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Module 8: Cell Adhesion

Lecture 1: Cell Adhesion Molecules
Multicellularity: Stable Interactions between Cells
- Single fertilized egg  multicellular organisms
- Development requires
o Reiterated mitotic divisions
o Cell differentiation that defines tissue-specific patterns of gene
expression
o Signaling between cells
o Cells are able to associated and maintain connections
- Embryogenesis: formation of the inner cell mass, forming the embryo
o As the number of cells increase, they separate into one of three
germ layers
 Endoderm, ectoderm, mesoderm
o Cells must establish and maintain connections

Sponge Cells Sort and Re-associate
- Ability of cells to recognize and adhere to one other –
demonstrated by H.V Wilson
- Cells of two species of sponges – cells of each
organism were separated using a fine mesh and then
mixed together
- Cells from the same species recognized and
associated back together, cells from different
species did not associate
Embryonic Cell Association is Cell
Specific
- Johannes Holtfreter used
cells form a frog embryo
- Took cells from two different
germ layers during
development and separated individual cells
- Two different types of cells mixed together, cells from like tissues
recognized one another and associated – mimicked the organization seen
in the original embryo
- How do like cells recognize and contact each other during
embryogenesis? How do they stay together?
o Requires a collection of transmembrane proteins that span the cell –
cell adhesion molecules (CAMs)
o After aggregation, cells form specialized cell junctions that stabilize
cell-cell interactions and facilitate communication between adjacent
cells
Different Types of Junctions in Epithelial Cells

gap junctions (e) and tight junctions (a) Tight Junctions . preventing fluid movement across the layer of cells  Restricts diffusion of small molecules  Prevents leakage of digestive enzymes o Linear arrays of occuldin and claudin proteins are closely arranged between neighbouring cells o Not a single junction. Epithelial sheets in the intestine o Basal surface (bottom) is anchored to extracellular structures that provide an underlying structure that includes the basal lamina or basement membrane  Adhesion complexes: hemidesmosomes connects cells to the extracellular matrix o Apical surface (top) is organized into microvilli that face the interior of the intestine . Epithelial cells connect to one another along lateral surfaces to form epithelial sheets o Forms inner lining of digestive system and outer layers of skin . Tight junctions o completely prevents the diffusion of molecules in the extracellular space between the cells (small electron-dense molecule: lanthanum hydroxide)  diffusing in the extracellular space from the basal surface  epithelial sheet. Tight junctions (zonula occludens): connect adjacent epithelial cells just below the apical surface o Seals off space between cells. 4 types of adhesion complexes connect the lateral surfaces o Desmosomes (c). but an accumulation of structures that form a complete band between cells . adherens junctions (b). EM image: freeze fracturing: cells instantly frozen in liquid nitrogen and broken apart o Fracture occurs at points of weakness  lateral surface between neighbouring cells . limited at the tight junctions and can move no further Gap Junctions – Animal cells . TEM image: see where cells are “pinched” together . .

secondary messengers (cAMP. specialized cells (companion cells) are closely associated with their development and function  Companion cells provide ATP. and other substances  Connected to phloem by plasmodesmata o Spans the cell walls in addition to the cell membranes Plasmodesmata in Action . Gap junctions allow rapid coordination of cardiac muscle contraction and uterine muscle contractions . proteins. Plasmodesmata: structures in plant cells that have similar function to gap junctions o Important to the structure and function of phloem in flowering plants  Phloem: system of elongated tubes formed from linear arrays of connected cells. Phloem: circulatory system for the plant . Molecules that can diffuse through the gap junction o Ions. Carries nutrients (products of photosynthesis.5 – 2. . calcium) .Hormonal stimulation of one cell can lead to a response shared by many cells  diffusion of secondary messengers directly o Ex. Fluorescent molecule injected into a single cell – diffuses into neighbouring cells joined by gap junctions Plasmodesmata between Plant Cells . Directly links the cytosol of one cell to the cytosol of the adjacent link o Pinches and holds cell membranes together o Integrates metabolic activities of all cells in a tissue o Each channel is 1. allowing the free diffusion of up to about 1 kDa in size through the channels o 6 connexon protein subunits comprise a hexagonal connexion hemichannel in each cell membrane  Two hemichannels = gap junction channel Gap junctions in Action .0 nm in diameter.sucrose) from the leaves to the rest of the plant o Cells (sieve-tube elements) are connected by modified and enlarged plasmodesmata which form the sieve tube plate o Sieve tube elements are metabolically inactive.

Cadherins 2. Ig-superfamily 3. P-cadherin (placental) . Some are responsible for association of similar cells (hemophilic interactions) others connect different cells together (heterophilic interactions) Cassical Cadherins . E-cadherin (epithelial) 2. hemidesmosomes o Distinguished from other junctions by association with the cytoskeleton in particular actin filaments o Desmosomes: links two cells together o Hemidesmosomes: attach one cell to the extracellular matrix o Adherens junctions: connect the actin cytoskeleton between neighbouring cells Major Families of CAMs . Under standard cell culture conditions (with calcium) epithelial cells do not aggregate into sheets . Cell adhesion molecules make up adherens junctions fall into 4 families 1. o Transports sucrose from source cells to the rest of the plant. Adhesion is mediated by multiprotein complexes involving the transmembrane cadherin’s. Integrins 4. Critical cell adhesion molecules of the adherens junctions . small RNAs) o Viral pathogens exploit these channels for intercellular spread Anchoring Junctions . adherens junctions. but also cytosolic cofactors (catenin’s) that anchor cadherin to the actin skeleton . N-cadherin (neural) 3. just below the tight junctions . 3 major classes 1. gene transcripts. Ca2+ dependent CAMs hat mediate hemophilic interactions . Anchoring junctions: desmosomes. Mediate epithelial cell-cell adhesion near the apical surface of the cells. Selectins . as well as informational macromolecules (transcription factors.

o Introduce gene expressing E- cadherin  aggregation into epithelial-like clumps  E-cadherin mediates Ca2+ dependent adhesion of epithelial cells  Calcium  adhesion  No calcium  no adhesion Cadherins and Cell Differentiation . GFP: E-cadherin  introduced into cultured cells o Mixed together in a calcium- containing medium  aggregation of E-cadherin expressing cells o Accumulates at the surfaces of neighbouring cells  cells adhere only to other cells expressing the same cadherin = hemophilic interactions . E-cadherin: mediates tissue-specific adhesive connections in cultured epithelial cells .