973:. Tight junctions are a pair of trans-membrane protein fused on outer plasma membrane. Adherens junctions are a plaque (protein layer on the inside plasma membrane) which attaches both cells' microfilaments. Desmosomes attach to the microfilaments of cytoskeleton made up of keratin protein. Hemidesmosomes resemble desmosomes on a section. They are made up of the integrin (a transmembrane protein) instead of cadherin. They attach the epithelial cell to the basement membrane. Gap junctions connect the cytoplasm of two cells and are made up of proteins called
1069:
630:
1394:
1382:
1370:
451:, Latin for "scale" – as on fish or snake skin. The cells fit closely together in tissues, providing a smooth, low-friction surface over which fluids can move easily. The shape of the nucleus usually corresponds to the cell form and helps to identify the type of epithelium. Squamous cells tend to have horizontally flattened, nearly oval-shaped nuclei because of the thin, flattened form of the cell. Squamous epithelium is found lining surfaces such as
767:
3719:
1358:
1077:
38:
649:, and both types of layering can be made up of any of the cell shapes. However, when taller simple columnar epithelial cells are viewed in cross section showing several nuclei appearing at different heights, they can be confused with stratified epithelia. This kind of epithelium is therefore described as
1060:. Here, cells fated for elimination are seamlessly squeezed out by contracting a band of actin and myosin around and below the cell, preventing any gaps from forming that could disrupt their barriers. Failure to do so can result in aggressive tumors and their invasion by aberrant basal cell extrusion.
1055:
Epithelia turn over at some of the fastest rates in the body. For epithelial layers to maintain constant cell numbers essential to their functions, the number of cells that divide must match those that die. They do this mechanically. If there are too few of the cells, the stretch that they experience
753:
to punakoidal. They are tightly packed and form a continuous sheet with almost no intercellular spaces. All epithelia is usually separated from underlying tissues by an extracellular fibrous basement membrane. The lining of the mouth, lung alveoli and kidney tubules are all made of epithelial tissue.
667:
Stratified or compound epithelium differs from simple epithelium in that it is multilayered. It is therefore found where body linings have to withstand mechanical or chemical insult such that layers can be abraded and lost without exposing subepithelial layers. Cells flatten as the layers become more
508:
that lines the walls of the seminferous tubules in the testes are also of the cuboidal type. Cuboidal cells provide protection and may be active in pumping material in or out of the lumen, or passive depending on their location and specialisation. Simple cuboidal epithelium commonly differentiates to
1283:
Epithelium grown in culture can be identified by examining its morphological characteristics. Epithelial cells tend to cluster together, and have a "characteristic tight pavement-like appearance". But this is not always the case, such as when the cells are derived from a tumor. In these cases, it is
1220:
of the extracellular environment by playing "a sensory role mediating specific signalling cues, including soluble factors in the external cell environment, a secretory role in which a soluble protein is released to have an effect downstream of the fluid flow, and mediation of fluid flow if the cilia
355:
There are three principal shapes of epithelial cell: squamous (scaly), columnar, and cuboidal. These can be arranged in a singular layer of cells as simple epithelium, either simple squamous, simple columnar, or simple cuboidal, or in layers of two or more cells deep as stratified (layered), or
690:
In this particular case, the most apical layers (exterior) of cells are dead and lose their nucleus and cytoplasm, instead contain a tough, resistant protein called keratin. This specialization makes the epithelium somewhat water-resistant, so is found in the mammalian skin. The lining of the
578:
are modified columnar cells and are found between the columnar epithelial cells of the duodenum. They secrete mucus, which acts as a lubricant. Single-layered non-ciliated columnar epithelium tends to indicate an absorptive function. Stratified columnar epithelium is rare but is found in
529:
Columnar epithelial cells are elongated and column-shaped and have a height of at least four times their width. Their nuclei are elongated and are usually located near the base of the cells. Columnar epithelium forms the lining of the stomach and intestines. The cells here may possess
619:, the wafting effect produced causes mucus secreted locally by the goblet cells (to lubricate and to trap pathogens and particles) to flow in that direction (typically out of the body). Ciliated epithelium is found in the airways (nose, bronchi), but is also found in the uterus and
1256:
1242:
with hematopoietic immune cells. The resulting immune functions of these non-hematopoietic, structural cells contribute to the mammalian immune system ("structural immunity"). Relevant aspects of the epithelial cell response to infections are encoded in the
2758:
717:
Transitional epithelia are found in tissues that stretch, and it can appear to be stratified cuboidal when the tissue is relaxed, or stratified squamous when the organ is distended and the tissue stretches. It is sometimes called
915:
and must be nourished by substances diffusing from the blood vessels in the underlying tissue. The basement membrane acts as a selectively permeable membrane that determines which substances will be able to enter the epithelium.
827:
stratified squamous epithelium. Other surfaces that separate body cavities from the outside environment are lined by simple squamous, columnar, or pseudostratified epithelial cells. Other epithelial cells line the insides of the
614:
pseudostratified epithelial cells have cilia. Cilia are capable of energy-dependent pulsatile beating in a certain direction through interaction of cytoskeletal microtubules and connecting structural proteins and enzymes. In the
491:
Cuboidal epithelial cells have a cube-like shape and appear square in cross-section. The cell nucleus is large, spherical and is in the center of the cell. Cuboidal epithelium is commonly found in secretive tissue such as the
609:
These are simple columnar epithelial cells whose nuclei appear at different heights, giving the misleading (hence "pseudo") impression that the epithelium is stratified when the cells are viewed in cross section.
1047:. Additionally, the filaments that support these mesoderm-derived tissues are very distinct. Outside of the field of pathology, it is generally accepted that the epithelium arises from all three germ layers.
360:, either squamous, columnar or cuboidal. In some tissues, a layer of columnar cells may appear to be stratified due to the placement of the nuclei. This sort of tissue is called pseudostratified. All
2790:
1039:(both derived from mesoderm) to be true epithelium. This is because such tissues present very different pathology. For that reason, pathologists label cancers in endothelium and mesothelium
423:
that separates it from the underlying connective tissue. In general, it is found where absorption and filtration occur. The thinness of the epithelial barrier facilitates these processes.
907:
and the free/apical surface faces body fluid or outside. The basement membrane acts as a scaffolding on which epithelium can grow and regenerate after injuries. Epithelial tissue has a
2720:
637:
By layer, epithelium is classed as either simple epithelium, only one cell thick (unilayered), or stratified epithelium having two or more cells in thickness, or multi-layered – as
1464:
1393:
2783:
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3148:
1131:
from the infolding of epithelium and subsequent growth in the underlying connective tissue. They may be specialized columnar or cuboidal tissues consisting of
1381:
82:
3382:
279:
942:
are especially abundant in epithelial tissues. They consist of protein complexes and provide contact between neighbouring cells, between a cell and the
2776:
1330:), "nipple". Epithelium is so called because the name was originally used to describe the translucent covering of small "nipples" of tissue on the
2619:
2558:
1972:
1938:
1808:
1745:
1712:
1622:
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650:
599:. This consists of a layer of cells resting on at least one other layer of epithelial cells, which can be squamous, cuboidal, or columnar.
163:
1056:
rapidly activates cell division. Alternatively, when too many cells accumulate, crowding triggers their death by activation epithelial
1909:
1833:
953:
Cell junctions are the contact points between plasma membrane and tissue cells. There are mainly 5 different types of cell junctions:
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2301:
1997:
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1415:
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89:
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often necessary to use certain biochemical markers to make a positive identification. The intermediate filament proteins in the
447:
Squamous cells have the appearance of thin, flat plates that can look polygonal when viewed from above. Their name comes from
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629:
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135:
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In this case, the most apical layers of cells are filled with keratin, but they still retain their nuclei. These nuclei are
496:, or in absorptive tissue such as the pancreas, the lining of the kidney tubules as well as in the ducts of the glands. The
2072:"An epithelial cell destined for apoptosis signals its neighbors to extrude it by an actin- and myosin-dependent mechanism"
1762:
1151:
Endocrine glands secrete their product into the extracellular space where it is rapidly taken up by the circulatory system.
77:
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230:
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form the secretory and duct portions of glands. Stratified cuboidal epithelium protects areas such as the ducts of
505:
235:
181:
153:
130:
2751:
2740:
2729:
2709:
265:
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and function of the cells. The basic cell types are squamous, cuboidal, and columnar, classed by their shape.
2824:
2819:
2814:
1068:
172:
144:
121:
65:
53:
1357:
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in the lungs. Specialized squamous epithelium also forms the lining of cavities such as in blood vessels (as
2941:
2407:
Krausgruber T, Fortelny N, Fife-Gernedl V, Senekowitsch M, Schuster LC, Lercher A, et al. (July 2020).
1439:
1434:
1430:
712:
656:
571:
381:
217:
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has cells that can change from squamous to cuboidal, depending on the amount of tension on the epithelium.
3175:
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1197:
947:
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373:
207:
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212:
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Stratified epithelia (of columnar, cuboidal, or squamous type) can have the following specializations:
70:
2747:
2736:
2725:
2705:
2420:
2197:
2140:
2083:
2026:
1862:
1444:
1261:
943:
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is covered with fast-growing, easily regenerated epithelial cells. A specialised form of epithelium,
427:
306:
202:
3748:
2987:
2714:
1459:
1420:
1201:
1112:, as well as the secretion of sweat, mucus, enzymes, and other products that are delivered by ducts
801:
392:
19:
This article is about epithelium in animal anatomy. For the fungal structure of the same name, see
3471:
3466:
3445:
3073:
2505:
2456:
2109:
1288:
group are almost exclusively found in epithelial cells, so they are often used for this purpose.
1276:
shown at (3); an uninfected cell shown at (2) and (4) showing the difference between an infected
1109:
456:
222:
2700:
2127:
Eisenhoffer GT, Loftus PD, Yoshigi M, Otsuna H, Chien CB, Morcos PA, Rosenblatt J (April 2012).
2186:"KRas-transformed epithelia cells invade and partially dedifferentiate by basal cell extrusion"
2184:
Fadul J, Zulueta-Coarasa T, Slattum GM, Redd NM, Jin MF, Redd MJ, et al. (December 2021).
1674:
703:, meaning that they are highly condensed. Parakeratinized epithelium is sometimes found in the
558:. Some columnar cells are specialized for sensory reception such as in the nose, ears and the
3407:
3179:
3100:
3090:
2681:
2646:
2615:
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2497:
2448:
2389:
2340:
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2223:
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2101:
2052:
1993:
1968:
1934:
1926:
1905:
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1804:
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1737:
1708:
1678:
1618:
1591:
1564:
1526:
1494:
1300:
1097:
958:
928:
904:
887:, or external "skin", consists of a single layer of epithelial ectoderm from which arises the
616:
464:
420:
416:
396:
341:
1962:
1612:
754:
The lining of the blood and lymphatic vessels are of a specialised form of epithelium called
668:
apical, though in their most basal layers, the cells can be squamous, cuboidal, or columnar.
3670:
3432:
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2638:
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861:
805:
337:
1931:
Lepidoptera, Moths and
Butterflies: Morphology, Physiology, and Development : Teilband
1732:
Permar's oral embryology and microscopic anatomy: a textbook for students in dental hygiene
3703:
3684:
3641:
3500:
3078:
3009:
3004:
2241:
Gu Y, Shea J, Slattum G, Firpo MA, Alexander M, Mulvihill SJ, et al. (January 2015).
1639:
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1307:
1205:
1155:
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841:
793:
723:
584:
547:
322:
2013:
Gudipaty SA, Lindblom J, Loftus PD, Redd MJ, Edes K, Davey CF, et al. (March 2017).
1154:
Exocrine glands secrete their products into a duct that then delivers the product to the
2424:
2201:
2144:
2129:"Crowding induces live cell extrusion to maintain homeostatic cell numbers in epithelia"
2087:
2030:
1866:
766:
3546:
3512:
3451:
3422:
3310:
3260:
3111:
3085:
3032:
2443:
2408:
2384:
2359:
2319:"Recent advances in the molecular pathology, cell biology and genetics of ciliopathies"
2269:
2242:
2218:
2185:
2161:
2128:
2047:
2014:
1730:
1239:
1144:
1057:
954:
939:
873:
837:
620:
580:
518:
493:
426:
In general, epithelial tissues are classified by the number of their layers and by the
349:
302:
2096:
2071:
3742:
3614:
3572:
3566:
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3417:
3301:
3288:
3167:
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3027:
2509:
2460:
1667:
1213:
1020:
966:
551:
514:
480:
400:
345:
326:
1255:
352:. These tissues also lack blood or lymph supply. The tissue is supplied by nerves.
3665:
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3583:
3296:
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3209:
3053:
3048:
2113:
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1028:
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853:
535:
330:
2609:
1702:
1488:
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of an organ or onto the free surface of the epithelium. Their secretions include
58:
3601:
3344:
3340:
3241:
3236:
3205:
2857:
2852:
2243:"Defective apical extrusion signaling contributes to aggressive tumor hallmarks"
1770:
1454:
1285:
1132:
1103:
1036:
1032:
986:
869:
865:
849:
755:
704:
691:
oesophagus is an example of a non-keratinized or "moist" stratified epithelium.
575:
555:
534:
for maximizing the surface area for absorption, and these microvilli may form a
510:
476:
472:
468:
377:
318:
20:
2375:
2209:
1885:
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3461:
3335:
3305:
3246:
3196:
2946:
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2475:
2433:
1425:
1339:
1102:
the regulation and exchange of chemicals between the underlying tissues and a
884:
877:
797:
746:
719:
596:
531:
452:
37:
1875:
1850:
633:
Summary showing different epithelial cells/tissues and their characteristics.
3727:
3695:
3651:
3588:
3315:
3231:
3191:
2999:
2982:
2335:
2318:
1410:
1335:
1292:
1247:
of these cells, which enables a rapid response to immunological challenges.
1244:
1089:
1044:
962:
812:
771:
567:
563:
559:
388:
369:
365:
310:
2685:
2676:
2659:
2650:
2600:
2501:
2452:
2393:
2360:"Deciphering cell-cell interactions and communication from gene expression"
2344:
2278:
2227:
2170:
2105:
2056:
2015:"Mechanical stretch triggers rapid epithelial cell division through Piezo1"
3659:
3635:
3629:
3619:
3541:
3479:
3275:
3270:
3218:
3213:
3184:
3126:
2994:
1235:
1222:
1016:
1005:
994:
974:
946:, or they build up the paracellular barrier of epithelia and control the
700:
2409:"Structural cells are key regulators of organ-specific immune responses"
2259:
2152:
2038:
1310:, sweat glands are also damaged, causing a frosty coating of the skin.
923:
is made up of laminin (glycoproteins) secreted by epithelial cells. The
3485:
3265:
2642:
2592:
1826:
Histology: A Text and Atlas: With
Correlated Cell and Molecular Biology
1519:
Freshney RI (2002). "Introduction". In
Freshney RI, Freshney M (eds.).
1296:
1040:
888:
824:
750:
731:
685:
592:
364:
are made up of epithelial cells. Functions of epithelial cells include
2579:
Green H (September 2008). "The birth of therapy with cultured cells".
1828:(7th ed.). Lippincott Williams & Wilkins. pp. 528, 604.
1375:
Human cheek cells (nonkeratinized stratified squamous epithelium) 500×
419:
is a single layer of cells with every cell in direct contact with the
3522:
3506:
3360:
3331:
3107:
1209:
1193:
1171:
1163:
1128:
1093:
927:
beneath the basal lamina is made up of collagen proteins secreted by
892:
845:
820:
816:
775:
727:
611:
334:
2768:
745:
Epithelial tissue cells can adopt shapes of varying complexity from
95:
3556:
3551:
3535:
2629:
Nagpal R, Patel A, Gibson MC (March 2008). "Epithelial topology".
1254:
1179:
1159:
1136:
1075:
1067:
912:
908:
857:
765:
628:
539:
501:
361:
395:
of substances from the underlying connective tissue, through the
3593:
3348:
1118:
Absorb water and digested food in the lining of digestive canal.
895:, the rigidity of which varies as per its chemical composition.
829:
789:
460:
314:
3364:
3130:
2772:
2358:
Armingol E, Officer A, Harismendy O, Lewis NE (February 2021).
1399:
Histology of sweat gland showing stratified cuboidal epithelium
1331:
1167:
588:
2551:
Essential IVF : basic research and clinical applications
985:
Epithelial tissues are derived from all of the embryological
778:, with luminal epithelial cells annotated near bottom right.
1964:
The nature of disease: pathology for the health professions
1614:
Color atlas of cytology, histology, and microscopic anatomy
1387:
Histology of female urethra showing transitional epithelium
1291:
Cancers originating from the epithelium are classified as
1127:
Glandular tissue is the type of epithelium that forms the
1803:(3rd ed.). John Wiley & Sons. pp. 110–115.
1669:
DiFiore's Atlas of
Histology with Functional Correlations
463:, enabling simple passive diffusion as also found in the
1851:"3D Organisation of Cells in Pseudostratified Epithelia"
1794:
1792:
1790:
1788:
1204:
forming a tube or tubule with cilia projecting into the
2755:
Stratified squamous epithelium of the skin (thick skin)
836:, the reproductive and urinary tracts, and make up the
2744:
Stratified squamous epithelium of the skin (thin skin)
1268:
The slide shows at (1) an epithelial cell infected by
333:. Epithelial tissue is one of the four basic types of
2658:
Yamaguchi Y, Brenner M, Hearing VJ (September 2007).
2317:
Adams M, Smith UM, Logan CV, Johnson CA (May 2008).
1704:
From cells to organs: a histology textbook and atlas
1084:
Epithelial tissues have as their primary functions:
3431:
3398:
3324:
3286:
3165:
3066:
3041:
3020:
2975:
2968:
2961:
2920:
2886:
2840:
2833:
2807:
1465:
List of distinct cell types in the adult human body
860:, and is known as vascular endothelium, and lining
321:) tissues line the outer surfaces of many internal
76:
64:
52:
47:
30:
2296:(4th ed.). New York : Garland. p. 1267.
2070:Rosenblatt J, Raff MC, Cramer LP (November 2001).
1992:(4th ed.). New York : Garland. p. 1067.
1729:
1552:
1306:When epithelial cells or tissues are damaged from
977:(six of which come together to make a connexion).
903:The basal surface of epithelial tissue rests on a
2553:. Boston: Kluwer Academic Publishers. p. 3.
1967:. Lippincott Williams & Wilkins. p. 55.
1696:
1694:
1139:. There are two major classifications of glands:
623:, where the cilia propel the ovum to the uterus.
1234:Epithelial cells express many genes that encode
384:. Compound epithelium has protective functions.
2759:Stratified squamous epithelium of the esophagus
1514:
1512:
1510:
1080:Different characteristics of glands of the body
811:Tissues that line the inside of the mouth, the
403:are especially abundant in epithelial tissues.
16:Tissue lining the surfaces of organs in animals
2611:Basement membranes: cell and molecular biology
1673:. Lippincott Williams & Wilkins. pp.
1665:Eroschenko VP (2008). "Integumentary System".
1587:Color atlas of human anatomy: Locomotor system
3376:
3142:
2784:
1736:. Lippincott Williams & Wilkins. p.
1546:
1544:
1542:
1482:
1480:
1088:to protect the tissues that lie beneath from
1043:, whereas true epithelial cancers are called
273:
8:
2733:Stratified squamous epithelium of the vagina
1925:Kristensen NP, Georges C (1 December 2003).
1559:(3rd ed.). Benjamin/Cummings. pp.
1212:on epithelial cells provide chemosensation,
722:since it is almost exclusively found in the
707:and in the upper regions of the oesophagus.
387:Epithelial layers contain no blood vessels (
1849:Iber, Dagmar; Vetter, Roman (12 May 2022).
1801:Anatomy and Physiology from Science to Life
1490:Dellmann's Textbook of Veterinary Histology
301:is a thin, continuous, protective layer of
3383:
3369:
3361:
3149:
3135:
3127:
2972:
2965:
2917:
2883:
2837:
2791:
2777:
2769:
1904:. Boston: Benjamin Cummings. p. 133.
280:
266:
105:
36:
2675:
2491:
2442:
2432:
2383:
2334:
2268:
2258:
2217:
2160:
2095:
2046:
1884:
1874:
864:as lymphatic endothelium. Another type,
1200:, and they commonly exist as a sheet of
673:
546:. Other ciliated cells are found in the
432:
391:), so they must receive nourishment via
2476:"A gene atlas of 'structural immunity'"
1476:
1353:
245:
194:
171:
143:
120:
113:
2721:Diagrams of simple squamous epithelium
1096:, invasion by pathogens, and physical
1072:Forms of secretion in glandular tissue
325:, the corresponding inner surfaces of
93:
27:
2660:"The regulation of skin pigmentation"
2608:Kefalides NA, Borel JP, eds. (2005).
1487:Eurell JA, Frappier BL, eds. (2006).
1187:Sensing the extracellular environment
7:
2928:Pseudostratified columnar epithelium
651:pseudostratified columnar epithelium
2664:The Journal of Biological Chemistry
1108:the secretion of hormones into the
788:Epithelium lines both the outside (
109:This article is part of a series on
2713:Simple squamous epithelium of the
1933:. Walter de Gruyter. p. 484.
796:of bodies. The outermost layer of
784:Table of epithelia of human organs
254:Table of epithelia of human organs
14:
2549:Van Blerkom J, Gregory L (2004).
1728:Melfi RC, Alley KE, eds. (2000).
1416:Epithelial-mesenchymal transition
574:which are similar to microvilli.
542:to move mucus in the function of
3718:
3717:
2614:. Gulf Professional Publishing.
1392:
1380:
1368:
1356:
1280:and an uninfected cell nucleus.
90:Anatomical terms of microanatomy
1799:Jenkins GW, Tortora GJ (2013).
1493:. Wiley-Blackwell. p. 18.
643:stratified cuboidal epithelium
2908:Stratified columnar epithelium
2901:Stratified cuboidal epithelium
2894:Stratified squamous epithelium
1769:. Amirsys, Inc. Archived from
1259:Epithelial cell infected with
792:) and the inside cavities and
647:stratified columnar epithelium
639:stratified squamous epithelium
1:
2294:Molecular Biology of the Cell
2097:10.1016/S0960-9822(01)00587-5
1990:Molecular biology of the cell
1326:), "on" or "upon", and θηλή (
313:, the outermost layer of the
1640:"Simple Cuboidal Epithelium"
1555:Human Anatomy and Physiology
1019:(e.g., the inner linings of
852:, forms the inner lining of
329:, and the inner surfaces of
2701:Epithelium Photomicrographs
2474:Minton K (September 2020).
2323:Journal of Medical Genetics
1824:Ross MH, Pawlina W (2015).
1522:Culture of epithelial cells
1342:senses; the plural form is
1314:Etymology and pronunciation
844:. The outer surface of the
3770:
2874:Simple columnar epithelium
2867:Simple cuboidal epithelium
2848:Simple squamous epithelium
2480:Nature Reviews. Immunology
2376:10.1038/s41576-020-00292-x
2210:10.1038/s41467-021-27513-z
1450:Retinal pigment epithelium
1322:uses the Greek roots ἐπί (
1192:Some epithelial cells are
781:
18:
3713:
2493:10.1038/s41577-020-0398-y
2434:10.1038/s41586-020-2424-4
1525:. John Wiley & Sons.
1238:and proteins involved in
1008:(e.g., the lining of the
868:, forms the walls of the
88:
35:
2364:Nature Reviews. Genetics
1902:Anatomy & Physiology
1876:10.3389/fphy.2022.898160
1363:Squamous epithelium 100×
173:Cuboidal epithelial cell
145:Columnar epithelial cell
122:Squamous epithelial cell
3176:Cell adhesion molecules
2942:Transitional epithelium
2336:10.1136/jmg.2007.054999
1617:. Thieme. p. 102.
1440:Iris pigment epithelium
1435:outer enamel epithelium
1240:cell-cell communication
891:, an outer covering of
657:Transitional epithelium
500:that covers the female
382:transcellular transport
2933:Respiratory epithelium
2764:Microanatomy Web Atlas
2677:10.1074/jbc.R700026200
1701:van Lommel AT (2002).
1265:
1227:
1198:respiratory epithelium
1081:
1073:
1010:gastrointestinal tract
948:paracellular transport
834:gastrointestinal tract
779:
634:
2190:Nature Communications
1961:McConnell TH (2006).
1590:. Thieme. p. 8.
1258:
1251:Clinical significance
1190:
1079:
1071:
769:
663:Stratified epithelium
632:
544:mucociliary clearance
538:. Other cells may be
195:Specialised epithelia
1855:Frontiers in Physics
1445:Neuroepithelial cell
1334:. The word has both
1270:Chlamydia pneumoniae
1262:Chlamydia pneumoniae
1230:Host immune response
1115:to provide sensation
944:extracellular matrix
800:is composed of dead
309:. An example is the
307:extracellular matrix
3630:Meristematic tissue
2670:(38): 27557–27561.
2425:2020Natur.583..296K
2260:10.7554/eLife.04069
2202:2021NatCo..12.7180F
2153:10.1038/nature10999
2145:2012Natur.484..546E
2088:2001CBio...11.1847R
2039:10.1038/nature21407
2031:2017Natur.543..118G
1886:20.500.11850/547113
1867:2022FrP....10.8160I
1773:on 19 December 2012
1460:Sulcular epithelium
1421:Epithelial polarity
883:In arthropods, the
802:stratified squamous
506:germinal epithelium
498:germinal epithelium
465:alveolar epithelium
42:Types of epithelium
3392:Biological tissues
3074:Myoepithelial cell
2643:10.1002/bies.20722
2593:10.1002/bies.20797
2528:Cancer Research UK
2292:Alberts B (2002).
1988:Alberts B (2002).
1763:"Epithelial Cells"
1584:Platzer W (2008).
1551:Marieb EM (1995).
1266:
1110:circulatory system
1082:
1074:
959:adherens junctions
819:, and part of the
808:epithelial cells.
780:
635:
71:H2.00.02.0.00002
3754:Tissues (biology)
3736:
3735:
3523:Phloem parenchyma
3358:
3357:
3257:membrane proteins
3180:Adherens junction
3124:
3123:
3120:
3119:
3062:
3061:
2957:
2956:
2916:
2915:
2882:
2881:
2748:Histology at KUMC
2737:Histology at KUMC
2726:Histology at KUMC
2706:Histology at KUMC
2621:978-0-12-153356-4
2560:978-1-4020-7551-3
2530:. 28 October 2014
2524:"Types of cancer"
2419:(7815): 296–302.
2139:(7395): 546–549.
2082:(23): 1847–1857.
2025:(7643): 118–121.
1974:978-0-7817-5317-3
1940:978-3-11-016210-3
1900:Marieb E (2011).
1810:978-1-118-12920-3
1747:978-0-683-30644-6
1714:978-1-4020-7257-4
1624:978-3-13-562404-4
1611:Kühnel W (2003).
1597:978-3-13-533306-9
1532:978-0-471-40121-6
1500:978-0-7817-4148-4
1350:Additional images
1301:connective tissue
929:connective tissue
905:basement membrane
899:Basement membrane
862:lymphatic vessels
738:
737:
627:
626:
617:respiratory tract
550:, the uterus and
421:basement membrane
417:Simple epithelium
412:Simple epithelium
397:basement membrane
342:connective tissue
299:epithelial tissue
290:
289:
104:
103:
99:
3761:
3721:
3720:
3671:Vascular cambium
3557:Xylem parenchyma
3385:
3378:
3371:
3362:
3151:
3144:
3137:
3128:
2973:
2966:
2918:
2884:
2838:
2793:
2786:
2779:
2770:
2689:
2679:
2654:
2625:
2604:
2565:
2564:
2546:
2540:
2539:
2537:
2535:
2520:
2514:
2513:
2495:
2471:
2465:
2464:
2446:
2436:
2404:
2398:
2397:
2387:
2355:
2349:
2348:
2338:
2314:
2308:
2307:
2289:
2283:
2282:
2272:
2262:
2238:
2232:
2231:
2221:
2181:
2175:
2174:
2164:
2124:
2118:
2117:
2099:
2067:
2061:
2060:
2050:
2010:
2004:
2003:
1985:
1979:
1978:
1958:
1952:
1951:
1949:
1947:
1922:
1916:
1915:
1897:
1891:
1890:
1888:
1878:
1846:
1840:
1839:
1821:
1815:
1814:
1796:
1783:
1782:
1780:
1778:
1758:
1752:
1751:
1735:
1725:
1719:
1718:
1698:
1689:
1688:
1672:
1662:
1656:
1655:
1653:
1651:
1635:
1629:
1628:
1608:
1602:
1601:
1581:
1575:
1574:
1558:
1548:
1537:
1536:
1516:
1505:
1504:
1484:
1396:
1384:
1372:
1360:
1274:inclusion bodies
1236:immune mediators
1218:mechanosensation
1196:, especially in
1176:digestive juices
1141:endocrine glands
1135:, which secrete
1123:Glandular tissue
1031:do not consider
925:reticular lamina
842:endocrine glands
823:are composed of
696:Parakeratinized
674:
605:Pseudostratified
479:), and in other
433:
282:
275:
268:
164:Pseudostratified
106:
96:edit on Wikidata
40:
28:
3769:
3768:
3764:
3763:
3762:
3760:
3759:
3758:
3739:
3738:
3737:
3732:
3709:
3642:Ground meristem
3501:Vascular tissue
3472:Subsidiary cell
3427:
3394:
3389:
3359:
3354:
3320:
3282:
3161:
3155:
3125:
3116:
3079:Serous demilune
3058:
3037:
3016:
2953:
2912:
2878:
2829:
2803:
2797:
2752:epithel-epith15
2741:epithel-epith14
2730:epithel-epith12
2710:epithel-epith02
2697:
2692:
2657:
2628:
2622:
2607:
2578:
2574:
2572:Further reading
2569:
2568:
2561:
2548:
2547:
2543:
2533:
2531:
2522:
2521:
2517:
2473:
2472:
2468:
2406:
2405:
2401:
2357:
2356:
2352:
2316:
2315:
2311:
2304:
2291:
2290:
2286:
2240:
2239:
2235:
2183:
2182:
2178:
2126:
2125:
2121:
2076:Current Biology
2069:
2068:
2064:
2012:
2011:
2007:
2000:
1987:
1986:
1982:
1975:
1960:
1959:
1955:
1945:
1943:
1941:
1924:
1923:
1919:
1912:
1899:
1898:
1894:
1848:
1847:
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1786:
1776:
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1760:
1759:
1755:
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1727:
1726:
1722:
1715:
1700:
1699:
1692:
1685:
1664:
1663:
1659:
1649:
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1637:
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1632:
1625:
1610:
1609:
1605:
1598:
1583:
1582:
1578:
1571:
1550:
1549:
1540:
1533:
1518:
1517:
1508:
1501:
1486:
1485:
1478:
1473:
1407:
1400:
1397:
1388:
1385:
1376:
1373:
1364:
1361:
1352:
1316:
1308:cystic fibrosis
1295:. In contrast,
1253:
1232:
1202:polarised cells
1189:
1145:exocrine glands
1125:
1092:, desiccation,
1066:
1053:
983:
955:tight junctions
937:
901:
786:
764:
743:
677:Specialization
665:
621:fallopian tubes
585:salivary glands
548:fallopian tubes
519:salivary glands
494:exocrine glands
414:
409:
286:
100:
43:
24:
17:
12:
11:
5:
3767:
3765:
3757:
3756:
3751:
3741:
3740:
3734:
3733:
3731:
3730:
3725:
3714:
3711:
3710:
3708:
3707:
3700:
3699:
3698:
3693:
3676:
3675:
3674:
3673:
3668:
3656:
3655:
3654:
3649:
3644:
3625:
3624:
3623:
3622:
3617:
3605:
3598:
3597:
3596:
3591:
3586:
3581:
3562:
3561:
3560:
3559:
3554:
3549:
3547:Vessel element
3544:
3532:
3531:
3530:
3525:
3520:
3515:
3513:Companion cell
3496:
3495:
3494:
3493:
3488:
3476:
3475:
3474:
3469:
3464:
3459:
3454:
3452:Bulliform cell
3437:
3435:
3429:
3428:
3426:
3425:
3420:
3415:
3410:
3404:
3402:
3396:
3395:
3390:
3388:
3387:
3380:
3373:
3365:
3356:
3355:
3353:
3352:
3338:
3328:
3326:
3322:
3321:
3319:
3318:
3313:
3311:Focal adhesion
3308:
3299:
3293:
3291:
3284:
3283:
3281:
3280:
3279:
3278:
3273:
3268:
3261:Tight junction
3252:
3251:
3250:
3249:
3244:
3239:
3224:
3223:
3222:
3221:
3202:
3201:
3200:
3199:
3189:
3188:
3187:
3172:
3170:
3163:
3162:
3156:
3154:
3153:
3146:
3139:
3131:
3122:
3121:
3118:
3117:
3115:
3114:
3105:
3104:
3103:
3098:
3093:
3082:
3081:
3076:
3070:
3068:
3064:
3063:
3060:
3059:
3057:
3056:
3051:
3045:
3043:
3039:
3038:
3036:
3035:
3033:Alveolar gland
3030:
3024:
3022:
3018:
3017:
3015:
3014:
3013:
3012:
3007:
2997:
2992:
2991:
2990:
2979:
2977:
2970:
2963:
2959:
2958:
2955:
2954:
2952:
2951:
2950:
2949:
2938:
2937:
2936:
2935:
2924:
2922:
2914:
2913:
2911:
2910:
2904:
2903:
2897:
2896:
2890:
2888:
2880:
2879:
2877:
2876:
2870:
2869:
2863:
2862:
2861:
2860:
2855:
2844:
2842:
2835:
2831:
2830:
2828:
2827:
2822:
2817:
2811:
2809:
2805:
2804:
2798:
2796:
2795:
2788:
2781:
2773:
2767:
2766:
2761:
2756:
2745:
2734:
2723:
2718:
2703:
2696:
2695:External links
2693:
2691:
2690:
2655:
2637:(3): 260–266.
2626:
2620:
2605:
2587:(9): 897–903.
2575:
2573:
2570:
2567:
2566:
2559:
2541:
2515:
2486:(9): 518–519.
2466:
2399:
2350:
2329:(5): 257–267.
2309:
2302:
2284:
2233:
2176:
2119:
2062:
2005:
1998:
1980:
1973:
1953:
1939:
1917:
1911:978-0321616401
1910:
1892:
1841:
1835:978-1451187427
1834:
1816:
1809:
1784:
1753:
1746:
1720:
1713:
1690:
1683:
1657:
1646:. Amirsys, Inc
1630:
1623:
1603:
1596:
1576:
1569:
1538:
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1428:
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1188:
1185:
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1183:
1152:
1124:
1121:
1120:
1119:
1116:
1113:
1106:
1100:
1065:
1062:
1058:cell extrusion
1052:
1049:
1025:
1024:
1013:
1002:
982:
979:
967:hemidesmosomes
940:Cell junctions
936:
935:Cell junctions
933:
900:
897:
825:nonkeratinized
763:
760:
742:
739:
736:
735:
715:
709:
708:
697:
693:
692:
688:
682:
681:
678:
664:
661:
625:
624:
607:
601:
600:
527:
523:
522:
515:mammary glands
489:
485:
484:
445:
441:
440:
437:
413:
410:
408:
407:Classification
405:
401:Cell junctions
368:, filtration,
350:nervous tissue
317:. Epithelial (
288:
287:
285:
284:
277:
270:
262:
259:
258:
257:
256:
248:
247:
243:
242:
241:
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239:
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233:
225:
220:
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210:
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197:
196:
192:
191:
190:
189:
184:
176:
175:
169:
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167:
166:
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156:
148:
147:
141:
140:
139:
138:
133:
125:
124:
118:
117:
111:
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102:
101:
92:
86:
85:
80:
74:
73:
68:
62:
61:
56:
50:
49:
45:
44:
41:
33:
32:
15:
13:
10:
9:
6:
4:
3:
2:
3766:
3755:
3752:
3750:
3747:
3746:
3744:
3729:
3726:
3724:
3716:
3715:
3712:
3706:
3705:
3701:
3697:
3694:
3692:
3689:
3688:
3687:
3686:
3681:
3678:
3677:
3672:
3669:
3667:
3664:
3663:
3662:
3661:
3657:
3653:
3650:
3648:
3645:
3643:
3640:
3639:
3638:
3637:
3632:
3631:
3627:
3626:
3621:
3618:
3616:
3613:
3612:
3611:
3610:
3606:
3604:
3603:
3599:
3595:
3592:
3590:
3587:
3585:
3582:
3580:
3577:
3576:
3575:
3574:
3569:
3568:
3567:Ground tissue
3564:
3563:
3558:
3555:
3553:
3550:
3548:
3545:
3543:
3540:
3539:
3538:
3537:
3533:
3529:
3526:
3524:
3521:
3519:
3516:
3514:
3511:
3510:
3509:
3508:
3503:
3502:
3498:
3497:
3492:
3489:
3487:
3484:
3483:
3482:
3481:
3477:
3473:
3470:
3468:
3467:Pavement cell
3465:
3463:
3460:
3458:
3455:
3453:
3450:
3449:
3448:
3447:
3442:
3441:Dermal tissue
3439:
3438:
3436:
3434:
3430:
3424:
3421:
3419:
3416:
3414:
3411:
3409:
3406:
3405:
3403:
3401:
3397:
3393:
3386:
3381:
3379:
3374:
3372:
3367:
3366:
3363:
3350:
3346:
3342:
3339:
3337:
3333:
3330:
3329:
3327:
3323:
3317:
3314:
3312:
3309:
3307:
3303:
3302:Hemidesmosome
3300:
3298:
3295:
3294:
3292:
3290:
3285:
3277:
3274:
3272:
3269:
3267:
3264:
3263:
3262:
3258:
3254:
3253:
3248:
3245:
3243:
3240:
3238:
3235:
3234:
3233:
3229:
3226:
3225:
3220:
3217:
3216:
3215:
3211:
3207:
3204:
3203:
3198:
3195:
3194:
3193:
3190:
3186:
3183:
3182:
3181:
3177:
3174:
3173:
3171:
3169:
3164:
3160:
3152:
3147:
3145:
3140:
3138:
3133:
3132:
3129:
3113:
3109:
3106:
3102:
3101:Intercalated
3099:
3097:
3094:
3092:
3089:
3088:
3087:
3084:
3083:
3080:
3077:
3075:
3072:
3071:
3069:
3065:
3055:
3054:Mucous glands
3052:
3050:
3049:Serous glands
3047:
3046:
3044:
3040:
3034:
3031:
3029:
3028:Tubular gland
3026:
3025:
3023:
3019:
3011:
3008:
3006:
3003:
3002:
3001:
2998:
2996:
2993:
2989:
2986:
2985:
2984:
2981:
2980:
2978:
2974:
2971:
2967:
2964:
2960:
2948:
2945:
2944:
2943:
2940:
2939:
2934:
2931:
2930:
2929:
2926:
2925:
2923:
2919:
2909:
2906:
2905:
2902:
2899:
2898:
2895:
2892:
2891:
2889:
2885:
2875:
2872:
2871:
2868:
2865:
2864:
2859:
2856:
2854:
2851:
2850:
2849:
2846:
2845:
2843:
2839:
2836:
2832:
2826:
2823:
2821:
2818:
2816:
2813:
2812:
2810:
2806:
2801:
2794:
2789:
2787:
2782:
2780:
2775:
2774:
2771:
2765:
2762:
2760:
2757:
2754:
2753:
2749:
2746:
2743:
2742:
2738:
2735:
2732:
2731:
2727:
2724:
2722:
2719:
2716:
2712:
2711:
2707:
2704:
2702:
2699:
2698:
2694:
2687:
2683:
2678:
2673:
2669:
2665:
2661:
2656:
2652:
2648:
2644:
2640:
2636:
2632:
2627:
2623:
2617:
2613:
2612:
2606:
2602:
2598:
2594:
2590:
2586:
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1211:
1210:Primary cilia
1207:
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1059:
1051:Cell turnover
1050:
1048:
1046:
1042:
1038:
1034:
1030:
1022:
1021:body cavities
1018:
1014:
1011:
1007:
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996:
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991:
990:
988:
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978:
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552:central canal
549:
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541:
537:
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528:
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516:
512:
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495:
490:
487:
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481:body cavities
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363:
359:
353:
351:
347:
346:muscle tissue
343:
340:, along with
339:
336:
332:
331:blood vessels
328:
327:body cavities
324:
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316:
312:
308:
304:
300:
296:
295:
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67:
63:
60:
57:
55:
51:
46:
39:
34:
29:
26:
22:
3702:
3683:
3679:
3666:Cork cambium
3658:
3634:
3628:
3609:Sclerenchyma
3607:
3600:
3584:Chlorenchyma
3571:
3565:
3534:
3518:Phloem fiber
3505:
3499:
3478:
3444:
3440:
3412:
3297:Basal lamina
3228:Cytoskeleton
3210:Gap junction
3206:Ion channels
3158:
3157:Proteins of
3091:Intralobular
2799:
2750:
2739:
2728:
2708:
2667:
2663:
2634:
2630:
2610:
2584:
2580:
2550:
2544:
2532:. Retrieved
2527:
2518:
2483:
2479:
2469:
2416:
2412:
2402:
2370:(2): 71–88.
2367:
2363:
2353:
2326:
2322:
2312:
2293:
2287:
2250:
2246:
2236:
2193:
2189:
2179:
2136:
2132:
2122:
2079:
2075:
2065:
2022:
2018:
2008:
1989:
1983:
1963:
1956:
1944:. Retrieved
1930:
1927:"Integument"
1920:
1901:
1895:
1858:
1854:
1844:
1825:
1819:
1800:
1777:28 September
1775:. Retrieved
1771:the original
1766:
1756:
1731:
1723:
1707:. Springer.
1703:
1668:
1660:
1650:28 September
1648:. Retrieved
1643:
1633:
1613:
1606:
1586:
1579:
1554:
1521:
1489:
1343:
1327:
1323:
1319:
1317:
1305:
1290:
1282:
1278:cell nucleus
1269:
1267:
1260:
1233:
1191:
1133:goblet cells
1126:
1083:
1054:
1029:pathologists
1026:
984:
952:
938:
921:basal lamina
918:
913:blood supply
909:nerve supply
902:
882:
810:
787:
744:
713:Transitional
680:Description
670:
666:
655:
636:
576:Goblet cells
536:brush border
511:sweat glands
448:
439:Description
425:
415:
386:
372:, selective
357:
354:
305:with little
298:
293:
292:
291:
218:Transitional
114:
25:
3602:Collenchyma
3552:Xylem fiber
3345:Stereocilia
3289:cell–matrix
3242:Plakoglobin
3237:Desmoplakin
2858:Mesothelium
2853:Endothelium
2196:(1): 7180.
1455:Skin cancer
1299:develop in
1286:cytokeratin
1104:body cavity
1037:mesothelium
1033:endothelium
997:(e.g., the
987:germ layers
981:Development
870:pericardium
866:mesothelium
850:endothelium
806:keratinized
756:endothelium
705:oral mucosa
686:Keratinized
581:lobar ducts
572:stereocilia
556:spinal cord
477:mesothelium
473:pericardium
469:endothelium
378:germination
319:mesothelial
208:Respiratory
48:Identifiers
21:Pileipellis
3749:Epithelium
3743:Categories
3691:Endodermis
3647:Procambium
3579:Aerenchyma
3573:Parenchyma
3528:Sieve tube
3491:Phelloderm
3462:Guard cell
3413:Epithelial
3408:Connective
3341:Microvilli
3336:Kinocilium
3306:Tonofibril
3247:Tonofibril
3197:Desmoglein
3159:epithelium
3067:Components
2947:Urothelium
2887:stratified
2800:Epithelial
2715:glomerulus
2534:13 October
2253:: e04069.
1946:10 January
1767:AnatomyOne
1644:AnatomyOne
1471:References
1426:Glycocalyx
1320:epithelium
1293:carcinomas
1045:carcinomas
963:desmosomes
885:integument
878:peritoneum
798:human skin
782:See also:
747:polyhedral
720:urothelium
597:sex organs
568:inner ears
564:Hair cells
560:taste buds
532:microvilli
504:, and the
471:), in the
374:absorption
294:Epithelium
213:Intestinal
187:Stratified
159:Stratified
136:Stratified
31:Epithelium
3728:Histology
3696:Exodermis
3660:Secondary
3652:Protoderm
3589:Mesophyll
3446:Epidermis
3316:Costamere
3232:Desmosome
3192:Desmosome
3168:cell–cell
3096:Striated
3042:Secretion
3010:Meibomian
3005:Sebaceous
3000:Holocrine
2983:Merocrine
2976:Mechanism
2820:Cuboidal
2815:Squamous
2631:BioEssays
2581:BioEssays
2510:220491226
2461:220295181
1761:Pratt R.
1638:Pratt R.
1411:Dark cell
1344:epithelia
1318:The word
1245:epigenome
1170:(sebum),
1090:radiation
1064:Functions
1027:However,
999:epidermis
975:connexins
911:, but no
813:esophagus
772:histology
751:scutoidal
741:Structure
526:Columnar
488:Cuboidal
444:Squamous
393:diffusion
389:avascular
370:secretion
366:diffusion
311:epidermis
227:Germinal
203:Olfactory
115:Epithelia
3723:Category
3620:Sclereid
3542:Tracheid
3480:Periderm
3418:Muscular
3276:MARVELD2
3271:Occludin
3219:Connexin
3214:Connexon
3185:Cadherin
3166:Lateral/
2995:Apocrine
2825:Columnar
2717:(kidney)
2686:17635904
2651:18293365
2601:18693268
2502:32661408
2453:32612232
2394:33168968
2345:18178628
2279:25621765
2228:34893591
2171:22504183
2106:11728307
2057:28199303
1405:See also
1297:sarcomas
1272:; their
1194:ciliated
1041:sarcomas
1017:mesoderm
1006:endoderm
995:ectoderm
856:and the
838:exocrine
762:Location
701:pyknotic
612:Ciliated
540:ciliated
358:compound
3636:Primary
3486:Phellem
3457:Cuticle
3423:Nervous
3400:Animals
3266:Claudin
2988:Eccrine
2444:7610345
2421:Bibcode
2385:7649713
2270:4337653
2219:8664939
2198:Bibcode
2162:4593481
2141:Bibcode
2114:5858676
2084:Bibcode
2048:5334365
2027:Bibcode
1863:Bibcode
1561:103–104
889:cuticle
874:pleurae
774:of the
770:Normal
732:urethra
728:ureters
724:bladder
593:pharynx
583:in the
566:in the
554:of the
459:in the
457:alveoli
223:Vaginal
59:D004848
3685:Cortex
3507:Phloem
3433:Plants
3325:Apical
3287:Basal/
3255:other
3108:Acinus
2962:Glands
2841:Simple
2802:tissue
2684:
2649:
2618:
2599:
2557:
2508:
2500:
2459:
2451:
2441:
2413:Nature
2392:
2382:
2343:
2300:
2277:
2267:
2226:
2216:
2169:
2159:
2133:Nature
2112:
2104:
2055:
2045:
2019:Nature
1996:
1971:
1937:
1908:
1832:
1807:
1744:
1711:
1681:
1677:–234.
1621:
1594:
1567:
1529:
1497:
1223:motile
1216:, and
1182:, etc.
1172:enzyme
1164:saliva
1129:glands
1098:trauma
1094:toxins
969:, and
893:chitin
876:, and
846:cornea
832:, the
821:rectum
817:vagina
815:, the
794:lumina
776:breast
645:, and
595:, and
591:, the
587:, the
517:, and
449:squāma
380:, and
362:glands
338:tissue
335:animal
323:organs
231:female
182:Simple
154:Simple
131:Simple
3704:Stele
3680:Mixed
3615:Fiber
3536:Xylem
3332:Cilia
3086:Ducts
3021:Shape
2969:Types
2921:Other
2834:Types
2808:Cells
2506:S2CID
2457:S2CID
2247:eLife
2110:S2CID
1431:Inner
1340:count
1328:thēlē
1206:lumen
1180:sweat
1160:tears
1156:lumen
1137:mucus
1015:from
1004:from
993:from
858:heart
830:lungs
570:have
502:ovary
436:Type
428:shape
303:cells
246:Other
94:[
3594:Pith
3349:STRC
3112:Lobe
2682:PMID
2647:PMID
2616:ISBN
2597:PMID
2555:ISBN
2536:2016
2498:PMID
2449:PMID
2390:PMID
2341:PMID
2298:ISBN
2275:PMID
2224:PMID
2167:PMID
2102:PMID
2053:PMID
1994:ISBN
1969:ISBN
1948:2013
1935:ISBN
1906:ISBN
1830:ISBN
1805:ISBN
1779:2012
1742:ISBN
1709:ISBN
1679:ISBN
1652:2012
1619:ISBN
1592:ISBN
1565:ISBN
1527:ISBN
1495:ISBN
1433:and
1338:and
1336:mass
1221:are
1143:and
1035:and
919:The
840:and
790:skin
730:and
475:(as
461:lung
453:skin
348:and
315:skin
236:male
83:9639
54:MeSH
2672:doi
2668:282
2639:doi
2589:doi
2488:doi
2439:PMC
2429:doi
2417:583
2380:PMC
2372:doi
2331:doi
2265:PMC
2255:doi
2214:PMC
2206:doi
2157:PMC
2149:doi
2137:484
2092:doi
2043:PMC
2035:doi
2023:543
1881:hdl
1871:doi
1675:212
1332:lip
1324:epi
1208:."
1168:oil
749:to
589:eye
455:or
297:or
78:FMA
3745::
3682::
3633::
3570::
3504::
3443::
3259::
3230::
3208::
3178::
2680:.
2666:.
2662:.
2645:.
2635:30
2633:.
2595:.
2585:30
2583:.
2526:.
2504:.
2496:.
2484:20
2482:.
2478:.
2455:.
2447:.
2437:.
2427:.
2415:.
2411:.
2388:.
2378:.
2368:22
2366:.
2362:.
2339:.
2327:45
2325:.
2321:.
2273:.
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2249:.
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2222:.
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2194:12
2192:.
2188:.
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2155:.
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2108:.
2100:.
2090:.
2080:11
2078:.
2074:.
2051:.
2041:.
2033:.
2021:.
2017:.
1929:.
1879:.
1869:.
1861:.
1859:10
1857:.
1853:.
1787:^
1765:.
1740:.
1693:^
1642:.
1563:.
1541:^
1509:^
1479:^
1346:.
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521:.
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3212:/
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98:]
23:.
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