Tissues - classsification, general structure and function Epithelial tissue Institute of Histology and Embryology Doc. MUDr. Zuzana Jirsová, CSc. Histology and Embryology – B81131 Lecture - 16. 10. 2013 TISSUES Aggregates of cells organized to perform one or more specific functions Common origin and morphological characteristic Four basic types of tissues: EPITHELIAL TISSUE CONNECTIVE TISSUE MUSCLE TISSUE NERVE TISSUE EPITHELIAL TISSUE Origin: tissue developing from the ectoderm, mesoderm and endoderm (all three germ layers ) Structure: tissue composed of closely aggregated cells Function: protection, secretion, absorption, transport, respiration, reception CONNECTIVE TISSUE Origin: tissue develops from mesenchyme Structure: CELLS EXTRACELLULAR MATRIX protein fibers and ground substance Types: connective tissue proper (loose and dense) CT with special properties (mucous, reticular, elastic and adipose) supporting CT (cartilage, bone, dentin) Functional specialization: supporting, structural, storage, defense, nutrition MUSCLE TISSUE Origin: mesoderm (skeletal and cardiac muscles) mesenchyme (smooth muscles) Functional property: contractibility. Actin and myosin are organized into the myofilaments, actin–myosin interaction is responsible for contraction. Two types: striated muscles - skeletal and cardiac muscles smooth muscles Structural unit: cell (cardiac and smooth muscles) muscle fiber (skeletal muscle) NERVE TISSUE Origin: tissue develops from neuroectoderm Structure: NT consists of two principal types of cells: neurons (nerve cells) supporting glial cells (neuroglia) Function Neurons: functional units of nervous system, gather and process information and generate signal, conduct electrical impulses Glial cells: protection, electric insulation, metabolic exchange, defense EPITHELIAL TISSUE Cell adhesion and intercellular junctions Cell surface modifications Cell polarity Classification of the epithelial tissue EPITHELIAL TISSUE Cells are closely apposed and adhere to one another by means of specific cell adhesion molecules (CAMs) Specialized cell junctions: OCCLUDING JUNCTIONS - link forming an impermeable barrier: zonula occludens, tight junction ANCHORING JUNCTIONS - provide mechanical stability by linking of the cytoskeleton of one cell to the cytoskeleton of adjacent cell or cell-to-extracellular matrix (*) zonula adherens (focal contacts*) - actin filaments macula adherens/desmosome (hemidesmosome*) - intermediate filaments COMMUNICATING JUNCTIONS: gap junctions – allow selective diffusion of molecules between adjacent cells CELL POLARITY Free surface/ apical domain Special structural surface modifications: Microvilli – cytoplasmic processes (core consisting of actin filaments) closed packed microvilli – brush border (absorptive epithelium) Stereocilia – long immotile microvilli: hair cells of the inner ear, epithelium of the male genital ducts Cilia – motile cytoplasmic processes (consisting of microtubule axoneme, basal bodies); epithelium of respiratory ways, and uterine tube CELL POLARITY Lateral domain In close contact with the opposed lateral domains of neighboring cells. Cell adhesion molecules: CADHERINS Cell-to-cell junctions: (a) occluding, (b) adhesive or anchoring, (c) and communicating Folding - invagination and evagination of lateral cell membrane, interdigitation Basal domain Attached to the basement membrane, cell adhesion molecules: INTEGRINS Cell-to-extracellular matrix junctions: hemidesmosomes and focal contacts Infoldings of basal cell membrane: ion – transporting epithelium (striated ducts of salivary glands, kidney tubules) BASEMENT MEMBRANE (term used in light microscopy) Basement membrane mediates attachment of epithelial cells to underlying connective tissue Basal lamina (in electron microscope) Lamina rara is an electron-lucent area (40 nm thick) between the lamina densa and epithelium; contains of extracellular portions of intergrins (CAM), mainly their fibronectin and laminin receptors Lamina densa is an electron-dense layer of 40 – 60 nm thick between the epithelium and the adjacent connective tissue composed of laminins, type IV collagen, proteoglycans (heparan sulfate and chondroitin sulfate proteoglycans), entactin BASEMENT MEMBRANE Attachment of the basal lamina to the underlying connective tissues mediated by anchoring fibrils (type VII collagen) and fibrillin microfibrils A layer of reticular fibers underlies the basal lamina reticular lamina (is not produced by the epithelium) belongs to the connective tissue Basement membrane is PAS positive and stains with acidic dyes Zonula occludens CELL JUNCTIONS Zonula adherens Free surface Apical domain Junctional complex Terminal bars in LM Scheme: Ross, Pawlina, Histology, 2003 Lateral domain Focal contacts Hemidesmosomes Desmosomes cell membranes b Scheme, electron micrographs: Ross, Pawlina, Histology, 2003 ZONULA OCCLUDENS Tight junction, that separates luminal space from the intercellular compartment. Focal junctions between cells by transmembrane proteins (claudin, occludin) form sealing strands which are arranged as intertwining lines. Electron micrograph of ZO (a) - close apposition of adjacent areas of cell membrane Freeze fracture technique (b) shows ZO as anastomosing network of ridges – arrows ZONULA ADHERENS (Scheme, electron micrograph: Ross, Pawlina, Histology, 2003) ZA is belt-like junction encircling the cell localized just bellow ZO. Actin filaments of terminal web of adjacent cells are attached to E cadherin-catenin complex by alfa-actinin and vinculin. Intercellular space measures 10-20 nm. DESMOSOM, MACULA ADHAERENS Disc-like intercellular attachment. Intermediate filaments (CF) are anchored to the attachment plaque (P) located on the cytoplasmatic side of cell membrane (CM). CM Cytokeratin filaments P Plaque: desmoplakins desmoglobin Transmembrane proteins: desmogleins X = intercellular space (20-30 nm) desmocollins Scheme, elektron micrograph: Histology, Ross, Pawlina, 2006 GAP JUNCTION, NEXUS – permit the direct passage of the ions, signaling molecules nexus Gap junction proteins (connexins) form hexamers CONNEXONS with hydrophilic pore (2 nm in diameter). Connexons in adjacent cell membranes are aligned to form a cylindrical transmembrane channel. Conformational changes in connexins lead to opening or closing of channels. Epithelium, smooth and cardiac muscles. CM CM 1: Electron micrograph (TEM): close apposition of the cell membranes in the gap junction (IS is only 2 nm) CM – cell membrane IS – intercellular space Ross, Pawlina, Histology, 2006 CM c 1 Cx 2: Freeze fracture technique: dense particles correspond to connexons Junquiera´s Basic Histology, Mescher, 2010 A 3: Scheme of a gap junction Ross, Pawlina, Histology, 2006 A – closed, B – open B 2 3 IS Diameter of connecting channel is regulated by reversible changes in the conformation of the individual connexins. Organization of actin filament in microvillus Brush (striated) border of the absorptive epithelium Microvillus is about 1µm heigh and 0.08 µm wide villin espin fimbrin Attachment of AF to the CM AF fascin Freeze fracture technique myosin I Bundles of AF constituting the core of microvillus spectrin AF rootlets terminal web (actin cortex) myosin II Spectrin binds AF to the actin cortex and to CM Cytokeratin intermediate filaments cytokeratin IMF Scheme: Histology, Ross, Pawlina, 2011 EM, Freeze fracture technique showing the arrangement of AF in the microvilli. Ross, Pawlina, Histology, 2006 STEREOCILIA - long immotile microvilli Epididymis, pseudostratified columnar epithelium - sterocilia (arrows) ezrin fimbrin actin filaments espin cytoplasmic bridges – α-actinin actin filaments α-actinin Scheme: Histology, Ross, Pawlina, 2011 Photomicrograph: W. Kühnel, Atlas, 2003 CILIA (lenght: 5-10 µm, diameter: 0,2 µm) Cilia are elongated, highly motile structures on the surface of some epithelial cells, inserted into basal bodies (BB). BB have a structure similar to centrioles and MTOC function. Cilium is composed of microtubules which form axoneme: 9 peripheral dublets (A microtubulus consists of 13 protofibrils, B – 10 protofibrils) and one central pair of microtubules. The motion of the cilia occurs due to activity of dynein (protein with ATP-ase activity), with ATP as the energy source. Occurrence: epithelium lining the respiratory tract and uterine tube B A Scheme, electron micrographs: Stevens, Lowe, Histology, 1993 A cross (b) and longitudival (c) section of cilia in EM CM = cytoplasmic membrane, BB = basal bodies, Cy = cytoplasm Basal surface modification: ion-transporting epithelium (proximal and distal tubules of the kidney, striated ducts of the salivary glands). Infoldings at the basal cell membrane enlarge basal surface, mitochondria provide energy for active transport of ions. microvilli tight junction G = Golgi complex tight junction longitudinally oriented elongated mitochondria infoldings of the basal cell membrane Na+K+ ATP-ase basement membrane Basal labyrinth Sodium pump Scheme: Krstic, Illustrated Encycl. of Human Histology, 1984 Classification of epithelium according arrangement of cells: a) PLANE: cells form tightly cohesive sheets traditional nomenclature is based on the: shape of cells: squamous, cuboidal, columnar number of layers: simple, stratified, pseudostratified b) TRABECULAR: cells are arranged in cords or plates (liver, endocrine glands) c) RETICULAR: cells form three-dimensional network (stroma of thymus, epithelium of crypts in tonsils, stellate reticulum of enamel organ) Classification of epithelium according functional specialization Covering epithelium Simple squamous, cuboidal, columnar, pseudostratified columnar Stratified squamous, columnar Transitional epithelium (urothelium) Secretory epithelium – exocrine and endocrine; polarization of cells mechanism of secretion: merocrine, apocrine, holocrine Absorptive epithelium Respiratory epithelium: exchange of respiratory gases, in the lung alveoli Sensory epithelium: cells react on the external stimuli by change of membrane potential Primary and secondary sensory cells Myoepithelium: cells with ability to contract Germinal epithelium - production of cells (seminiferous epithelium of testis produce spermatozoa) Ion-transporting epithelium (modification of the basal cell surface) REQUESTED KNOWLEDGE Epithelial tissue, overview of the structure and function, classification Secretory epithelium (endocrine, exocrine – types of secretion) Sensory epithelium - structure and function. Myoepithelium