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Mammary gland

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1427: 3854: 1371: 1399: 38: 1343: 1413: 1357: 439: 1441: 447: 493:). 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 proliferate and sprout down into the mesenchymal layer until they reach the fat pad to begin the first round of branching. At the same time, the embryonic mesenchymal cells around the epithelial bud receive secreting factors activated by 1385: 1331: 1312:. One theory proposes that mammary glands evolved from glands that were used to keep the eggs of early mammals moist and free from infection (monotremes still lay eggs). Other theories suggest that early secretions were used directly by hatched young, or that the secretions were used by young to help them orient to their mothers. 545:(ER) expression in mammary gland tissue, which is a downstream effect of mammary gland GHR activation. In any case, unlike IGF-1, GH itself is not essential for mammary gland development, and IGF-1 in conjunction with estrogen can induce normal mammary gland development without the presence of GH. In addition to IGF-1, other 346:) hanging from it. For instance, cows and buffalo udders have two pairs of mammary glands and four teats, whereas sheep and goat udders have one pair of mammary glands with two teats protruding from the udder. Each gland produces milk for a single teat. These mammary glands are evolutionarily derived from sweat glands. 338:. In the case of prototherians, both males and females have functional mammary glands, but their mammary glands are without nipples. These mammary glands are modified sebaceous glands. Concerning metatherians and eutherians, only females have functional mammary glands. Their mammary glands can be termed as breasts or 962:
in mammary glands can be induced biochemically by abnormal expression level of circulating hormones or local ECM components, or from a mechanical change in the tension of mammary stroma. Under either of the two circumstances, mammary epithelial cells would grow out of control and eventually result in
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of alveolar structures after weaning, in humans there is only partial involution and the level of involution in humans appears to be highly individual. The glands in the breast do secrete fluid also in nonlactating women. In some other species (such as cows), all alveoli and secretory duct structures
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secreted from mammary epithelial cells act paracrinely to induce more neighboring cells' branching. 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
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on epithelial surface again, is essential in this process. Their binding ensures correct placement of prolactin receptors on the basal lateral side of alveoli cells and directional secretion of milk into lactiferous ducts. Suckling of the baby causes release of the hormone oxytocin, which stimulates
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By the pubertal stage, the mammary ducts have invaded to the end of the mammary fat pad. At this point, the terminal end buds become less proliferative and decrease in size. Side branches form from the primary ducts and begin to fill the mammary fat pad. Ductal development decreases with the arrival
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During embryonic development, IGF-1 levels are low, and gradually increase from birth to puberty. At puberty, the levels of GH and IGF-1 reach their highest levels in life and estrogen begins to be secreted in high amounts in females, which is when ductal development mostly takes place. Under the
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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. It is the (passive) process of a child or animal ceasing to be dependent on the mother for
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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. On the other hand, basement membrane along the mature mammary ducts is thicker, with strong adhesion to epithelial cells via binding to
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All the milk-secreting tissue leading to a single lactiferous duct is collectively called a "simple mammary gland"; in a "complex mammary gland", all the simple mammary glands serve one nipple. Humans normally have two complex mammary glands, one in each breast, and each complex mammary gland
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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. Degraded basement membrane fragments
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Mammary glands develop during different growth cycles. They exist in both sexes during the embryonic stage, forming only a rudimentary duct tree at birth. In this stage, mammary gland development depends on systemic (and maternal) hormones, but is also under the (local) regulation of
805:(TEB), secondary branches sprouting from primary ducts 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' " 669:
mammary gland development can be divided into a series of specific stages. Initially, the formation of the milk lines that run between the fore and hind limbs bilaterally on each side of the midline occurs around embryonic day 10.5 (E10.5). The second stage occurs at E11.5 when
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component of mammary gland development, and act synergistically to mediate it. Neither estrogen nor GH are capable of inducing ductal development without the other. The role of GH in ductal development has been found to be mostly mediated by its induction of the secretion of
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The breasts of female humans vary from most other mammals that tend to have less conspicuous mammary glands. The number and positioning of mammary glands varies widely in different mammals. The protruding teats and accompanying glands can be located anywhere along the two
708:. Terminal end buds, the highly proliferative structures found at the tips of the invading ducts, expand and increase greatly during this stage. This developmental period is characterized by the emergence of the terminal end buds and lasts until an age of about 7–8 weeks. 1937:
Wysolmerski, J. J.; Philbrick, W. M.; Dunbar, M. E.; Lanske, B.; Kronenberg, H.; Broadus, A. E. (1998). "Rescue of the parathyroid hormone-related protein knockout mouse demonstrates that parathyroid hormone-related protein is essential for mammary gland development".
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Nicholas L. Petrakis; Lynn Mason; Rose Lee; Barbara Sugimoto; Stella Pawson; Frank Catchpool (1975). "Association of Race, Age, Menopausal Status, and Cerumen Type With Breast Fluid Secretion in Nonlactating Women, as Determined by Nipple Aspiration".
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and other growth inhibiting hormones and local factors. This major structural change leads loose fat tissue to fill the empty space afterward. But a functional lactiferous duct tree can be formed again when a female is pregnant again.
501:. These mesenchymal cells can transform into a dense, mammary-specific mesenchyme, which later develop into connective tissue with fibrous threads, forming blood vessels and the lymph system. A basement membrane, mainly containing 985:. 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 teats varies from 2 (in most primates) to 18 (in pigs). The 809:" 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. Important players include 429:
and the connective tissue. They not only help to support mammary basic structure, but also serve as a communicating bridge between mammary epithelia and their local and global environment throughout this organ's development.
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Petrakis NL, Barnes S, King EB, Lowenstein J, Wiencke J, Lee MM, Miike R, Kirk M, Coward L (October 1996). "Stimulatory influence of soy protein isolate on breast secretion in pre- and postmenopausal people AFAB".
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Developmentally, mammary gland epithelium is constantly produced and maintained by rare epithelial cells, dubbed as mammary progenitors which are ultimately thought to be derived from tissue-resident stem cells.
509:, formed afterward by differentiated myoepithelial cells, keeps the polarity of this primary duct tree. These components of the extracellular matrix are strong determinants of duct morphogenesis. 342:. In the case of breasts, each mammary gland has its own nipple (e.g., human mammary glands). In the case of udders, pairs of mammary glands comprise a single mass, with more than one nipple (or 1300:. Because mammary glands do not fossilize well, supporting such theories with fossil evidence is difficult. Many of the current theories are based on comparisons between lines of living mammals— 916:
contraction of the myoepithelial cells. In this combined control from ECM and systemic hormones, milk secretion can be reciprocally amplified so as to provide enough nutrition for the baby.
720:(proestrus, estrus, metestrus, and diestrus). As a result of estrous cycling, the mammary gland undergoes dynamic changes where cells proliferate and then regress in an ordered fashion. 462:(first milk) when giving birth. Mammary glands can be identified as apocrine because they exhibit striking "decapitation" secretion. Many sources assert that mammary glands are modified 3336:
Muschler, J.; Levy, D.; Boudreau, R.; Henry, M.; Campbell, K.; Bissell, M. J. (2002). "A role for dystroglycan in epithelial polarization: Loss of function in breast tumor cells".
704:, the mammary ducts elongate into the mammary fat pad. Then, starting around four weeks of age, mammary ductal growth increases significantly with the ducts invading towards the 772:, due to much lower levels of estrogen, and due to lower levels of GH and IGF-1, which decrease with age, mammary gland tissue atrophies and the mammary glands become smaller. 3528:"Control of mammary epithelial differentiation: Basement membrane induces tissue-specific gene expression in the absence of cell-cell interaction and morphological polarity" 3733:
Motyl, T.; Gajkowska, B.; Zarzyńska, J.; Gajewska, M.; Lamparska-Przybysz, M. (2006). "Apoptosis and autophagy in mammary gland remodeling and breast cancer chemotherapy".
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Light micrograph of a human proliferating mammary gland during estrous cycle. Sprouting gland tissue can be seen in the upper left field (haematoxylin eosin staining)
3762:"Normal and tumor-derived myoepithelial cells differ in their ability to interact with luminal breast epithelial cells for polarity and basement membrane deposition" 402:
consists of 10–20 simple glands. The opening of each simple gland on the surface of the nipple is called a "pore." The presence of more than two nipples is known as
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surrounding the ductal system in the mammary glands also grows. After puberty, GH and IGF-1 levels progressively decrease, which limits further development until
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cap cells on TEB tips through a thin, leaky layer of basement membrane. These hormones promote specific gene expression. Hence cap cells can differentiate into
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There are many theories on how mammary glands evolved. For example, it is thought that the mammary gland is a transformed sweat gland, more closely related to
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and the presence of another important hormone prolactin, which mediates further alveologenesis, milk protein production, and regulates osmotic balance and
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expressed in mammary gland tissue, and in conjunction with relatively low estrogen levels, are the cause of the lack of developed mammary glands in males.
890:, serum progesterone remains at a stably high concentration so signaling through its receptor is continuously activated. As one of the transcribed genes, 1514: 989:
has 13, one of the few mammals with an odd number. The following table lists the number and position of teats and glands found in a range of mammals:
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formation begins along the mammary milk line. This will eventually give rise to the nipple. Lastly, the third stage occurs at E12.5 and involves the
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begins to suck, the oxytocin-mediated "let down reflex" ensues, and the mother's milk is secreted—not sucked—from the gland into the infant's mouth.
2681:"Breast size in relation to endogenous hormone levels, body constitution, and oral contraceptive use in healthy nulligravid women aged 19–25 years" 859:. These complex "Yin-yang" balancing crosstalks between mammary ECM and epithelial cells "instruct" healthy mammary gland development until adult. 4405:
Lefèvre, C. M.; Sharp, J. A.; Nicholas, K. R. (2010). "Evolution of Lactation: Ancient Origin and Extreme Adaptations of the Lactation System".
4357: 3965:"Conversion of the Nipple to Hair-Bearing Epithelia by Lowering Bone Morphogenetic Protein Pathway Activity at the Dermal-Epidermal Interface" 3870: 1487: 196: 413:
Maintaining the correct polarized morphology of the lactiferous duct tree requires another essential component – mammary epithelial cells
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Wiseman, B. S.; Sternlicht, M. D.; Lund, L. R.; Alexander, C. M.; Mott, J.; Bissell, M. J.; Soloway, P.; Itohara, S.; Werb, Z. (2003).
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Sekhri, KK; Pitelka, DR; Deome, KB (September 1967). "Studies of mouse mammary glands. I. Cytomorphology of the normal mammary gland".
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Brisken, C.; Heineman, A.; Chavarria, T.; Elenbaas, B.; Tan, J.; Dey, S. K.; McMahon, J. A.; McMahon, A. P.; Weinberg, R. A. (2000).
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for lack of growth promoting factors either from the ECM or circulating hormones. At the same time, apoptosis of blood capillary
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Watson, C. J.; Khaled, W. T. (2008). "Mammary development in the embryo and adult: A journey of morphogenesis and commitment".
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Male mammals typically have rudimentary mammary glands and nipples, with a few exceptions: male mice do not have nipples, male
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inhibits it. A mature duct tree reaching the limit of the fat pad of the mammary gland comes into being by bifurcation of duct
2228:"Insulin-like growth factor I is essential for terminal end bud formation and ductal morphogenesis during mammary development" 4489: 4023: 3921: 3809:
Provenzano, P. P.; Inman, D. R.; Eliceiri, K. W.; Knittel, J. G.; Yan, L.; Rueden, C. T.; White, J. G.; Keely, P. J. (2008).
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Kleinberg DL, Feldman M, Ruan W (2000). "IGF-I: an essential factor in terminal end bud formation and ductal morphogenesis".
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surrounding the alveoli will cause the milk to be ejected through the ducts and into the nipple for the nursing infant. Upon
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Graves, B. M.; Duvall, D. (1983). "A Role for Aggregation Pheromones in the Evolution of Mammallike Reptile Lactation".
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Vorbach, C.; Capecchi, M. R.; Penninger, J. M. (2006). "Evolution of the mammary gland from the innate immune system?".
3198:"Site-specific inductive and inhibitory activities of MMP-2 and MMP-3 orchestrate mammary gland branching morphogenesis" 2904:"Key stages of mammary gland development: molecular mechanisms involved in the formation of the embryonic mammary gland" 1975:"Key stages of mammary gland development: Molecular mechanisms involved in the formation of the embryonic mammary gland" 1507: 3296:"Cell adhesion to laminin 1 or 5 induces isoform-specific clustering of integrins and other focal adhesion components" 3899: 2722:"Testosterone inhibits estrogen-induced mammary epithelial proliferation and suppresses estrogen receptor expression" 1921: 1875: 1582: 942:
speeds up the regression of lactation ductal beds. Shrinkage of the mammary duct tree and ECM remodeling by various
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Streuli, C. H.; Schmidhauser, C.; Bailey, N.; Yurchenco, P.; Skubitz, A. P.; Roskelley, C.; Bissell, M. J. (1995).
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inhibit estrogen-mediated mammary gland development (e.g., by reducing local ER expression) through activation of
425:, inflammatory cells, and others, constitute mammary stroma. Mammary epithelial ECM mainly contains myoepithelial 4175:
Li, Peng; Knabe, Darrell A.; Kim, Sung Woo; Lynch, Christopher J.; Hutson, Susan M.; Wu, Guoyao (1 August 2009).
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Dog breeds vary in the number of mammary glands: larger breeds tend to have 5 pairs, smaller breeds have 4 pairs.
557: 4177:"Lactating Porcine Mammary Tissue Catabolizes Branched-Chain Amino Acids for Glutamine and Aspartate Synthesis" 1257: 215: 179: 3147:"Regulation of mammary gland branching morphogenesis by the extracellular matrix and its remodeling enzymes" 4546: 541:(GHR). However, GH itself also acts independently of IGF-1 to stimulate ductal development by upregulating 530: 693:
The primitive (stem) cells are detected in embryo and their numbers increase steadily during development
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Kleinberg DL, Ruan W (2008). "IGF-I, GH, and sex steroid effects in normal mammary gland development".
2026:"Modeling mammary organogenesis from biological first principles: Cells and their physical constraints" 1315:
Lactation is thought to have developed long before the evolution of the mammary gland and mammals; see
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Gudjonsson, T.; Rønnov-Jessen, L.; Villadsen, R.; Rank, F.; Bissell, M. J.; Petersen, O. W. (2002).
4687: 2620:"The potential clinical applications of insulin-like growth factor-1 ligand in human breast cancer" 826: 569: 565: 126: 4481:
Histologic Diagnosis of Inflammatory Skin Diseases An Algorithmic Method Based on Pattern Analysis
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Tharmapalan, Pirashaanthy; Mahendralingam, Mathepan; Berman, Hal K; Khokha, Rama (15 July 2019).
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Newton, Michael; Newton, Niles Rumely (December 1948). "The let-down reflex in human lactation".
553: 538: 296:, the occasional production of milk by the glands, can occur in any mammal, but in most mammals, 130: 51: 3047:"Key stages in mammary gland development: The cues that regulate ductal branching morphogenesis" 1398: 1370: 394:, excreting the milk secreted by alveolar units into the lobule lumen toward the nipple. As the 4260:
Oftedal, O. T. (2002). "The origin of lactation as a water source for parchment-shelled eggs".
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Zarzynska, J.; Motyl, T. (2008). "Apoptosis and autophagy in involuting bovine mammary gland".
3476:"Essential function of Wnt-4 in mammary gland development downstream of progesterone signaling" 1459: 830: 640: 577: 572:(HGF) are involved in breast development as mediators downstream to sex hormones and GH/IGF-1. 227: 203: 191: 158: 142: 2650: 2503: 756:. This process involves the controlled collapse of mammary epithelial cells where cells begin 4589: 2587: 2032:. From the Century of the Genome to the Century of the Organism: New Theoretical Approaches. 1297: 926: 891: 489:
between neighboring epithelial and mesenchymal cells by parathyroid hormone-related protein (
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function. Laminin and collagen in myoepithelial basement membrane interacting with beta-1
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of the infant, lactation stops and the mammary gland turns in on itself, a process called
8: 4656: 2847:"Mammary stem cells and progenitors: targeting the roots of breast cancer for prevention" 2364:"Mouse models of transforming growth factor beta impact in breast development and cancer" 1273: 745: 4418: 4110: 2585: 1777: 1667: 1342: 4640: 4630: 4457: 4340: 4285: 4211: 4176: 4122: 4067: 4040: 3989: 3964: 3837: 3810: 3786: 3761: 3715: 3604: 3579: 3552: 3527: 3456: 3295: 3271: 3246: 3222: 3197: 3122: 3097: 3073: 3046: 3027: 2979: 2954: 2930: 2903: 2879: 2846: 2795: 2770: 2751: 2697: 2680: 2563: 2530: 2435: 2408: 2344: 2301: 2203: 2155: 2102: 2077: 2050: 2025: 2001: 1974: 1843: 1818: 1794: 1761: 1737: 1684: 1651: 1356: 475: 371: 162: 3626: 3500: 3475: 3173: 3146: 1628: 4625: 4485: 4422: 4387: 4332: 4277: 4216: 4198: 4157: 4072: 4019: 3994: 3960: 3917: 3842: 3791: 3742: 3707: 3672: 3645: 3609: 3557: 3505: 3448: 3407: 3386: 3345: 3318: 3276: 3227: 3178: 3127: 3113: 3078: 3019: 2984: 2935: 2884: 2866: 2827: 2800: 2743: 2702: 2656: 2631: 2593: 2568: 2550: 2509: 2481: 2440: 2385: 2336: 2293: 2249: 2195: 2147: 2107: 2055: 2041: 2024:
Montévil, Maël; Speroni, Lucia; Sonnenschein, Carlos; Soto, Ana M. (1 October 2016).
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Daniel, CW; Smith, GH (January 1999). "The mammary gland: a model for development".
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development in estrogen-primed mammary gland tissue, which occurs in preparation of
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can occur. With humans, male lactation can occur only under specific circumstances.
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Fadok, V. A. (1999). "Clearance: The last and often forgotten stage of apoptosis".
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Oftedal, OT (2002). "The mammary gland and its origin during synapsid evolution".
847:(laminin-5) roles to lead the way of mammary epithelial cells migration. Whereas, 4692: 1900: 1819:"Adipose Stroma Induces Branching Morphogenesis of Engineered Epithelial Tubules" 1537:
Macéa, José Rafael; Fregnani, José Humberto Tavares Guerreiro (1 December 2006).
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occurs within the mammary gland; lactation involves the secretion of milk by the
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Jay R. Harris; Marc E. Lippman; C. Kent Osborne; Monica Morrow (28 March 2012).
2546: 2093: 366:(hollow cavities, a few millimeters large), which are lined with milk-secreting 4594: 4502: 4153: 4057: 3980: 1316: 1284:, and the production in live mammals is much cheaper than the use of mammalian 1246: 908: 879: 833:, and others. Elevated circulating level of growth hormone and estrogen get to 736:
structures within the branches to be used for milk production. After delivery,
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Koshikawa, N.; Giannelli, G.; Cirulli, V.; Miyazaki, K.; Quaranta, V. (2000).
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in a controlled manner, reverting the mammary gland back to a pubertal state.
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Mammary gland development is characterized by the unique process by which the
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Ackerman, A. Bernard; Almut Böer; Bruce Bennin; Geoffrey J. Gottlieb (2005).
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intake mildly stimulates the breast glands in pre- and postmenopausal women.
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by Sir Astley Paston Cooper (1840). Numerous drawings, in the public domain.
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Kleinberg DL (1997). "Early mammary development: growth hormone and IGF-1".
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Kunz, T; Hosken, D (2009). "Male lactation: why, why not and is it care?".
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Lactiferous duct development occurs in females in response to circulating
438: 3777: 3491: 3443: 3426: 2531:"Mammary Gland Growth Factors: Roles in Normal Development and in Cancer" 1477: 834: 655:
which establishes the basic arboreal network of ducts emanating from the
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Human Microscopic Anatomy: An Atlas for Students of Medicine and Biology
1577:(7th ed.). Maryland Heights, Maryland: Mosby/Elsevier. p. 54. 4495: 4383: 4090:
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3580:"Laminin mediates tissue-specific gene expression in mammary epithelia" 1891: 1725: 1650:
Zucca-Matthes, Gustavo; Urban, Cícero; Vallejo, André (February 2016).
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Secretory alveoli develop mainly in pregnancy, when rising levels of
757: 737: 729: 701: 601: 593: 585: 546: 533:(IGF-1), which occurs both systemically (mainly originating from the 459: 418: 309: 297: 265: 4012:
Patricia J. Armati; Chris R. Dickman; Ian D. Hume (17 August 2006).
3811:"Collagen density promotes mammary tumor initiation and progression" 3759: 3063: 2920: 2721: 2425: 2130:
Kleinberg DL (1998). "Role of IGF-I in normal mammary development".
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in recent months or years. It is directed by hormonal guidance from
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do not have mammary glands, and male horses lack nipples. The male
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Cunningham, Merle; LaTour, Mickey A. & Acker, Duane (2005).
3473: 1936: 1762:"Stromal Effects on Mammary Gland Development and Breast Cancer" 4635: 4577: 4563: 4235:"BBC News – The goats with spider genes and silk in their milk" 3732: 3098:"Hormonal and local control of mammary branching morphogenesis" 2720:
Zhou J, Ng S, Adesanya-Famuiya O, Anderson K, Bondy CA (2000).
2023: 1362: 1061: 856: 666: 656: 537:) and locally in the mammary fat pad through activation of the 395: 387: 355: 305: 277: 257: 121: 66: 61: 3889: 3808: 3195: 2719: 3096:
Sternlicht, M. D.; Kouros-Mehr, H.; Lu, P.; Werb, Z. (2006).
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Moore, Keith L.; Dalley, Arthur F.; Agur, Anne M. R. (2018).
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Lawrence, Ruth A.; Lawrence, Robert M. (30 September 2010).
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negatively regulates this side branching process in case of
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and the presence of more than two complex mammary glands as
390:. The myoepithelial cells contract under the stimulation of 3424: 3095: 1404: 1261: 1030: 643:. The development of the mammary gland occurs mainly after 498: 343: 261: 137: 3425:
Robinson, G. W.; Hennighausen, L.; Johnson, P. F. (2000).
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nourishment. In some species there is complete or partial
903:) begins a few days later due to reduction in circulating 732:, the ductal systems undergo rapid proliferation and form 19:"Mammary" redirects here. For the mountain in Alaska, see 4089: 3335: 1816: 1649: 1390: 1376: 1348: 1265: 1156: 1137: 1099: 1080: 967:
originate in the lobules or ducts of the mammary glands.
4369: 3365:"Hormonal effects of soy in premenopausal women and men" 2768: 1817:
Pavlovich, A. L.; Manivannan, S.; Nelson, C. M. (2010).
4041:"Development and Pathology of the Equine Mammary Gland" 3293: 360:
The basic components of a mature mammary gland are the
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in primates (for example, humans and chimpanzees), the
1268:, to produce proteins for pharmaceutical use. Complex 3573: 3571: 3521: 3519: 2618:
Chong YM, Subramanian A, Sharma AK, Mokbel K (2007).
2460:"Hepatocyte growth factor profile with breast cancer" 797:
promotes branching differentiation, whereas in males
288:(for example, cows, goats, sheep, and deer), and the 4404: 3525: 2275: 1755: 1753: 1751: 3526:Streuli, C. H.; Bailey, N.; Bissell, M. J. (1991). 2592:. Lippincott Williams & Wilkins. pp. 94–. 1707: 1705: 1703: 1515:
List of distinct cell types in the adult human body
3568: 3516: 2901: 2817: 2648: 4005: 2579: 1748: 1575:Breastfeeding: A Guide for the Medical Profession 1539:"Anatomy of the Thoracic Wall, Axilla and Breast" 4674: 3400:Cancer Epidemiology, Biomarkers & Prevention 3294:Dogic, D.; Rousselle, P.; Aumailley, M. (1998). 2769:Lemaine V, Cayci C, Simmons PS, Petty P (2013). 2492: 1700: 1572: 4174: 3959:Julie Ann Mayer; John Foley; Damon De La Cruz; 3144: 3001: 2678: 2613: 2611: 2609: 2522: 2271: 2269: 2267: 2265: 2263: 2221: 2219: 2217: 2173: 2171: 2169: 1972: 1865: 292:of other animals (for example, dogs and cats). 272:, "breast". The mammary glands are arranged in 4317:Journal of Mammary Gland Biology and Neoplasia 4262:Journal of Mammary Gland Biology and Neoplasia 4045:Journal of Mammary Gland Biology and Neoplasia 3692:Journal of Mammary Gland Biology and Neoplasia 3662: 3145:Fata, J. E.; Werb, Z.; Bissell, M. J. (2003). 3004:Journal of Mammary Gland Biology and Neoplasia 2952: 2674: 2672: 2125: 2123: 2121: 4547: 2955:"Stem Cells and the Developing Mammary Gland" 2457: 2318: 2225: 2075: 2071: 2069: 1810: 1536: 1336:Cross section of the breast of a human female 1280:cannot be produced by genetically engineered 1256:factories, and several labs have constructed 374:. These alveoli join to form groups known as 268:. Mammals get their name from the Latin word 4439: 4407:Annual Review of Genomics and Human Genetics 3873:. Digitalcollections.fiu.edu. Archived from 2762: 2713: 2606: 2498: 2451: 2402: 2396: 2361: 2355: 2312: 2260: 2214: 2166: 2030:Progress in Biophysics and Molecular Biology 1884: 1711: 1614: 3952: 2902:Hens, JR; Wysolmerski JJ (10 August 2005). 2669: 2642: 2528: 2508:. Elsevier Health Sciences. pp. 413–. 2118: 1759: 4554: 4540: 4139: 3044: 2535:Cold Spring Harbor Perspectives in Biology 2082:Cold Spring Harbor Perspectives in Biology 2066: 42:Cross-section of the human mammary gland. 36: 16:Exocrine gland in humans and other mammals 4561: 4305:. Smithsonian National Zoo, 14 July 2003. 4210: 4192: 4133: 4066: 4056: 3988: 3836: 3826: 3785: 3603: 3551: 3499: 3442: 3380: 3362: 3270: 3221: 3172: 3162: 3121: 3072: 3062: 2978: 2929: 2919: 2878: 2794: 2737: 2696: 2562: 2475: 2434: 2424: 2379: 2243: 2177: 2129: 2101: 2049: 2000: 1990: 1842: 1793: 1683: 1557: 1532: 1530: 1528: 3630:Journal of the National Cancer Institute 1973:Hens, J. R.; Wysolmerski, J. J. (2005). 1905: 1859: 1652:"Anatomy of the nipple and breast ducts" 954: 851:-1 interacts with non-integrin receptor 445: 437: 4314: 4259: 4675: 4038: 3735:Journal of Physiology and Pharmacology 3665:Journal of Physiology and Pharmacology 1911: 1525: 454:A mammary gland is a specific type of 4535: 3689: 2078:"Hormone Action in the Mammary Gland" 1508:List of glands of the human body#Skin 1488:Hypothalamic–pituitary–prolactin axis 1249:occurs infrequently in some species. 458:gland specialized for manufacture of 108:(blood vessels and connective tissue) 3963:; Randall Widelitz (November 2008). 3871:"With the Wild Things – Transcripts" 2529:Hynes, N. E.; Watson, C. J. (2010). 1912:Krstic, Radivoj V. (18 March 2004). 1597: 1322: 300:, the production of enough milk for 4583:Areolar gland (gland of Montgomery) 4419:10.1146/annurev-genom-082509-141806 2505:Anatomy and Physiology for Midwives 2076:Brisken; Malley (2 December 2010). 1668:10.3978/j.issn.2227-684X.2015.05.10 1546:International Journal of Morphology 930:collapse by programmed cell death ( 862:There is preliminary evidence that 780: 744:in the alveoli. Contraction of the 651:, tubule formation is coupled with 480:Parathyroid hormone-related protein 13: 2953:Makarem, M; Eaves C (April 2013). 2771:"Gynecomastia in adolescent males" 2698:10.1093/oxfordjournals.aje.a009153 588:, if it occurs. During pregnancy, 14: 4719: 4520:Comparative Mammary Gland Anatomy 4513: 4142:Trends in Ecology & Evolution 2655:. Academic Press. pp. 770–. 1760:Wiseman, B. S.; Werb, Z. (2002). 763: 386:that drains into openings in the 3852: 3114:10.1111/j.1432-0436.2006.00105.x 2042:10.1016/j.pbiomolbio.2016.08.004 1439: 1425: 1411: 1397: 1383: 1369: 1355: 1341: 1329: 970: 963:cancer. Almost all instances of 815:epidermal growth factor receptor 4471: 4433: 4398: 4363: 4351: 4308: 4296: 4253: 4227: 4168: 4083: 4032: 3939: 3930: 3905: 3802: 3753: 3726: 3683: 3656: 3620: 3467: 3418: 3356: 3329: 3287: 3238: 3189: 3138: 3089: 3038: 2995: 2946: 2895: 2838: 2811: 2017: 1966: 1930: 1559:10.4067/S0717-95022006000500030 630: 558:transforming growth factor beta 512: 442:Normal histology of the breast. 4092:"Lactation in male fruit bats" 4018:. Cambridge University Press. 3947:Red-Sided Short-Tailed Opossum 3900:Raising Orphaned Baby Opossums 2959:J Mammary Gland Biol Neoplasia 2679:Jernström H, Olsson H (1997). 2458:El-Attar HA, Sheta MI (2011). 2321:J Mammary Gland Biol Neoplasia 2278:J Mammary Gland Biol Neoplasia 2180:J Mammary Gland Biol Neoplasia 1643: 1591: 1566: 1245:has lactating mammary glands. 869: 469: 316:. In a few mammalian species, 1: 4708:Human female endocrine system 2226:Ruan W, Kleinberg DL (1999). 1629:10.1016/S0022-3476(48)80075-2 1519: 899:is secreted. Milk secretion ( 831:heparan sulfate proteoglycans 775: 180:Pectoral axillary lymph nodes 168:(lateral and medial branches) 4526:On the anatomy of the breast 2652:Essential Medical Physiology 2362:Serra R, Crowley MR (2005). 1291: 723: 596:are essential for mediating 531:insulin-like growth factor 1 433: 349: 304:, occurs only in phenotypic 7: 4507:Clinically Oriented Anatomy 3914:Animal Science and Industry 3584:The Journal of Cell Biology 3532:The Journal of Cell Biology 3251:The Journal of Cell Biology 3202:The Journal of Cell Biology 2649:Leonard R. Johnson (2003). 2547:10.1101/cshperspect.a003186 2502:; Dunstall, Melvyn (2011). 2094:10.1101/cshperspect.a003178 1868:Clinically oriented anatomy 1453: 625: 524:(GH) are essential for the 417:(ECM) which, together with 326:are divided into 3 groups: 10: 4724: 4154:10.1016/j.tree.2008.09.009 4058:10.1007/s10911-020-09471-2 4039:Hughes, Katherine (2021). 3981:10.2353/ajpath.2008.070920 3902:. Alabama Wildlife Center. 3045:Sternlicht, M. D. (2006). 1457: 1218:Southern red-sided opossum 975: 919: 473: 353: 18: 4649: 4570: 3916:. Pearson Prentice Hall. 2971:10.1007/s10911-013-9284-6 2464:Indian J Pathol Microbiol 2333:10.1007/s10911-008-9103-7 1916:. Springer. p. 466. 1835:10.1089/ten.TEA.2009.0836 1823:Tissue Engineering Part A 1617:The Journal of Pediatrics 1601:Anatomy of the Human Body 696: 226: 214: 202: 190: 185: 173: 152: 136: 120: 96: 91: 35: 30: 4181:The Journal of Nutrition 3369:The Journal of Nutrition 3363:Kurzer MS (March 2002). 2863:10.15252/embj.2018100852 2245:10.1210/endo.140.11.7095 2132:Breast Cancer Res. Treat 1252:Mammary glands are true 946:is under the control of 827:matrix metalloproteinase 570:hepatocyte growth factor 566:fibroblast growth factor 127:Internal thoracic artery 4442:The American Naturalist 4329:10.1023/A:1022896515287 4274:10.1023/A:1022848632125 3766:Journal of Cell Science 3704:10.1023/A:1011384009787 3480:Genes & Development 3431:Genes & Development 3303:Journal of Cell Science 3016:10.1023/A:1018796301609 2477:10.4103/0377-4929.85083 2290:10.1023/A:1009507030633 2192:10.1023/A:1026373513521 2144:10.1023/a:1005998832636 1786:10.1126/science.1067431 686:, leading to a mammary 653:branching morphogenesis 576:influence of estrogen, 554:epidermal growth factor 539:growth hormone receptor 487:paracrine communication 131:Lateral thoracic artery 3898:Stockard, Mary (2005) 3828:10.1186/1741-7015-6-11 3544:10.1083/jcb.115.5.1383 3151:Breast Cancer Research 3051:Breast Cancer Research 2787:10.1055/s-0033-1347166 2589:Diseases of the Breast 1979:Breast Cancer Research 1952:10.1242/dev.125.7.1285 1460:anatomical terminology 1317:evolution of lactation 451: 443: 228:Anatomical terminology 159:Supraclavicular nerves 143:Internal thoracic vein 4194:10.3945/jn.109.105957 3642:10.1093/jnci/54.4.829 3596:10.1083/jcb.129.3.591 3382:10.1093/jn/132.3.570S 3315:10.1242/jcs.111.6.793 3263:10.1083/jcb.148.3.615 3214:10.1083/jcb.200302090 2739:10.1096/fj.99-0863com 2368:Endocr. Relat. Cancer 1899:21 April 2011 at the 1890:Ackerman (2005) ch.1 1298:apocrine sweat glands 1274:monoclonal antibodies 955:Clinical significance 474:Further information: 449: 441: 3778:10.1242/jcs.115.1.39 3671:(Suppl 9): 275–288. 3492:10.1101/gad.14.6.650 3444:10.1101/gad.14.8.889 1598:Gray, Henry (1918). 678:of cells within the 415:extracellular matrix 380:. Each lobule has a 4657:Dense breast tissue 4484:. Ardor Scribendi. 4111:1994Natur.367..691F 2381:10.1677/erc.1.00936 1778:2002Sci...296.1046W 1772:(5570): 1046–1049. 746:myoepithelial cells 372:myoepithelial cells 115:(cellular elements) 4641:Retromammary space 4631:Intermammary cleft 4590:Cooper's ligaments 4384:10.1002/bies.20423 4360:. scienceblogs.com 3741:(Suppl 7): 17–32. 2820:J Natl Cancer Inst 1726:10.1242/dev.005439 1458:This article uses 1258:transgenic animals 620:androgen receptors 476:Breast development 452: 444: 370:and surrounded by 163:Intercostal nerves 52:Pectoralis muscles 4670: 4669: 4626:Inframammary fold 4498:on 21 April 2011. 4491:978-1-893357-25-9 4358:Breast beginnings 4303:Lactating on Eggs 4241:. 17 January 2012 4105:(6465): 691–692. 4025:978-1-139-45742-2 3961:Cheng-Ming Chuong 3949:. Fauna Paraguay 3923:978-0-13-046256-5 3344:(23): 7102–7109. 2908:Breast Cancer Res 2662:978-0-12-387584-6 2599:978-1-4511-4870-1 2515:978-0-7020-3489-3 2413:Breast Cancer Res 1829:(12): 3719–3726. 1323:Additional images 1235: 1234: 940:endothelial cells 803:terminal end buds 543:estrogen receptor 427:basement membrane 242: 241: 237: 169: 116: 109: 4715: 4619:Mammary alveolus 4612:Terminal end bud 4607:Lactiferous duct 4556: 4549: 4542: 4533: 4532: 4499: 4494:. Archived from 4466: 4465: 4437: 4431: 4430: 4402: 4396: 4395: 4367: 4361: 4355: 4349: 4348: 4312: 4306: 4300: 4294: 4293: 4257: 4251: 4250: 4248: 4246: 4231: 4225: 4224: 4214: 4196: 4187:(8): 1502–1509. 4172: 4166: 4165: 4137: 4131: 4130: 4119:10.1038/367691a0 4096: 4087: 4081: 4080: 4070: 4060: 4036: 4030: 4029: 4009: 4003: 4002: 3992: 3956: 3950: 3943: 3937: 3934: 3928: 3927: 3909: 3903: 3896: 3887: 3886: 3884: 3882: 3877:on 23 March 2013 3867: 3858: 3857: 3856: 3850: 3840: 3830: 3806: 3800: 3799: 3789: 3757: 3751: 3750: 3730: 3724: 3723: 3687: 3681: 3680: 3660: 3654: 3653: 3624: 3618: 3617: 3607: 3575: 3566: 3565: 3555: 3538:(5): 1383–1395. 3523: 3514: 3513: 3503: 3471: 3465: 3464: 3446: 3422: 3416: 3415: 3394: 3384: 3375:(3): 570S–573S. 3360: 3354: 3353: 3333: 3327: 3326: 3300: 3291: 3285: 3284: 3274: 3242: 3236: 3235: 3225: 3208:(6): 1123–1133. 3193: 3187: 3186: 3176: 3166: 3142: 3136: 3135: 3125: 3093: 3087: 3086: 3076: 3066: 3042: 3036: 3035: 2999: 2993: 2992: 2982: 2950: 2944: 2943: 2933: 2923: 2899: 2893: 2892: 2882: 2851:The EMBO Journal 2842: 2836: 2835: 2815: 2809: 2808: 2798: 2775:Semin Plast Surg 2766: 2760: 2759: 2741: 2717: 2711: 2710: 2700: 2685:Am. J. Epidemiol 2676: 2667: 2666: 2646: 2640: 2639: 2615: 2604: 2603: 2583: 2577: 2576: 2566: 2526: 2520: 2519: 2496: 2490: 2489: 2479: 2455: 2449: 2448: 2438: 2428: 2400: 2394: 2393: 2383: 2359: 2353: 2352: 2316: 2310: 2309: 2273: 2258: 2257: 2247: 2223: 2212: 2211: 2175: 2164: 2163: 2127: 2116: 2115: 2105: 2073: 2064: 2063: 2053: 2021: 2015: 2014: 2004: 1994: 1970: 1964: 1963: 1946:(7): 1285–1294. 1934: 1928: 1927: 1909: 1903: 1888: 1882: 1881: 1863: 1857: 1856: 1846: 1814: 1808: 1807: 1797: 1757: 1746: 1745: 1709: 1698: 1697: 1687: 1647: 1641: 1640: 1612: 1606: 1605: 1595: 1589: 1588: 1570: 1564: 1563: 1561: 1543: 1534: 1443: 1429: 1415: 1401: 1387: 1373: 1359: 1345: 1333: 1199:Virginia opossum 992: 991: 987:Virginia opossum 781:Hormonal control 688:anlage (biology) 383:lactiferous duct 234:edit on Wikidata 231: 167: 114: 107: 40: 28: 27: 4723: 4722: 4718: 4717: 4716: 4714: 4713: 4712: 4688:Exocrine system 4673: 4672: 4671: 4666: 4645: 4566: 4562:Anatomy of the 4560: 4522:by W. L. Hurley 4516: 4503:Moore, Keith L. 4492: 4474: 4469: 4438: 4434: 4403: 4399: 4368: 4364: 4356: 4352: 4313: 4309: 4301: 4297: 4258: 4254: 4244: 4242: 4233: 4232: 4228: 4173: 4169: 4138: 4134: 4094: 4088: 4084: 4037: 4033: 4026: 4010: 4006: 3957: 3953: 3944: 3940: 3935: 3931: 3924: 3910: 3906: 3897: 3890: 3880: 3878: 3869: 3868: 3861: 3851: 3807: 3803: 3772:(Pt 1): 39–50. 3758: 3754: 3731: 3727: 3688: 3684: 3661: 3657: 3625: 3621: 3576: 3569: 3524: 3517: 3472: 3468: 3423: 3419: 3361: 3357: 3338:Cancer Research 3334: 3330: 3298: 3292: 3288: 3243: 3239: 3194: 3190: 3143: 3139: 3102:Differentiation 3094: 3090: 3064:10.1186/bcr1368 3043: 3039: 3000: 2996: 2951: 2947: 2921:10.1186/bcr1306 2900: 2896: 2857:(14): e100852. 2843: 2839: 2816: 2812: 2767: 2763: 2732:(12): 1725–30. 2718: 2714: 2677: 2670: 2663: 2647: 2643: 2630:(3B): 1617–24. 2616: 2607: 2600: 2584: 2580: 2527: 2523: 2516: 2497: 2493: 2456: 2452: 2426:10.1186/bcr1740 2401: 2397: 2360: 2356: 2317: 2313: 2274: 2261: 2238:(11): 5075–81. 2224: 2215: 2176: 2167: 2128: 2119: 2088:(12): a003178. 2074: 2067: 2022: 2018: 1992:10.1186/bcr1306 1971: 1967: 1935: 1931: 1924: 1910: 1906: 1901:Wayback Machine 1889: 1885: 1878: 1864: 1860: 1815: 1811: 1758: 1749: 1720:(6): 995–1003. 1710: 1701: 1648: 1644: 1613: 1609: 1596: 1592: 1585: 1571: 1567: 1541: 1535: 1526: 1522: 1512: 1463: 1456: 1449: 1444: 1435: 1433:Dromedary camel 1430: 1421: 1416: 1407: 1402: 1393: 1388: 1379: 1374: 1365: 1360: 1351: 1346: 1337: 1334: 1325: 1294: 1243:dayak fruit bat 1041: 1017: 1008: 999: 978: 973: 957: 922: 872: 811:beta-1 integrin 783: 778: 766: 726: 714:sexual maturity 699: 633: 628: 515: 482: 472: 436: 358: 352: 238: 165: 161: 145: 129: 112: 110: 105: 87: 86: 24: 17: 12: 11: 5: 4721: 4711: 4710: 4705: 4703:Female mammals 4700: 4698:Mammal anatomy 4695: 4690: 4685: 4683:Breast anatomy 4668: 4667: 4665: 4664: 4659: 4653: 4651: 4647: 4646: 4644: 4643: 4638: 4633: 4628: 4623: 4622: 4621: 4616: 4615: 4614: 4599: 4598: 4597: 4595:Tail of Spence 4587: 4586: 4585: 4574: 4572: 4568: 4567: 4559: 4558: 4551: 4544: 4536: 4530: 4529: 4523: 4515: 4514:External links 4512: 4511: 4510: 4505:et al. (2010) 4500: 4490: 4473: 4470: 4468: 4467: 4454:10.1086/284177 4432: 4397: 4378:(6): 606–616. 4362: 4350: 4307: 4295: 4268:(3): 253–266. 4252: 4226: 4167: 4132: 4082: 4051:(2): 121–134. 4031: 4024: 4004: 3975:(5): 1339–48. 3951: 3938: 3929: 3922: 3904: 3888: 3859: 3801: 3752: 3725: 3698:(2): 203–211. 3682: 3655: 3636:(4): 829–834. 3619: 3590:(3): 591–603. 3567: 3515: 3486:(6): 650–654. 3466: 3437:(8): 889–894. 3417: 3406:(10): 785–94. 3395:Also cited by 3355: 3328: 3309:(6): 793–802. 3286: 3257:(3): 615–624. 3237: 3188: 3164:10.1186/bcr634 3137: 3108:(7): 365–381. 3088: 3057:(1): 201–203. 3037: 2994: 2945: 2894: 2837: 2810: 2761: 2712: 2668: 2661: 2641: 2624:Anticancer Res 2605: 2598: 2578: 2541:(8): a003186. 2521: 2514: 2491: 2450: 2395: 2354: 2311: 2259: 2213: 2165: 2117: 2065: 2016: 1985:(5): 220–224. 1965: 1929: 1922: 1904: 1893:Apocrine Units 1883: 1876: 1858: 1809: 1747: 1699: 1642: 1623:(6): 698–704. 1607: 1590: 1583: 1565: 1523: 1521: 1518: 1511: 1510: 1505: 1500: 1495: 1490: 1485: 1480: 1475: 1470: 1464: 1455: 1452: 1451: 1450: 1445: 1438: 1436: 1431: 1424: 1422: 1417: 1410: 1408: 1403: 1396: 1394: 1389: 1382: 1380: 1375: 1368: 1366: 1361: 1354: 1352: 1347: 1340: 1338: 1335: 1328: 1324: 1321: 1293: 1290: 1247:Male lactation 1233: 1232: 1229: 1226: 1223: 1220: 1214: 1213: 1210: 1207: 1204: 1201: 1195: 1194: 1191: 1188: 1185: 1182: 1172: 1171: 1168: 1165: 1162: 1159: 1153: 1152: 1149: 1146: 1143: 1140: 1134: 1133: 1130: 1127: 1124: 1121: 1115: 1114: 1111: 1108: 1105: 1102: 1096: 1095: 1092: 1089: 1086: 1083: 1077: 1076: 1073: 1070: 1067: 1064: 1058: 1057: 1054: 1051: 1048: 1045: 1027: 1026: 1023: 1014: 1005: 996: 977: 974: 972: 969: 956: 953: 921: 918: 909:tight junction 895:giving birth, 880:adipose tissue 871: 868: 782: 779: 777: 774: 765: 764:Postmenopausal 762: 725: 722: 718:estrous cycles 716:and undergoes 698: 695: 632: 629: 627: 624: 598:lobuloalveolar 550:growth factors 522:growth hormone 514: 511: 471: 468: 435: 432: 368:cuboidal cells 351: 348: 318:male lactation 264:to feed young 260:that produces 250:exocrine gland 240: 239: 230: 224: 223: 218: 212: 211: 206: 200: 199: 194: 188: 187: 183: 182: 177: 171: 170: 156: 150: 149: 140: 134: 133: 124: 118: 117: 100: 94: 93: 89: 88: 85: 84: 79: 74: 69: 64: 59: 54: 49: 43: 41: 33: 32: 15: 9: 6: 4: 3: 2: 4720: 4709: 4706: 4704: 4701: 4699: 4696: 4694: 4691: 4689: 4686: 4684: 4681: 4680: 4678: 4663: 4662:Mammary ridge 4660: 4658: 4655: 4654: 4652: 4648: 4642: 4639: 4637: 4634: 4632: 4629: 4627: 4624: 4620: 4617: 4613: 4610: 4609: 4608: 4605: 4604: 4603: 4602:Mammary gland 4600: 4596: 4593: 4592: 4591: 4588: 4584: 4581: 4580: 4579: 4576: 4575: 4573: 4569: 4565: 4557: 4552: 4550: 4545: 4543: 4538: 4537: 4534: 4527: 4524: 4521: 4518: 4517: 4508: 4504: 4501: 4497: 4493: 4487: 4483: 4482: 4476: 4475: 4463: 4459: 4455: 4451: 4447: 4443: 4436: 4428: 4424: 4420: 4416: 4412: 4408: 4401: 4393: 4389: 4385: 4381: 4377: 4373: 4366: 4359: 4354: 4346: 4342: 4338: 4334: 4330: 4326: 4323:(3): 225–52. 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Index

Mammary Peak

Chest wall
Pectoralis muscles
Lobules
Nipple
Areola
Milk duct
Fatty tissue
Skin
Precursor
Mesoderm
Artery
Internal thoracic artery
Lateral thoracic artery
Vein
Internal thoracic vein
Axillary vein
Nerve
Supraclavicular nerves
Intercostal nerves
Lymph
Pectoral axillary lymph nodes
TA98
A16.0.02.006
TA2
7099
FMA
60088
Anatomical terminology

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