Y-shaped pseudopods are more frequent than *de novo* ones, which explain the preference of the cell to keep moving to the same direction. This persistence is modulated by [PLA2](/source/Phospholipase_A2) and cGMP signalling pathways.[7]

## Functions

The functions of pseudopodia include locomotion and ingestion:

- Pseudopodia are critical in sensing targets which can then be engulfed; the engulfing pseudopodia are called [phagocytosis](/source/Phagocytosis) pseudopodia. A common example of this type of amoeboid cell is the [macrophage](/source/Macrophage).

- They are also essential to amoeboid-like locomotion. Human [mesenchymal stem cells](/source/Mesenchymal_stem_cell) are a good example of this function: these migratory cells are responsible for in-utero remodeling; for example, in the formation of the [trilaminar germ disc](/source/Trilaminar_germ_disc) during [gastrulation](/source/Gastrulation).[13]

## Morphology

The forms of pseudopodia, from left: polypodial and lobose; monopodial and lobose; filose; conical; reticulose; tapering actinopods; non-tapering actinopods

Pseudopods can be classified into several varieties according to the number of projections (monopodia and polypodia), and according to their appearance.

Some pseudopodial cells are able to use multiple types of pseudopodia depending on the situation. Most use a combination of [lamellipodia](/source/Lamellipodia) and [filopodia](/source/Filopodia) to migrate[14] (e.g. metastatic cancer cells).[15] Human [foreskin](/source/Foreskin) fibroblasts can either use lamellipodia- or lobopodia-based migration in a 3D matrix depending on the matrix elasticity.[16]

### Lamellipodia

Lamellipodia are broad and flat pseudopodia used in locomotion.[4] They are supported by microfilaments which form at the leading edge, creating a mesh-like internal network.[17]

### Filopodia

Filopodia (or filose pseudopods) are slender and filiform with pointed ends, consisting mainly of [ectoplasm](/source/Ectoplasm_(cell_biology)). These formations are supported by [microfilaments](/source/Microfilament) which, unlike the filaments of lamellipodia with their net-like actin, form loose bundles by [cross-linking](/source/Cross-link). This formation is partly due to bundling proteins such as [fimbrins](/source/Fimbrin) and [fascins](/source/Fascin).[17][18] Filopodia are observed in some animal cells: in part of [Filosa](/source/Filosa) ([Rhizaria](/source/Rhizaria)), in "[Testaceafilosia](/source/Testaceafilosia)", in [Vampyrellidae](/source/Vampyrellidae) and [Pseudosporida](/source/Pseudosporida) ([Rhizaria](/source/Rhizaria)) and in [Nucleariida](/source/Nucleariida) ([Opisthokonta](/source/Opisthokonta)).[4]

### Lobopodia

Lobopodia (or lobose pseudopods) are bulbous, short, and blunt in form.[19] These finger-like, tubular pseudopodia contain both [ectoplasm](/source/Ectoplasm_(cell_biology)) and [endoplasm](/source/Endoplasm). They can be found in different kind of cells, notably in [Lobosa](/source/Lobosa) and other [Amoebozoa](/source/Amoebozoa) and in some [Heterolobosea](/source/Heterolobosea) ([Excavata](/source/Excavata)).

High-pressure lobopodia can also be found in human [fibroblasts](/source/Fibroblast) travelling through a complex network of 3D [matrix](/source/Extracellular_matrix) (e.g. mammalian [dermis](/source/Dermis), cell-derived matrix). Contrarily to other pseudopodia using the pressure exerted by actin polymerization on the membrane to extend, fibroblast lobopods use the nuclear piston mechanism consisting in pulling the nucleus via actomyosin contractility to push the [cytoplasm](/source/Cytoplasm) that in turn push the membrane, leading to pseudopod formation. To occur, this lobopodia-based fibroblast migration needs [nesprin 3](/source/Nesprin), [integrins](/source/Integrin), [RhoA](/source/RHOA), [ROCK](/source/Rho-associated_protein_kinase) and [myosin II](/source/Myosin#Myosin_II). Otherwise, lobopods are often accompanied with small lateral [blebs](/source/Bleb_(cell_biology)) forming along the side of the cell, probably due to the high intracellular pressure during lobopodia formation increasing the frequency of plasma membrane-cortex rupture.[20][16][21]

### Reticulopodia

Reticulopodia (or reticulose pseudopods),[22] are complex formations in which individual pseudopods are merged and form irregular nets. The primary function of reticulopodia, also known as myxopodia, is food ingestion, with locomotion a secondary function. Reticulopods are typical of [Foraminifera](/source/Foraminifera), [Chlorarachnea](/source/Chlorarachnea), *[Gromia](/source/Gromia)* and *[Filoreta](/source/Filoreta)* (Rhizaria).[4]

If a pseudopod is branched as a reticulopod is but not forming a network, it is known as a rhizopod.[23]

### Axopodia

Axopodia (also known as actinopodia) are narrow pseudopodia containing complex arrays of [microtubules](/source/Microtubule) enveloped by cytoplasm. Axopodia are mostly responsible for phagocytosis by rapidly retracting in response to physical contact. These pseudopodia are primarily food-collecting structures, but also provide a means of hydrological transportation via the expansion of their surface areas. They are observed in "[Radiolaria](/source/Radiolaria)" and "[Heliozoa](/source/Heliozoa)".[4]

## References

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