Histology of Glands
Number of interconnected units acinus / tubule / cord Arrangement of secretory cells Destination of product Nature of product: Duct function Duct location Tissue composition
Examples of glands
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Introduction to Glandular Tissue
Glands are organized arrangements of secretory cells. All exocrine glands (and also most endocrine glands), are composed of epithelial tissue. Even the lung and the kidney, although not properly glands, have a glandular pattern of tissue organization.
Although most glands give the appearance of being "solid" tissue, their epithelial nature is expressed by the organization of their cells, with each cell attached laterally to its neighbors. Every exocrine secretory cell has some portion of its plasma membrane exposed to an external surface, communicating with the outside of the body by way of a system of ducts.
In most glands, the secretory cells are organized into secretory units, which are described according to their shape as tubules, acini, or cords.
In the diagrams here and below, secretory units are colored orange and ducts are colored blue.
The basic organization of epithelial tissue into a gland can be best seen in a very simple gland, like the one from frog skin shown here. Click on here or on the image to open an enlarged view.
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Histologically, glands are described using some standard vocabulary.
Simple / Compound
The simple / compound distinction is based on on duct shape.
A simple gland has an unbranched duct (or no duct at all). There is only a single secretory unit (acinus or tubule). Examples include sweat glands, gastric glands, intestinal crypts, and uterine glands.
A compound gland has a branching duct. Salivary glands and pancreas are familiar examples. Compound glands are typically fairly bulky and contain very many individual secretory units (acini or tubules).
Random tissue sections seldom show ducts branching. Nevertheless, the appearance of multiple duct profiles, in various sizes, provides evidence of a branching duct system.
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Acinus / Tubule / Cord
Each secretory unit of a gland consists of cells arranged into an acinus, a tubule, or a cord. Each of of these arrangements has a different and characteristic appearance when viewed in section.
Acinus (or alveolus)
An acinus (from Latin, grape) is a small ball of secretory epithelial cells containing a tiny central lumen. Acini are usually formed by serous cells. [Acini are sometimes called alveoli, from L., small cavity.]
A typical acinar cell is shaped like a pyramid. Its basal surface, located at the periphery of the acinus, rests on the basement membrane, which separates the acinus from the underlying stroma. Its lateral surfaces (the sides of the pyramid) are attached to adjacent secretory cells. Its apical surface is free and faces the acinar lumen, which communicates by duct with the outside. The acinar cell's cytoplasm is also visibly polarized, usually with basophilic basal cytoplasm and variously-staining secretory granules concentrated in apical cytoplasm. For more, see serous cells.
A compound acinar gland can be quite accurately modelled as a bunch of grapes embedded in Jello. The grapes are the acini, the branching stems are the ducts, and the Jello represents the rest of the stroma. Major and minor branches of the bunch represent lobes and lobules, respectively, separated by greater amounts of connective tissue.
In routine tissue sections, most acini are cut in random planes and look like solid lumps, made of cells having various sizes and shapes. The lumen of an acinus is typically quite tiny (i.e., much smaller than a cell) and so is visible only when an acinus is sliced neatly across the middle. In such a slice, the cells look like slices of pie, with the lumen in the center, as in the upper thumbnail image at right.
Secretory cells may also arrange themselves into secretory tubules (in contrast to the small balls of cells which comprise secretory acini). Sweat glands are probably the most familiar tubular glands. Other tubular glands include gastric glands (in the lining of the stomach), uterine glands, and various mucous glands of the GI system.
Because tubules are elongated, random sections commonly include the lumen as well as the secretory cells themselves (in contrast to the situation with acini). But interpretation of the sectioned appearance of tubular glands will depend on whether the tubules are simple or branched, on whether they are straight or twisted, and on whether or not adjacent tubules lie parallel to one another.
Cords are epithelial cells arranged into sheets separated by vascular sinusoids. In section, the predominant pattern appears linear (hence, "cord"), even though the lines may twist and branch.
Cords are a common arrangement for endocrine secretion. The cells retain an epithelial character, attached to neighboring cells, even though they may no longer comprise a surface barrier between interstitial space and a secretory lumen that leads to the outside. Examples of endocrine cells arranged into cords include the epithelial cells of pancreatic islets, parathyroid, adrenal cortex, and liver.
The liver (in the thumbnails) is notable for having cells arranged into cords in spite of its major exocrine function. In order to maintain communication with ducts, the liver cords contain a network of intercellular channels called bile canaliculi.
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Endocrine / Exocrine
The suffix -crine refers to secretion; the prefix endo- or exo- tells where the secretory product goes.
The product of exocrine glands leaves the body proper, either by direct secretion onto the body's surface (e.g., sweat) or into the lumen of an organ (e.g., gastric juice) or else by flowing through a system of ducts (e.g., saliva, pancreatic enzymes, bile). The cells of exocrine glands are generally arranged into secretory units in the form of acini or tubules (although the liver has a remarkable arrangement of cords).
The product of endocrine glands is secreted into interstitial fluid and hence into capillaries and general circulation. The cells of endocrine glands are often arranged into cords adjacent to capillaries or sinusoids.
Link to the endocrine system.
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Serous / Mucous / Mixed
The serous / mucous distinction is based on the secretory cell's product -- whether it is a clear, watery solution of enzymes (serous, like serum) or else a glycoprotein mixture (mucous, like mucin). These two categories of secretory products come from two distinct categories of cells, each with a characteristic appearance.
Mixed glands (e.g., most salivary glands) contain both types of cells. Glands which contain only one of these two cell types may be described either as serous glands (e.g., parotid gland or pancreas) or as mucous glands (e.g., Brunner's glands of the duodenum).
Serous cells are specialized to secrete an enzyme solution. Examples include serous cells of the salivary glands, exocrine cells of the pancreas, gastric chief cells, and Paneth cells of intestinal crypts. Serous cells of the pancreas and the salivary glands are typically organized into secretory units called acini.
In routine light microscopy, serous cells are distinguished by basophilic basal cytoplasm, a centrally-located nucleus, and variously-staining secretory vesicles (zymogen granules) in apical cytoplasm. These features are all associated with organized mass production of protein for export. More.
Cells which are specialized to secrete mucus are called mucous cells. Examples include secretory cells of the salivary glands, esophageal glands, stomach surface, pyloric glands, and Brunner's glands of the duodenum. These cells are typically organized into tubular secretory units.
Goblet cells are mucous cells which stand alone within the intestinal epithelium. Goblet cells take their name from their characteristic shape, with a broad opening at the apical end and a narrow, "pinched" base. Cells with this goblet shape are also characteristic of the respiratory tract and the female reproductive tract.
In routine light microscopy, mucous cells are most conspicuously distinguished by "empty"-appearing (i.e., poorly stained) apical cytoplasm and by densely-stained, basal nuclei. (More on mucous cells of the gastro-intestinal system.)
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Ducts are relatively simple tubular structures which are (usually) easily distinguished from blood vessels by their conspicuous cuboidal to columnar epithelial lining. Blood vessels, in contrast, are lined by simple squamous endothelium.
Cells which comprise ducts generally receive much less attention than those which actually secrete the gland's product. However, complete understanding of a gland requires some awareness of and attention to the duct system through which the gland drains. Ducts are not just passive "plumbing." Some duct segments actively modify the secretory product passing through, concentrating it by removing water.
In general, cells lining ducts may often be distinguished by one or more of the following:
- cytoplasm rather pale (relative to most serous secretory cells);
- nucleus centrally located (as opposed to somewhat basal for most mucous secretory cells).
- basal and apical cytoplasm not obviously differentiated (but this isn't always obvious for secretory cells either, and striated ducts do have specialized basal cytoplasm);
- cells relatively short (cuboidal) relative to many (but not all) secretory cells (and larger ducts may be lined by columnar cells);
For the purpose of describing duct structure and function, especially in compound glands which include branching ducts of various sizes and appearances, some special terminology can be useful. (By and large, the distinctions that these terms allow represent minor details rather than essential knowledge.)
Intercalated / Striated
Intercalated ducts are small ducts which drain individual secretory units. These are usually inconspicuous, lined by a simple epithelium consisting of low cuboidal cells.
In some glands, intercalated ducts lead to striated ducts lined by a simple epithelium consisting of conspicuous cuboidal to columnar cells. In the basal cytoplasm of these cells, fine striations are visible at high magnification.
The cells of the striated ducts are specialized for concentrating the secretory product that is flowing through the duct. They do this by pumping water and ions across the duct epithelium, from the duct lumen and into interstitial fluid. This striated duct function is carried to extremes in the proximal and distal tubules of the kidney.
Ultrastructurally, striated duct cells display extensive infoldings of the basal membrane. These folds are closely associated with mitochondria that provide ATP for the membrane pumps. In light microscopy, the basal folds and mitochondria are sometimes visible as basal striations, hence the name striated duct.
Secretory / Excretory
Both intercalated and striated ducts are sometimes called secretory ducts. They are located within lobules (intralobular). More distal ducts (interlobular), sometimes called excretory ducts, are generally passive conducting tubes. Their size varies, depending on how many branches have converged proximally. Larger excretory ducts may be lined by stratified cuboidal epithelium.
It is sometimes convenient to refer to ducts by location within the gland. The following terms are all directly descriptive. Intra- means within. Inter- means between. Lobes and lobules are clusters of secretory units served, respectively, by major and minor branches of the duct tree. Within a lobule, individual secretory units are separated from one another by little more than basement membranes and capillaries. In contrast, the stroma which separates lobules and lobes consists of thicker septa of connective tissue. (The distinction between lobes and lobules is arbitrary; lobes are evident upon gross inspection while lobules are evident to low power microscopy.)
Intralobular -- Located within lobules, with no more connective tissue intervening between ducts and secretory units (i.e., acini or tubules) than between adjacent secretory units. Intercalated and striated ducts are intralobular.
Interlobular -- Located between lobules, within the thin connective tissue septa that separate lobules. All interlobular ducts are excretory.
Interlobar -- Located between lobes, within conspicuous, thick connective tissue septa that separate lobes. All interlobar ducts are excretory.
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Parenchyma / Stroma
The parenchyma of an organ consists of those cells which carry out the specific function of the organ and which usually comprise the bulk of the organ. Stroma is everything else -- connective tissue, blood vessels, nerves, and ducts.
In most glands, the parenchyma consists of secretory epithelial cells. However, the parenchyma / stroma distinction can be convenient for describing not only glands but also other organs and even tumors. Examples:
- Cardiac muscle cells comprise the parenchyma of the heart. Everything else is stroma.
- Nephrons comprise the parenchyma of the kidney. Everything else is stroma.
- Hepatocytes comprise the parenchyma of the liver. Everything else is stroma.
- Neurons and glia comprise the parenchyma of the brain. Everything else is stroma.
- Cancer cells comprise the parenchyma of malignant neoplasms. Everything else is stroma.
Because parenchyma often seems more interesting, stroma is commonly ignored as just boring background tissue. But no organ can function without the mechanical and nutritional support provided by the stroma. In any gland, connective tissue and capillaries of the stroma envelop every acinus, tubule, or cord, although they are often inconspicuous.
Pay attention to the stroma. If an organ is inflamed, the signs of inflammation appear first in the stroma.
Historical note: Ignoring inconspicuous tissue features can have consequences. Stromal capillaries are seldom evident in tissue specimens. Nothing calls them to one's attention, so they are often ignored and forgotten. Unfortunately, just such inattention may have delayed for decades the realization that interfering with tumor vasculature might powerfully inhibit tumor growth.
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SOME EXAMPLES of ordinary exocrine glands
Note: As with all topics in this website, this sample of glands is not comprehensive.
See Other Tissues for a list of glands (and other anatomical features) which are not described on these pages.
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Comments and questions: firstname.lastname@example.org
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of Medicine / Anatomy / David
Last updated: 24 January 2023 / dgk