Convergent extension (CE), sometimes called convergence and extension (C&E), is the process by which the tissue of an embryo is restructured to converge (narrow) along one axis and extend (elongate) along a perpendicular axis by cellular movement. An example of this process is where the

anteroposterior axis Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...

(the axis drawn between the head and tail end of an embryo) becomes longer as the lateral tissues (those that make up the left and right sides of the embryo) move in towards the dorsal midline (the middle of the back of the animal). This process plays a crucial role in shaping the body plan during

embryogenesis An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...

and occurs 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 multilayered structure known as the gastrula. Be ...

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 ...

, axis elongation, and

organogenesis Organogenesis is the phase of embryonic development that starts at the end of gastrulation and continues until birth. During organogenesis, the three germ layers formed from gastrulation (the ectoderm, endoderm, and mesoderm) form the internal orga ...

in both

vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...

and

invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

embryos. In

chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...

animals, this process is utilized within a vast population of cells; from the smaller populations in the

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consis ...

of the sea squirt (ascidian) to the larger populations of the dorsal

mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...

and neural

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...

of frogs (Xenopus) and fish. Many characteristics of convergent extension are conserved in the teleost fish, the bird, and very likely within mammals at the molecular, cellular, and tissue level. Convergent extension has been primarily studied in frogs and fish due to their large embryo size and their development outside of a maternal host (in egg clutches in the water, as opposed to in a uterus). Within frogs and fish, however, there exist fundamental differences in how convergent extension is achieved. Frog embryogenesis utilizes cell rearrangement as the sole player of this process. Fish, on the other hand, utilize both cell rearrangement as well as directed migration (Fig. 1) . Cellular rearrangement is the process by which individual cells of a tissue rearrange to reshape the tissue as a whole, while cellular migration is the directed movement of a singular cell or small group of cells across a substrate such as a membrane or tissue. Frog (Xenopus), as well as other amphibian,

serves as an excellent example of the role of convergent extension in embryogenesis. During gastrulation in frogs, the driving force of convergent extension is the morphogenic activity of the presumptive dorsal mesodermal cells; this activity is driven by the

mesenchymal Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...

cells that lie beneath the presumptive mesodermal and neural tissues. These tissues exist within the involuting marginal zone (IMZ) of the embryo which lies between the vegetal

endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...

and the posterior neural tissue. The IMZ is integral to gastrulation and R. Keller et al. eloquently exemplify the importance of convergent extension in Xenopus gastrulation. “…the IMZ, true to its name, involutes or rolls over the blastoporal lip and turns inside out. As it involutes, it converges along the mediolateral axis and extends along the future anterior-posterior axis of the notochordal and somitic mesoderm. Convergence and extension of these tissues squeezes the blastopore shut and simultaneously elongate the body axis. Elongation continues through the neurula and tailbud stages…As these involuted dorsal mesodermal tissues converge and extend on the inside of the gastrula, the presumptive posterior neural tissue converges and extends on the outside of the embryo, parallel to the underlying mesoderm, and then rolls up to form the neural tube, which later forms the hindbrain and spinal cord of the central nervous system". Should convergent extension be interrupted or incomplete, the resulting organism will have a short anteroposterior axis, wide notochord, and broad, open neural tube. The cellular signals required for convergent extension are not fully understood, however, it is known that the non-canonical

Wnt signaling pathway The Wnt signaling pathways are a group of signal transduction pathways which begin with proteins that pass signals into a cell through cell surface receptors. The name Wnt is a portmanteau created from the names Wingless and Int-1. Wnt signaling p ...

plays an important role,. Current research is shedding light on the cellular mechanisms of convergent extension and recently the planar cell polarity (PCP) pathway was implicated in regulating the cell polarity of the factors involved in convergent extension. This is an interesting development as the PCP pathway is an integral and well-studied pathway in flies, but has classically been thought to be unemployed by vertebrates. In addition to the non-canonical Wnt and PCP pathways involvement in convergent extension, the down-regulation of certain cell-cell adhesion molecules, such as C-cadherin and fibronectin/integrin interactions, may also play a role. Reduction of the activity of these cell-cell adhesion molecules allows for cells undergoing convergent extension to move more freely. Consistent with a role for a reduction of cell-cell adhesion in convergent extension, when cell-cell adhesion is not reduced, convergent extension cannot occur.

References

{{reflist Developmental biology