Cell junction

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Cell junction
Tight cell junction.png
Tight junction between two cells
Details
Identifiers
Latin junctiones cellulares
TH H1.00.01.0.00012
FMA 67394
Anatomical terminology

Cell junctions [1] or junctional complexes are a class of cellular structures consisting of multiprotein complexes that provide contact or adhesion between neighboring cells or between a cell and the extracellular matrix in animals. [2] They also maintain the paracellular barrier of epithelia and control paracellular transport. Cell junctions are especially abundant in epithelial tissues. Combined with cell adhesion molecules and extracellular matrix, cell junctions help hold animal cells together.

Contents

Cell junctions are also especially important in enabling communication between neighboring cells via specialized protein complexes called communicating (gap) junctions. Cell junctions are also important in reducing stress placed upon cells.

In plants, similar communication channels are known as plasmodesmata, and in fungi they are called septal pores. [3]

Types

Some examples of cell junctions Cell junction simplified en.svg
Some examples of cell junctions

In vertebrates, there are three major types of cell junction:

Invertebrates have several other types of specific junctions, for example septate junctions or the C. elegans apical junction. In multicellular plants, the structural functions of cell junctions are instead provided for by cell walls. The analogues of communicative cell junctions in plants are called plasmodesmata.

Anchoring junctions

Cells within tissues and organs must be anchored to one another and attached to components of the extracellular matrix. Cells have developed several types of junctional complexes to serve these functions, and in each case, anchoring proteins extend through the plasma membrane to link cytoskeletal proteins in one cell to cytoskeletal proteins in neighboring cells as well as to proteins in the extracellular matrix. [5]

Three types of anchoring junctions are observed, and differ from one another in the cytoskeletal protein anchor as well as the transmembrane linker protein that extends through the membrane:

JunctionCytoskeletal anchorTransmembrane linkerTies cell to:
Desmosomes Intermediate filaments Cadherin Other cells
Hemidesmosomes Intermediate filaments Integrins EC matrix
Adherens junctions (Adhesion belt, Focal adhesion) Actin filaments Cadherin / Integrin / Nectins Other cells / EC matrix

Anchoring-type junctions not only hold cells together but provide tissues with structural cohesion. These junctions are most abundant in tissues that are subject to constant mechanical stress such as skin and heart. [5]

Desmosomes

This image shows a desmosome junction between cells of the epidermal layer of the skin. Desmosome Cell Junction.png
This image shows a desmosome junction between cells of the epidermal layer of the skin.

Desmosomes, also termed as maculae adherentes, can be visualized as rivets through the plasma membrane of adjacent cells. Intermediate filaments composed of keratin or desmin are attached to membrane-associated attachment proteins that form a dense plaque on the cytoplasmic face of the membrane. Cadherin molecules form the actual anchor by attaching to the cytoplasmic plaque, extending through the membrane and binding strongly to cadherins coming through the membrane of the adjacent cell. [6]

Hemidesmosomes

Hemidesmosomes form rivet-like links between cytoskeleton and extracellular matrix components such as the basal laminae that underlie epithelia. Like desmosomes, they tie to intermediate filaments in the cytoplasm, but in contrast to desmosomes, their transmembrane anchors are integrins rather than cadherins. [7]

Adherens junctions

Adherens junctions share the characteristic of anchoring cells through their cytoplasmic actin filaments. Similarly to desmosomes and hemidesmosomes, their transmembrane anchors are composed of cadherins in those that anchor to other cells and integrins (focal adhesion) in those that anchor to extracellular matrix. There is considerable morphologic diversity among adherens junctions. Those that tie cells to one another are seen as isolated streaks or spots, or as bands that completely encircle the cell. The band-type of adherens junctions is associated with bundles of actin filaments that also encircle the cell just below the plasma membrane. Spot-like adherens junctions called focal adhesions help cells adhere to extracellular matrix. The cytoskeletal actin filaments that tie into adherens junctions are contractile proteins and in addition to providing an anchoring function, adherens junctions are thought to participate in folding and bending of epithelial cell sheets. Thinking of the bands of actin filaments as being similar to 'drawstrings' allows one to envision how contraction of the bands within a group of cells would distort the sheet into interesting patterns. [5]

Communicating (gap) junctions

Communicating junctions, or gap junctions allow for direct chemical communication between adjacent cellular cytoplasm through diffusion without contact with the extracellular fluid. [8] This is possible due to six connexin proteins interacting to form a cylinder with a pore in the centre called a connexon. [9] The connexon complexes stretches across the cell membrane and when two adjacent cell connexons interact, they form a complete gap junction channel. [8] [9] Connexon pores vary in size, polarity and therefore can be specific depending on the connexin proteins that constitute each individual connexon. [8] [9] Whilst variation in gap junction channels do occur, their structure remains relatively standard, and this interaction ensures efficient communication without the escape of molecules or ions to the extracellular fluid. [9]

Gap junctions play vital roles in the human body, [10] including their role in the uniform contractile of the heart muscle. [10] They are also relevant in signal transfers in the brain, and their absence shows a decreased cell density in the brain. [11] Retinal and skin cells are also dependent on gap junctions in cell differentiation and proliferation. [10] [11]

Tight junctions

Found in vertebrate epithelia, tight junctions act as barriers that regulate the movement of water and solutes between epithelial layers. Tight junctions are classified as a paracellular barrier which is defined as not having directional discrimination; however, movement of the solute is largely dependent upon size and charge. There is evidence to suggest that the structures in which solutes pass through are somewhat like pores.

Physiological pH plays a part in the selectivity of solutes passing through tight junctions with most tight junctions being slightly selective for cations. Tight junctions present in different types of epithelia are selective for solutes of differing size, charge, and polarity.

Proteins

There have been approximately 40 proteins identified to be involved in tight junctions. These proteins can be classified into four major categories;signalling proteins.

Roles
  • Scaffolding proteins – organise the transmembrane proteins, couple transmembrane proteins to other cytoplasmic proteins as well as to actin filaments.
  • Signaling proteins – involved in junctions assembly, barrier regulation, and gene transcription.
  • Regulation proteins – regulate membrane vesicle targeting.
  • Transmembrane proteins – including junctional adhesion molecule, occludin, and claudin.

It is believed that claudin is the protein molecule responsible for the selective permeability between epithelial layers.

A three-dimensional image is still yet to be achieved and as such specific information about the function of tight junctions is yet to be determined.

Tricellular junctions

Tricellular junctions seal epithelia at the corners of three cells. Due to the geometry of three-cell vertices, the sealing of the cells at these sites requires a specific junctional organization, different from those in bicellular junctions. In vertebrates, components tricellular junctions are tricellulin and lipolysis-stimulated lipoprotein receptors. In invertebrates, the components are gliotactin and anakonda. [12]

The cartoon of epithelium cells connected by tricellular junctions at the regions where three cells meet. Epithelium TCJ.png
The cartoon of epithelium cells connected by tricellular junctions at the regions where three cells meet.

Tricellular junctions are also implicated in the regulation of cytoskeletal organization and cell divisions. In particular they ensure that cells divide according to the Hertwig rule. In some Drosophila epithelia, during cell divisions tricellular junctions establish physical contact with spindle apparatus through astral microtubules. Tricellular junctions exert a pulling force on the spindle apparatus and serve as a geometrical clue to determine orientation of cell divisions. [13]

Cell junction molecules

The molecules responsible for creating cell junctions include various cell adhesion molecules. There are four main types: selectins, cadherins, integrins, and the immunoglobulin superfamily. [14]

Selectins are cell adhesion molecules that play an important role in the initiation of inflammatory processes. [15] The functional capacity of selectin is limited to leukocyte collaborations with vascular endothelium. There are three types of selectins found in humans; L-selectin, P-selectin and E-selectin. L-selectin deals with lymphocytes, monocytes and neutrophils, P-selectin deals with platelets and endothelium and E-selectin deals only with endothelium. They have extracellular regions made up of an amino-terminal lectin domain, attached to a carbohydrate ligand, growth factor-like domain, and short repeat units (numbered circles) that match the complementary binding protein domains. [16]

Cadherins are calcium-dependent adhesion molecules. Cadherins are extremely important in the process of morphogenesis – fetal development. Together with an alpha-beta catenin complex, the cadherin can bind to the microfilaments of the cytoskeleton of the cell. This allows for homophilic cell–cell adhesion. [17] The β-catenin–α-catenin linked complex at the adherens junctions allows for the formation of a dynamic link to the actin cytoskeleton. [18]

Integrins act as adhesion receptors, transporting signals across the plasma membrane in multiple directions. These molecules are an invaluable part of cellular communication, as a single ligand can be used for many integrins. Unfortunately, these molecules still have a long way to go in the ways of research. [19]

Immunoglobulin superfamily are a group of calcium independent proteins capable of homophilic and heterophilic adhesion. Homophilic adhesion involves the immunoglobulin-like domains on the cell surface binding to the immunoglobulin-like domains on an opposing cell's surface while heterophilic adhesion refers to the binding of the immunoglobulin-like domains to integrins and carbohydrates instead. [20]

Cell adhesion is a vital component of the body. Loss of this adhesion effects cell structure, cellular functioning and communication with other cells and the extracellular matrix and can lead to severe health issues and diseases.

Related Research Articles

<span class="mw-page-title-main">Epithelium</span> Tissue lining the surfaces of organs in animals

Epithelium or epithelial tissue is a thin, continuous, protective layer of compactly packed cells with little extracellular matrix. Epithelial tissues line the outer surfaces of organs and blood vessels throughout the body, as well as the inner surfaces of cavities in many internal organs. An example is the epidermis, the outermost layer of the skin. Epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. These tissues also lack blood or lymph supply. The tissue is supplied by nerves.

<span class="mw-page-title-main">Cell adhesion</span> Process of cell attachment

Cell adhesion is the process by which cells interact and attach to neighbouring cells through specialised molecules of the cell surface. This process can occur either through direct contact between cell surfaces such as cell junctions or indirect interaction, where cells attach to surrounding extracellular matrix, a gel-like structure containing molecules released by cells into spaces between them. Cells adhesion occurs from the interactions between cell-adhesion molecules (CAMs), transmembrane proteins located on the cell surface. Cell adhesion links cells in different ways and can be involved in signal transduction for cells to detect and respond to changes in the surroundings. Other cellular processes regulated by cell adhesion include cell migration and tissue development in multicellular organisms. Alterations in cell adhesion can disrupt important cellular processes and lead to a variety of diseases, including cancer and arthritis. Cell adhesion is also essential for infectious organisms, such as bacteria or viruses, to cause diseases.

<span class="mw-page-title-main">Desmosome</span> Cell junction involved in cell-to-cell adhesion

A desmosome, also known as a macula adherens, is a cell structure specialized for cell-to-cell adhesion. A type of junctional complex, they are localized spot-like adhesions randomly arranged on the lateral sides of plasma membranes. Desmosomes are one of the stronger cell-to-cell adhesion types and are found in tissue that experience intense mechanical stress, such as cardiac muscle tissue, bladder tissue, gastrointestinal mucosa, and epithelia.

<span class="mw-page-title-main">Cadherin</span> Calcium-dependent cell adhesion molecule

Cadherins (named for "calcium-dependent adhesion") are cell adhesion molecules important in forming adherens junctions that let cells adhere to each other. Cadherins are a class of type-1 transmembrane proteins, and they depend on calcium (Ca2+) ions to function, hence their name. Cell-cell adhesion is mediated by extracellular cadherin domains, whereas the intracellular cytoplasmic tail associates with numerous adaptors and signaling proteins, collectively referred to as the cadherin adhesome.

Cell adhesion molecules (CAMs) are a subset of cell surface proteins that are involved in the binding of cells with other cells or with the extracellular matrix (ECM), in a process called cell adhesion. In essence, CAMs help cells stick to each other and to their surroundings. CAMs are crucial components in maintaining tissue structure and function. In fully developed animals, these molecules play an integral role in generating force and movement and consequently ensuring that organs are able to execute their functions normally. In addition to serving as "molecular glue", CAMs play important roles in the cellular mechanisms of growth, contact inhibition, and apoptosis. Aberrant expression of CAMs may result in a wide range of pathologies, ranging from frostbite to cancer.

<span class="mw-page-title-main">Tight junction</span> Structure preventing inter-cell leakage

Tight junctions, also known as occluding junctions or zonulae occludentes, are multiprotein junctional complexes whose canonical function is to prevent leakage of solutes and water and seals between the epithelial cells. They also play a critical role maintaining the structure and permeability of endothelial cells. Tight junctions may also serve as leaky pathways by forming selective channels for small cations, anions, or water. The corresponding junctions that occur in invertebrates are septate junctions.

<span class="mw-page-title-main">Vinculin</span> Mammalian protein found in Homo sapiens

In mammalian cells, vinculin is a membrane-cytoskeletal protein in focal adhesion plaques that is involved in linkage of integrin adhesion molecules to the actin cytoskeleton. Vinculin is a cytoskeletal protein associated with cell-cell and cell-matrix junctions, where it is thought to function as one of several interacting proteins involved in anchoring F-actin to the membrane.

<span class="mw-page-title-main">Adherens junction</span> Protein complexes at cell junctions

In cell biology, adherens junctions are protein complexes that occur at cell–cell junctions and cell–matrix junctions in epithelial and endothelial tissues, usually more basal than tight junctions. An adherens junction is defined as a cell junction whose cytoplasmic face is linked to the actin cytoskeleton. They can appear as bands encircling the cell or as spots of attachment to the extracellular matrix.

<span class="mw-page-title-main">Juxtacrine signalling</span> Contact-based cell-cell signalling

In biology, juxtracrine signalling is a type of cell–cell or cell–extracellular matrix signalling in multicellular organisms that requires close contact. In this type of signalling, a ligand on one surface binds to a receptor on another adjacent surface. Hence, this stands in contrast to releasing a signaling molecule by diffusion into extracellular space, the use of long-range conduits like membrane nanotubes and cytonemes or the use of extracellular vesicles like exosomes or microvesicles. There are three types of juxtracrine signaling:

  1. A membrane-bound ligand and a membrane protein of two adjacent cells interact.
  2. A communicating junction links the intracellular compartments of two adjacent cells, allowing transit of relatively small molecules.
  3. An extracellular matrix glycoprotein and a membrane protein interact.
α-Catenin Primary protein link between cadherins and the actin cytoskeleton

α-Catenin (alpha-catenin) functions as the primary protein link between cadherins and the actin cytoskeleton. It has been reported that the actin binding proteins vinculin and α-actinin can bind to alpha-catenin. It has been suggested that alpha-catenin does not bind with high affinity to both actin filaments and the E-cadherin-beta-catenin complex at the same time. It has been observed that when α-catenin is not in a molecular complex with β-catenin, it dimerizes and functions to regulate actin filament assembly, possibly by competing with Arp2/3 protein. α-Catenin exhibits significant protein dynamics. However, a protein complex including a cadherin, actin, β-catenin and α-catenin has not been isolated.

<span class="mw-page-title-main">Tight junction protein 1</span> Protein found in humans

Zonula occludens-1 ZO-1, also known as Tight junction protein-1 is a 220-kD peripheral membrane protein that is encoded by the TJP1 gene in humans. It belongs to the family of zonula occludens proteins, which are tight junction-associated proteins and of which, ZO-1 is the first to be cloned. It was first isolated in 1986 by Stevenson and Goodenough using a monoclonal antibody raised in rodent liver to recognise a 225-kD polypeptide in whole liver homogenates and in tight junction-enriched membrane fractions. It has a role as a scaffold protein which cross-links and anchors Tight Junction (TJ) strand proteins, which are fibril-like structures within the lipid bilayer, to the actin cytoskeleton.

<span class="mw-page-title-main">Epithelial cell adhesion molecule</span> Transmembrane glycoprotein

Epithelial cell adhesion molecule (EpCAM), also known as CD326 among other names, is a transmembrane glycoprotein mediating Ca2+-independent homotypic cell–cell adhesion in epithelia. EpCAM is also involved in cell signaling, migration, proliferation, and differentiation. Additionally, EpCAM has oncogenic potential via its capacity to upregulate c-myc, e-fabp, and cyclins A & E. Since EpCAM is expressed exclusively in epithelia and epithelial-derived neoplasms, EpCAM can be used as diagnostic marker for various cancers. It appears to play a role in tumorigenesis and metastasis of carcinomas, so it can also act as a potential prognostic marker and as a potential target for immunotherapeutic strategies.

<span class="mw-page-title-main">Stress fiber</span> Contractile actin bundles found in non-muscle cells

Stress fibers are contractile actin bundles found in non-muscle cells. They are composed of actin (microfilaments) and non-muscle myosin II (NMMII), and also contain various crosslinking proteins, such as α-actinin, to form a highly regulated actomyosin structure within non-muscle cells. Stress fibers have been shown to play an important role in cellular contractility, providing force for a number of functions such as cell adhesion, migration and morphogenesis.

Desmocollins are a subfamily of desmosomal cadherins, the transmembrane constituents of desmosomes. They are co-expressed with desmogleins to link adjacent cells by extracellular adhesion. There are seven desmosomal cadherins in humans, three desmocollins and four desmogleins. Desmosomal cadherins allow desmosomes to contribute to the integrity of tissue structure in multicellular living organisms.

Epithelial polarity is one example of the cell polarity that is a fundamental feature of many types of cells. Epithelial cells feature distinct 'apical', 'lateral' and 'basal' plasma membrane domains. Epithelial cells connect to one another via their lateral membranes to form epithelial sheets that line cavities and surfaces throughout the animal body. Each plasma membrane domain has a distinct protein composition, giving them distinct properties and allowing directional transport of molecules across the epithelial sheet. How epithelial cells generate and maintain polarity remains unclear, but certain molecules have been found to play a key role.

Cell–cell interaction refers to the direct interactions between cell surfaces that play a crucial role in the development and function of multicellular organisms. These interactions allow cells to communicate with each other in response to changes in their microenvironment. This ability to send and receive signals is essential for the survival of the cell. Interactions between cells can be stable such as those made through cell junctions. These junctions are involved in the communication and organization of cells within a particular tissue. Others are transient or temporary such as those between cells of the immune system or the interactions involved in tissue inflammation. These types of intercellular interactions are distinguished from other types such as those between cells and the extracellular matrix. The loss of communication between cells can result in uncontrollable cell growth and cancer.

<span class="mw-page-title-main">Role of cell adhesions in neural development</span>

Cellular adhesions can be defined as proteins or protein aggregates that form mechanical and chemical linkages between the intracellular and extracellular space. Adhesions serve several critical processes including cell migration, signal transduction, tissue development and repair. Due to this functionality, adhesions and adhesion molecules have been a topic of study within the scientific community. Specifically, it has been found that adhesions are involved in tissue development, plasticity, and memory formation within the central nervous system (CNS), and may prove vital in the generation of CNS-specific therapeutics.

<span class="mw-page-title-main">Synaptic stabilization</span> Modifying synaptic strength via cell adhesion molecules

Synaptic stabilization is crucial in the developing and adult nervous systems and is considered a result of the late phase of long-term potentiation (LTP). The mechanism involves strengthening and maintaining active synapses through increased expression of cytoskeletal and extracellular matrix elements and postsynaptic scaffold proteins, while pruning less active ones. For example, cell adhesion molecules (CAMs) play a large role in synaptic maintenance and stabilization. Gerald Edelman discovered CAMs and studied their function during development, which showed CAMs are required for cell migration and the formation of the entire nervous system. In the adult nervous system, CAMs play an integral role in synaptic plasticity relating to learning and memory.

Tight junction proteins are molecules situated at the tight junctions of epithelial, endothelial and myelinated cells. This multiprotein junctional complex has a regulatory function in passage of ions, water and solutes through the paracellular pathway. It can also coordinate the motion of lipids and proteins between the apical and basolateral surfaces of the plasma membrane. Thereby tight junction conducts signaling molecules, that influence the differentiation, proliferation and polarity of cells. So tight junction plays a key role in maintenance of osmotic balance and trans-cellular transport of tissue specific molecules. Nowadays is known more than 40 different proteins, that are involved in these selective TJ channels.

<span class="mw-page-title-main">Intercellular communication</span>

Intercellular communication (ICC) refers to the various ways and structures that biological cells use to communicate with each other directly or through their environment. Often the environment has been thought of as the extracellular spaces within an animal. More broadly cells may also communicate with other animals, either of their own group or species, or other species in the wider ecosystem. Different types of cells use different proteins and mechanisms to communicate with one another using extracellular signalling molecules or electric fluctuations which could be likened to an intercellular ethernet. Components of each type of intercellular communication may be involved in more than one type of communication making attempts at clearly separating the types of communication listed somewhat futile. Broadly speaking, intercellular communication may be categorized as being within a single animal, or between an animal and other animals in the ecosystem in which it lives. In this article intercellular communication has been further collated into various areas of research rather than by functional or structural characteristics.

References

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