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A unique anatomical structure of mammals that secretes milk for the nourishment of the newborn. The mammary gland contains thousands of milk-producing units called alveoli, each of which consists of a unicellular layer of epithelial cells arranged in a spheroid structure. The alveolar epithelial cells take up a variety of nutrients from the blood that perfuses the outer surface of the alveolar structures. Some of the nutrients are then secreted directly into the alveolar lumen; other nutrients are used to synthesize the unique constituents of milk which are then secreted. Each alveolus is connected to a duct through which milk flows. The ducts from many alveoli are connected via a converging ductal system which opens externally by way of the lactiferous pore.
Surrounding each alveolus and its associated small ducts are smooth muscle cells called myoepithelial cells. These cells contract in response to the posterior pituitary hormone oxytocin; milk is thus forced out of the alveoli, through the ductal system, and out the lactiferous pore for the nourishment of the newborn. The release of oxytocin is a neuroendocrine reflex triggered by the stimulation of sensory receptors by the suckling of the newborn.
Mammary glands are basically highly modified and specialized sebaceous glands which derive from ectoderm. In the embryo, mammary lines, formed on both sides of the midventral line, mark the location of future mammary glands. Along the mammary lines discrete ectodermal ingrowths, called mammary buds, produce a rudimentary branched system of ducts at birth. In all species (except the monotremes) a nipple or teat develops in concert with the mammary buds. In the most primitive mammal (the duckbill or platypus), which lacks nipples or teats, milk simply oozes out of the two mammary gland areas and is lapped up by the young.
From birth to sexual maturity the mammary gland consists of a nipple and a rudimentary ductal system in both males and females. At the onset of puberty in the female, the enhanced secretion of estrogen causes a further development of the mammary ductal system and an accumulation of lipids in fat cells. After puberty in women, the mammary gland consists of about 85% fat cells and a partially developed ductal system. See also Estrogen.
During pregnancy the mammary gland comes under the influence of estrogen and progesterone which are derived from both the ovary and placenta. These hormones cause a further branching of the ductal system and the development of milk-secreting structures, the alveoli. In humans, approximately 200 alveoli are surrounded by a connective tissue sheath forming a structure called a lobule. About 26 lobules are packaged via another connective tissue sheath into a larger structure called a lobe. Each of 15–20 lobes is exteriorized into the nipple via separate lactiferous pores. See also Progesterone.
A complement of hormones maximizes the development of the ductal andlobuloalveolar elements in the mammary gland. Optimal ductal growth is attainedwith estrogen, a glucocorticoid, prolactin, and insulin. Maximal lobuloalveolargrowth is obtained with estrogen, progesterone, growth hormone, prolactin, aglucocorticoid, and insulin. During pregnancy estrogen and progesteronestimulate mammary development but inhibit milk production.
During the final third of pregnancy, the alveolar epithelial cells beginsecreting a fluid called colostrum. This fluid fills the alveoli and causes agradual enlargement of the breast or udder. At parturition, the inhibitoryinfluence of estrogen and progesterone is removed, and the gland can secretemilk under the influence of a further complement of hormones includingprolactin, a glucocorticoid, insulin, and the thyroid hormones. See also Gland; Lactation; Mammalia; Milk; Pregnancy.
In humans, there is one pair of mammary glands, also known as mammae, or breasts. They are rudimentary in both sexes until the age of puberty when, in response to ovarian hormones, they begin to develop in the female. During pregnancy, they distend still further in preparation for nursing the infant. Pregnant women are prevented from lactating (producing milk) by the presence in the blood of high levels of estrogen and progesterone, secreted by the placenta until birth occurs.
After birth, response to prolactin, the milk-stimulating hormone, is no longer inhibited by placental hormones, and lactation begins. Mammary tissue contains between 15 and 20 compartments called lobes, each of which is divided into smaller compartments called lobules. The lobes and lobules are connected by a network of tubes whose cells manufacture the liquid and fatty substances that form milk. The tubes of each lobe connect with a duct, and all ducts lead to the nipple, where the milk is secreted when the nipple is sucked by the young. The letdown of milk during the nursing process is aided by oxytocin, a hormone secreted by the pituitary. The physical force of an infant's sucking on the breast is a major stimulus to milk production. Disorders of the mammary gland include mastitis and breast cancer.
| mammalian artificial chromosome, mammal, mamma | |
| mammotropic, mammotropic hormone, mammotropin |

| Mammary gland in a human female | |
|---|---|
| Cross section of the breast of a human female. | |
| Dissection of a lactating breast. 1 – Fat 2 – Lactiferous duct/lobule 3 – Lobule 4 – Connective tissue 5 – Sinus of lactiferous duct 6 – Lactiferous duct |
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| Latin | glandula mammaria |
| Gray's | subject #271 1267 |
A mammary gland is an organ in female mammals that produces milk to feed young offspring. Mammals get their name from the word "mammary". In humans, the mammary glands are situated on the breasts. In ruminants such as cows, goats, and deer, the mammary glands are contained in their udders. The mammary glands of other mammals that have more than two breasts, such as dogs and cats, are sometimes called dugs.
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A mammary gland is a specific type of apocrine gland specialized for manufacture of colostrum at the time of parturition. Mammary glands can be identified as apocrine because they exhibit striking "decapitation" secretion. Whether mammary glands are modified sweat glands or sebaceous glands still remains controversial.[1]
The basic components of a mature mammary gland are the alveoli (hollow cavities, a few millimetres large) lined with milk-secreting cuboidal cells and surrounded by myoepithelial cells. These alveoli join up to form groups known as lobules, and each lobule has a lactiferous duct that drains into openings in the nipple. The myoepithelial cells can contract under the stimulation of oxytocin thereby excreting milk secreted from alveolar units into the lobule lumen toward the nipple, where it collects in sinuses of the ducts. As the infant begins to suck, the hormonally (oxytocin) mediated "let down reflex" ensues and the mother's milk is secreted – not sucked from the gland – into the baby's mouth.
All the milk-secreting tissue leading to a single lactiferous duct is called a "simple mammary gland"; a "complex mammary gland" is all the simple mammary glands serving one nipple. Humans normally have two complex mammary glands, one in each breast, and each complex mammary gland consists of 10–20 simple glands. The presence of more than two nipples is known as polythelia and the presence of more than two complex mammary glands as polymastia.
To keep the correct polarized morphology of the lactiferous duct tree requires another essential component – mammary epithelial cells extracellular matrix (ECM), which together with adipocytes, fibroblast, inflammatory cells etc. constitute mammary stroma.[2] Mammary epithelial ECM mainly contains myoepithelial basement membrane and the connective tissue. They not only help to support mammary basic structure, but also serve as a communicating bridge between mammary epithelials and their local and global environment throughout this organ's development.[3] [4]
Mammary glands develop during different growth cycles. They exist in both sexes during embryonic stage, forming only a rudimentary duct tree at birth. In this stage, mammary gland development depends on systemic (and maternal) hormones,[2] but is also under the (local) regulation of paracrine communication between neighboring epithelial and mesenchymal cells by parathyroid hormone-related protein(PTHrP).[5] This locally secreted factor gives rise to a series of outside-in and inside-out positive feedback between these two types of cells, so that mammary bud epithelial cells can get to proliferate and sprout down into the mesenchymal layer until they reach the fat pad to begin the first round of branching.[2] At the same time, the embryonic mesenchymal cells around the epithelial bud get secrecting factors activated by PTHrP, such as BMP4, can transform into a dense, mammary-specific mesenchyme, which later develop into connective tissue with fibrous threads, forming blood vessels and the lymph system.[6] Basement membrane, mainly containing laminin and collagen, formed thereafter by differentiated myoepithelial cells keeps the polarity of this primary duct tree.
Lactiferous duct development occurs in females in response to circulating hormones, a first development is frequently seen during pre- and postnatal stages and later during puberty. Estrogen promotes branching differentiation,[7] whereas in males testosterone inhibits it. A mature duct tree reaching the limit of the fat pad of the mammary gland comes into being by bifurcation of duct terminal end buds (TEB), secondary branches sprouting from primary ducts[3][8] and proper duct lumen formation. These processes are tightly modulated by components of mammary epithelial ECM interacting with systemic hormones and local secreting factors. However, for each mechanism the epithelial cells' "niche" can be delicately unique with different membrane receptor profiles and basement membrane thickness from specific branching area to area, so as to regulate cell growth or differentiation sub-locally.[9] Important players include beta-1 integrin, epidermal growth factor receptor (EGFR), laminin-1/5, collagen-IV, matrix metalloproteinase(MMPs), heparan sulfate proteoglycans etc. Elevated circulating level of growth hormone and estrogen get to multipotent cap cells on tip of TEB through a leaky thin layer of basement membrance and promote specific gene expression. Hence cap cells can differentiate into myoepithelial and luminal (duct) epithelial cells, and the increased amount of activated MMPs can degrade surrounding ECM helping duct buds to reach further in the fat pads.[10][11] On the other hand, basement membrane along the mature mammary ducts is thicker with strong adhesion to epithelial cells via binding to integrin and non-integrin receptors. When side branches develop, it is a much more “pushing-forward” working process including extending through myoepithelial cells, degrading basement membrane and then invading into a periductal layer of fibrous stromal tissue.[3] Degraded basement membrane fragments (laminin-5) roles to lead the way of mammary epithelial cells migration.[12] Whereas, laminin-1 interacts with non-integrin receptor dystroglycan negatively regulates this side branching process in case of cancer.[13] These complex "Yin-yang" balancing crosstalks between mammary ECM and epithelial cells "instruct" healthy mammary gland development until adult.
Secretory alveoli develop mainly in pregnancy, when rising levels of prolactin, estrogen and progesterone cause further branching, together with an increase in adipose tissue and a richer blood flow. In gestation, serum progesterone remains at a stably high concentration so signaling through its receptor is continuously activated. As one of the transcribed genes, Wnts secreted from mammary epithelial cells act paracrinely to induce more neighboring cells branching.[14][15] When the lactiferous duct tree is almost ready, "leaves" alveoli are differentiated from luminal epithelial cells and added at the end of each branch. In late pregnancy and for the first few days after giving birth, colostrum is secreted. Milk secretion (lactation) begins a few days later due to reduction in circulating progesterone and the presence of another important hormone prolactin, which mediates further alveologenesis, milk protein production, and regulates osmotic balance and tight junction function. Laminin and collagen in myoepithelial basement membrane interacting with beta-1 integrin on epithelial surface again, is essential in this process.[16][17] Their binding ensures correct placement of prolactin receptors on basal lateral side of alveoli cells and directional secretion of milk into lactiferous ducts.[16][17] Suckling of the baby causes release of hormone oxytocin which stimulates contraction of the myoepithelial cells. In this way of combined control from ECM and systemic hormones, milk secretion can be reciprocally amplified so as to provide enough nutrition for the baby.
During weaning, decreased prolactin, missing mechanical stimulation (baby suckling) and changes in osmotic balance caused by milk stasis and leaking of tight junctions cause cessation of milk production. In some species there is complete or partial involution of alveolar structures after weaning, in humans there is only partial involution and the level of involution in humans appears to be highly individual. In some other species (such as cows) all alveoli and secretory duct structure collapse by programmed cell death (apoptosis) and autophagy for lack of growth promoting factors either from the ECM or circulating hormones.[18][19] At the same time, apoptosis of blood capillary endothelial cells speeds up the regression of lactation ductal beds. Shrinkage of the mammary duct tree and ECM remodeling by various proteinase is under the control of somatostatin and other growth inhibiting hormones and local factors.[20] This big structure change leads loose fat tissue to fill up the empty space thereafter. But a functional lactiferous duct tree can be formed again when a female is pregnant again.
Tumorigenesis in mammary glands can be induced biochemically by abnormal expression level of circulating hormones or local ECM components,[21] or from a mechanical change in the tension of mammary stroma.[22] Under either of the two circumstances, mammary epithelial cells would grow out of control and eventually result in cancer. Almost all instances of breast cancer originate in the lobules or ducts of the mammary glands.
The constantly protruding breasts of the adult human female, unusually large relative to body size, are a unique evolutionary development whose purpose is not yet fully known (see breasts); other mammals tend to have less conspicuous mammary glands that protrude only while actually filling with milk. The number and positioning of complex and simple mammary glands varies widely in different mammals. The nipples and glands can occur anywhere along the two milk lines, two roughly-parallel lines along the ventral aspect of the body. In general most mammals develop mammary glands in pairs along these lines, with a number approximating the number of young typically birthed at a time. The number of nipples varies from 2 (in most primates) to 18 (in pigs). The Virginia Opossum has 13, one of the few mammals with an odd number.[23][24] The following table lists the number and position of glands normally found in a range of mammals:
| Species [25] | Anterior (thoracic) |
Intermediate (abdominal) |
Posterior (inguinal) |
Total |
|---|---|---|---|---|
| Goat, sheep, horse guinea pig |
0 | 0 | 2 | 2 |
| Cattle | 0 | 0 | 4 | 4 |
| Cat | 2 | 2 | 4 | 8 |
| Dog [26] | 4 | 2 | 2 or 4 | 8 or 10 |
| Mouse | 6 | 0 | 4 | 10 |
| Rat | 6 | 2 | 4 | 12 |
| Pig | 6 | 6 | 6 | 18 |
| proboscideans, primates | 2 | 0 | 0 | 2 |
Male mammals typically have rudimentary mammary glands and nipples, with a few exceptions: male mice don't have nipples, and male horses lack nipples and mammary glands.[citation needed] The male Dayak fruit bat has lactating mammary glands;[27] male lactation occurs infrequently in some species, including humans.
Mammary glands are true protein factories, and several labs have constructed transgenic animals, mainly goats and cows, in order to produce proteins for pharmaceutical use.[28] Complex glycoproteins such as monoclonal antibodies or antithrombin cannot be produced by genetically engineered bacteria, and the production in live mammals is much cheaper than the use of mammalian cell cultures.
It is believed that the mammary gland is a transformed sweat gland, more closely related to Apocrine sweat glands.[29] There are many theories of how they evolved, but since they do not fossilize well, supporting such theories is difficult. Many of the current theories are based on comparisons between lines of living mammals- monotremes, marsupials and eutherians. One theory proposes that mammary glands evolved from glands that were used to keep the eggs of early mammals moist[30][31] and free from infection[32][33] (monotremes still lay eggs). Other theories propose that early secretions were used directly by hatched young,[34] or that the secretions were used by young to help them orient to their mothers.[35]
Lactation developed long before the evolution of the mammary gland and mammals, see evolution of lactation.
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Dansk (Danish)
n. - kvindebryst, brystkasse
v. tr. - tage kampen op imod, trodse
idioms:
Nederlands (Dutch)
borst, boezem, ribbenkast, confronteren, heuveltop bereiken
Français (French)
n. - sein, poitrine, poitrail, devant (d'une chemise), (Tech) ventre (de haut-fourneau), (Minér) front de taille ou d'abattage
v. tr. - affronter, faire front à, gravir (une colline)
idioms:
Deutsch (German)
n. - Brust
v. - trotzen
idioms:
Ελληνική (Greek)
n. - θώρακας, στέρνο, στήθος, μαστός, βυζί
v. - αντιμετωπίζω άφοβα, αψηφώ, τα βγάζω πέρα, σκαρφαλώνω στην κορυφή (υψώματος)
idioms:
Italiano (Italian)
petto, seno
idioms:
Português (Portuguese)
n. - mama (f)
v. - enfrentar, aplicar ou opor o peito
idioms:
Русский (Russian)
грудь, противиться
idioms:
Español (Spanish)
n. - pecho, seno, mama
v. tr. - acometer de frente, dar pecho, amamantar
idioms:
Svenska (Swedish)
n. - bröst, barm, bringa, gå rätt emot, möta, trotsa
中文(简体)(Chinese (Simplified))
乳房, 胸膛, 胸部, 胸脯, 以胸对着, 对付
idioms:
中文(繁體)(Chinese (Traditional))
n. - 乳房, 胸膛, 胸部, 胸脯
v. tr. - 以胸對著, 對付
idioms:
한국어 (Korean)
n. - 흉부, 유방, 마음
v. tr. - ~을 가슴으로 받다, ~에 과감히 맞서다
日本語 (Japanese)
n. - 乳房, 胸, 胸部, 胸中
v. - 立ち向かう, 胸に受ける
idioms:
العربيه (Arabic)
(الاسم) صدر, نهد, ثدي, واجه, جابه
עברית (Hebrew)
n. - חזה, שדיים, בגד חזה
v. tr. - התייצב מול, נאבק עם, דחף בחזה, נגע בחזהו ב-
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