Lamellipodia
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The lamellipodium (plural lamellipodia) (from
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''lamella'', related to ', "thin sheet", and the Greek radical ''pod-'', "foot") is a
cytoskeletal The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compo ...
protein actin projection on the leading edge of the cell. It contains a quasi-two-dimensional actin mesh; the whole structure propels the cell across a substrate. Within the lamellipodia are ribs of actin called microspikes, which, when they spread beyond the lamellipodium frontier, are called filopodia. The lamellipodium is born of actin nucleation in the plasma membrane of the cell and is the primary area of actin incorporation or
microfilament 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 ...
formation of the cell.


Description

Lamellipodia are found primarily in all mobile cells, such as the
keratinocytes Keratinocytes are the primary type of cell found in the epidermis, the outermost layer of the skin. In humans, they constitute 90% of epidermal skin cells. Basal cells in the basal layer (''stratum basale'') of the skin are sometimes referre ...
of fish and frogs, which are involved in the quick repair of wounds. The lamellipodia of these keratinocytes allow them to move at speeds of 10–20 μm / min over
epithelial Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellu ...
surfaces. When separated from the main part of a cell, a lamellipodium can still crawl about freely on its own. Lamellipodia are a characteristic feature at the front, leading edge, of motile cells. They are believed to be the actual motor which pulls the cell forward during the process of
cell migration Cell migration is a central process in the development and maintenance of multicellular organisms. Tissue formation during embryonic development, wound healing and immune responses all require the orchestrated movement of cells in particular dir ...
. The tip of the lamellipodium is the site where exocytosis occurs in migrating mammalian cells as part of their clathrin-mediated endocytic cycle. This, together with actin-polymerisation there, helps extend the lamella forward and thus advance the cell's front. It thus acts as a steering device for cells in the process of chemotaxis. It is also the site from which particles or aggregates attached to the cell surface migrate in a process known as cap formation. ;Structure Structurally, the barbed ends of the microfilaments (localized actin
monomer In chemistry, a monomer ( ; '' mono-'', "one" + '' -mer'', "part") is a molecule that can react together with other monomer molecules to form a larger polymer chain or three-dimensional network in a process called polymerization. Classification ...
s in an ATP-bound form) face the "seeking" edge of the cell, while the pointed ends (localized actin monomers in an
ADP Adp or ADP may refer to: Aviation * Aéroports de Paris, airport authority for the Parisian region in France * Aeropuertos del Perú, airport operator for airports in northern Peru * SLAF Anuradhapura, an airport in Sri Lanka * Ampara Airp ...
-bound form) face the lamella behind. This creates treadmilling throughout the lamellipodium, which aids in the retrograde flow of particles throughout. Arp2/3 complexes are present at microfilament-microfilament junctions in lamellipodia, and help create the actin meshwork. Arp 2/3 can only join onto previously existing microfilaments, but once bound it creates a site for the extension of ''new'' microfilaments, which creates branching. Another molecule that is often found in polymerizing actin with Arp2/3 is cortactin, which appears to link tyrosine kinase signalling to cytoskeletal reorganization in the lamellipodium and its associated structures. Rac and Cdc42 are two
Rho Rho (uppercase Ρ, lowercase ρ or ; el, ρο or el, ρω, label=none) is the 17th letter of the Greek alphabet. In the system of Greek numerals it has a value of 100. It is derived from Phoenician letter res . Its uppercase form uses the sa ...
-family GTPases which are normally cytosolic but can also be found in the cell membrane under certain conditions. When Cdc42 is activated, it can interact with Wiskott–Aldrich syndrome protein (WASp) family receptors, in particular
N-WASp Neural Wiskott-Aldrich syndrome protein is a protein that in humans is encoded by the ''WASL'' gene. The Wiskott-Aldrich syndrome (WAS) family of proteins share similar domain structure, and are involved in transduction of signals from receptor ...
, which then activates Arp2/3. This stimulates actin branching and increases cell
motility Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
. Rac1 induces cortactin to localize to the cell membrane, where it simultaneously binds F-actin and Arp2/3. The result is a structural reorganization of the lamellipodium and ensuing cell motility. Rac promotes lamellipodia while cdc42 promotes filopodia. Ena/VASP proteins are found at the leading edge of lamellipodia, where they promote actin polymerization necessary for lamellipodial protrusion and chemotaxis. Further, Ena/VASP prevents the action of capping protein, which halts actin polymerization.


References


External links


MBInfo - Lamellipodia

MBInfo - Lamellipodia Assembly

Video tour of cell motility
{{Authority control Cell movement Cytoskeleton Actin-based structures de:Lamellipodium