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Actin Dynamics and Remodeling of Cell-Cell Junctions in Epithelial Morphogenesis pp. 1-59 $100.00
Authors:  (Marc D.H. Hansen, Michael R. Stark, Department of Physiology and Developmental Biology, Brigham Young University, Provo, Utah, USA)
Cell-cell adhesion systems play a critical role in integrating individual cells into
tissues. Crucial to cell-cell adhesion is the structure and dynamics of the actin
cytoskeleton, particularly at cell-cell junctions. Such organization forms the basis of
proper cell and tissue morphology. Control of actin dynamics at adhesion sites allows
epithelial tissues to undergo dramatic remodeling events. These epithelial morphogenesis
events occur throughout development. The staggering variety of cell and tissue
architecture changes during epithelial morphogenesis has resulted in events often being
categorized by specific types of cellular rearrangements, with little regard to
understanding epithelial morphogenesis in a unified manner. All morphogenetic
rearrangements rely on movements and shape changes made by individual cells within
the tissue. When these movement and shape changes occur collectively, they result in
repositioning of individual cells within the tissue or an overall deformation of the tissue.
The unfiying feature of such events is that cell-cell adhesion are maintained, allowing
individual cell morphology changes to alter the overall tissue architecture. Thus, actin
dynamics in individual cells are coordinated thorough cell-cell contacts to drive changes
in morphology at the tissue level. Epithelial morphogenesis also includes tissue
remodeling events where cell shape changes and movement occur in isolation and do not
impact the entire tissue, allowing cells to behave in a solitary fashion. An example is
epithelial-mesenchymal transition (EMT), where cells completely detach from the
epithelial tissue and migrate to new locations. Importantly, epithelial morphogenesis
events that result in individual cells detaching from the tissue highlight how cells can
uncouple actin dynamics from cell-cell adhesion systems to generate completely different
outcomes on tissue architecture. Given the central role of actin-membrane connections at
adhesion sites in determining whether actin dynamics of individual cells occur
collectively or independently, it is important that the molecular basis of membrane
connections at adhesion sites has been under recently renewed scrutiny. The prevailing
dogma that cadherin-based adhesions are directly anchored to actin filaments, thus
providing the majority of functional actin-membrane linkages at adhesion sites, has been
challenged and the precise molecular organization of actin-membrane connections of
cell-cell adhesions is under active revision. Actin-membrane anchor sites at epithelial
cell-cell junctions are typically symmetrical, with actin membrane linkages occurring on
both sides of the adhesion site. A benefit of such organization it that symmetrical
connection points allow the actin cytoskeleton of individual cells to be integrated into a
larger multi-cellular actin network, connected through points of cell-cell adhesion. This
network can be regulated at the level of the whole tissue to drive collective epithelial
morphogenesis events. From the perspective of actin dynamics and actin-membrane
connections, the entire variety of epithelial morphogenesis events can be viewed as a
spectrum of outcomes that result from changes in actin organization and contractility, as
well as whether these changes are coordinated through actin-membrane connections with
cell-cell adhesion systems. In this view, the extent, order, and location of changes in the
actin cytoskeleton in each cell constrain the outcome of epithelial morphogenesis in terms
of tissue architecture. A detailed understanding of how actin reorganization and
contractility are coordinated through cell-cell contacts of epithelial cells will be critical in
developing such a unified view of epithelial morphogenesis events. A major goal of cell
biologists has been to characterize actin dynamics in live cells and identify how specific
molecular machines generate those actin dynamic events. A major emphasis of
developmental biologists has been to understand how signaling networks are regulated in
time and space to drive specific developmental events. This chapter will focus on how
spatiotemporal signaling and actin rearrangements of cell-cell adhesions interface during
development to generate specific changes in tissue morphology. 

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Actin Dynamics and Remodeling of Cell-Cell Junctions in Epithelial Morphogenesis pp. 1-59