morphogenesis Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of deve ...

, apical constriction is the process in which contraction of the apical side of a cell causes the cell to take on a wedged shape. Generally, this shape change is coordinated across many cells of an

epithelial Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of man ...

layer, generating forces that can bend or fold the cell sheet.

Morphogenetic role

Apical constriction plays a central role in important morphogenetic events in both

invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...

and

vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

. It is typically the first step in any invagination process and is also important in folding tissues at specified hingepoints. During

gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...

in both invertebrates and vertebrates, apical constriction of a ring of cells leads to blastopore formation. These cells are known as bottle cells, for their eventual shape. Because all of the cells constrict on the apical side, the epithelial sheet bends convexly on the basal side. In vertebrates, apical constriction plays a role in a range of other morphogenetic processes such

neurulation Neurulation refers to the folding process in vertebrate embryos, which includes the transformation of the neural plate into the neural tube. The embryo at this stage is termed the neurula. The process begins when the notochord induces the formati ...

placode In embryology, a neurogenic placode is an area of thickening of the epithelium in the embryonic head ectoderm layer that gives rise to neurons and other structures of the sensory nervous system. Placodes are embryonic structures that give rise t ...

formation, and

primitive streak The primitive streak is a structure that forms in the early embryo in amniotes. In amphibians, the equivalent structure is the blastopore. During early embryonic development, the embryonic disc becomes oval shaped, and then pear-shaped with the ...

formation.

Mechanism

Apical constriction occurs primarily through the contraction of

cytoskeletal The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all Cell (biology), cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane ...

elements. The specific mechanism depends on the species, the cell type, and the morphogenetic movement.

Model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s that have been studied include the

frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...

Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos'' = strange, πους, ''pous'' = foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described with ...

'', and the fly ''

Drosophila ''Drosophila'' (), from Ancient Greek δρόσος (''drósos''), meaning "dew", and φίλος (''phílos''), meaning "loving", is a genus of fly, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or p ...

''.

''Xenopus''

During ''

, bottle cells are located in the dorsal marginal zone and apically constrict inwards to initiate

involution Involution may refer to: Mathematics * Involution (mathematics), a function that is its own inverse * Involution algebra, a *-algebra: a type of algebraic structure * Involute, a construction in the differential geometry of curves * Exponentiati ...

of the blastopore. In these cells, apical constriction occurs when

actomyosin Myofilaments are the three protein filaments of myofibrils in muscle cells. The main proteins involved are myosin, actin, and titin. Myosin and actin are the ''contractile proteins'' and titin is an elastic protein. The myofilaments act toget ...

contractility folds the

cell membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...

to reduce the apical surface area.

Endocytosis Endocytosis is a cellular process in which Chemical substance, substances are brought into the cell. The material to be internalized is surrounded by an area of cell membrane, which then buds off inside the cell to form a Vesicle (biology and chem ...

of the membrane at the apical side further reduces surface area. Active trafficking of these endocytosed vesicles along

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 nanometer, nm and have an inner diameter bet ...

tracks is also believed to be important, since the depolymerization (but not stabilization) of microtubules reduces the extent of apical constriction. Although apical constriction is always observed, it is not necessary for gastrulation, indicating that there are other morphogenetic forces working in parallel. Researchers have shown that the removal of bottle cells does not inhibit gastrulation, but simply makes it less efficient. Bottle cell removal does, however, result in deformed embryos.

Neural tube In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural folds become elevated, ...

cells in ''Xenopus'' apically constrict during the initial invagination as well as during hingepoint folding. Here, the mechanism depends upon the protein Shroom3, which is sufficient to drive apical constriction. Because Shroom3 is an actin-binding protein and accumulates on the apical side, the most likely mechanism is that Shroom3 aggregates the

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 ...

meshwork, generating a squeezing force. Ectopic Shroom3 has been shown to be sufficient to induce apical constriction, but only in cells with apico-basal polarity.

''Drosophila''

The molecular picture of apical constriction is most complete for ''

''. During ''Drosophila'' gastrulation, apical constriction of midline cells initiates invagination to create the ventral furrow. Like in ''Xenopus'', actomyosin contractility plays a major role in constricting the apical side of the cell. The constricting cells have an actin meshwork directly beneath the apical membrane as well as circumferential actin belts lining the adherens junctions between cells. Pulsed contractions of the actin meshwork are believed to be primarily responsible for reducing the apical surface area. In ''Drosophila'', researchers have also pinpointed the molecules responsible for ''coordinating'' apical constriction in time. ''Protein folded gastrulation'' (Fog), a secreted protein and ''Concertina'', a G alpha protein, are members of the same pathway that ensure that apical constriction is initiated in the right cells at the right time. The transmembrane protein T48 is part of a redundant pathway that is also needed for coordination of apical constriction. Both pathways must be disrupted in order to completely block ventral furrow formation. Both pathways also regulate the localization of RhoGEF2, a member of the Rho family GTPases, which are known to regulate actin dynamics.

References

External links

* http://worms.zoology.wisc.edu/urchins/SUgast_primary4.html * http://www.sdbonline.org/fly/newgene/foldgs1.htm {{DEFAULTSORT:Apical Constriction Cell biology