WordNet

fry in a pan until it changes color; "brown the meat in the pan"
of a color similar to that of wood or earth (同)brownish, chocolate-brown, dark-brown
an orange of low brightness and saturation (同)brownness
(of skin) deeply suntanned (同)browned
a soft thin (usually translucent) paper (同)tissue_paper
part of an organism consisting of an aggregate of cells having a similar structure and function
abolitionist who was hanged after leading an unsuccessful raid at Harpers Ferry, Virginia (1800-1859) (同)John Brown
Scottish botanist who first observed the movement of small particles in fluids now known a Brownian motion (1773-1858) (同)Robert Brown
composed of animal fat; "adipose tissue constitutes the fat of meat"
the part of the face above the eyes (同)forehead

PrepTutorEJDIC

『褐色の』,『茶色の』 / 日に焼けた / 『褐色』,『茶色』,トビ色 / …'を'褐色にする / 褐色になる
〈U〉〈C〉(生物体の)『組織』 / 〈U〉〈C〉『薄織物』 / 〈U〉〈C〉水を吸収する柔らかな薄紙 / 〈C〉カーボンコピー用薄紙 / 〈C〉《a ~》(…を)織り交ぜて作ったもの《+of+名》 / =tissue paper
脂肪質の
《複数形》『まゆ』,『まゆ毛』 / 『額』(forehead) / がけ(山)の端,斜面の上端

Wikipedia preview

出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2017/03/15 14:23:39」(JST)

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Brown adipose tissue (BAT) or brown fat makes up the adipose organ together with white adipose tissue (or white fat).^[1] BAT is found in almost all mammals.

Classification of brown fat refers to two distinct cell populations with similar functions. The first shares a common embryological origin with muscle cells, found in larger "classic" deposits. The second develops from white adipocytes that are stimulated by the sympathetic nervous system. These adipocytes are found interspersed in white adipose tissue and are also named 'beige' or 'brite' .^[2]^[3]^[4]

BAT is especially abundant in newborns and in hibernating mammals.^[5] It is also present and metabolically active in adult humans,^[6]^[7] but its prevalence decreases as humans age.^[8] Its primary function is thermoregulation. In addition to heat produced by shivering muscle, BAT produces heat by non-shivering thermogenesis.

In contrast to white adipocytes, which contain a single lipid droplet, brown adipocytes contain numerous smaller droplets and a much higher number of (iron-containing) mitochondria, which gives BAT its brown appearance.^[2] Brown fat also contains more capillaries than white fat, to supply the tissue with oxygen and nutrients and distribute the produced heat throughout the body.

Anatomical location and histology

The presence of BAT in adult humans was discovered during FDG-PET scans to detect metastatic cancers.^[9]^[10] Using these scans and data from human autopsies, several BAT depots have been identified. In infants, BAT depots include, but are not limited to: interscapular, supraclavicular, suprarenal, pericardial, para-aortic and around the pancreas, kidney and trachea.^[11] These depots gradually get more white fat-like during adulthood. In adults, the depots that are most often detected in FDG-PET scans are the supraclavicular, paravertebral, mediastinal, para-aortic and suprarenal ones.^[12]^[13] It remains to be determined whether these depots are 'classical' BAT or beige/brite fat.^[14]^[15]

Brown fat in humans in the scientific and popular literature refers to two cell populations defined by both anatomical location and cellular morphology. Both share presence of small lipid droplets and numerous iron-rich mitochondria, giving the brown appearance.

"Classical" brown fat is found in highly vascularized deposits in somewhat consistent anatomical locations, such as between the shoulder blades, surrounding the kidneys, the neck, and supraclavicular area, and along the spinal cord. This is the smaller of the two types and has numerous small lipid droplets.
Beige fat is the adrenergically inducible cell type that is dispersed throughout adipose tissue. It has greater variability in lipid droplet size, and a greater proportion of lipid droplets to mitochondria giving it a lighter brown appearance.^[16]

Development

Brown fat cells come from the middle embryo layer, mesoderm, also the source of myocytes (muscle cells), adipocytes, and chondrocytes (cartilage cells).

The classic population of brown fat cells and muscle cells both seem to be derived from the same population of stem cells in the mesoderm, paraxial mesoderm. Both have the intrinsic capacity to activate the myogenic factor 5 (Myf5) promoter, a trait only associated with myocytes and this population of brown fat. Progenitors of traditional white fat cells and adrenergically induced brown fat do not have the capacity to activate the Myf5 promoter. Both adipocytes and brown adipocyte may be derived from pericytes, the cells which surround the blood vessels that run through white fat tissue.^[2]^[17] Notably, this is not the same as the presence of Myf5 protein, which is involved in the development of many tissues.

Additionally, muscle cells that were cultured with the transcription factor PRDM16 were converted into brown fat cells, and brown fat cells without PRDM16 were converted into muscle cells.^[2]

Function

The mitochondria in a eukaryotic cell utilize fuels to produce energy in the form of adenosine triphosphate (ATP). This process involves storing energy as a proton gradient, also known as the proton motive force (PMF), across the mitochondrial inner membrane. This energy is used to synthesize ATP when the protons flow across the membrane (down their concentration gradient) through the ATP synthase enzyme; this is known as chemiosmosis.

In endotherms, body heat is maintained by signaling the mitochondria to allow protons to run back along the gradient without producing ATP (proton leak). This can occur since an alternative return route for the protons exists through an uncoupling protein in the inner membrane. This protein, known as uncoupling protein 1 (thermogenin), facilitates the return of the protons after they have been actively pumped out of the mitochondria by the electron transport chain. This alternative route for protons uncouples oxidative phosphorylation and the energy in the PMF is instead released as heat.

To some degree, all cells of endotherms give off heat, especially when body temperature is below a regulatory threshold. However, brown adipose tissue is highly specialized for this non-shivering thermogenesis. First, each cell has a higher number of mitochondria compared to more typical cells. Second, these mitochondria have a higher-than-normal concentration of thermogenin in the inner membrane.

Infants

In neonates (newborn infants), brown fat makes up about 5% of the body mass and is located on the back, along the upper half of the spine and toward the shoulders. It is of great importance to avoid hypothermia, as lethal cold is a major death risk for premature neonates. Numerous factors make infants more susceptible to cold than adults:

The higher ratio of body surface area (proportional to heat loss) to body volume (proportional to heat production)
The higher proportional surface area of the head
The low amount of musculature and the inability or reluctance to shiver
A lack of thermal insulation, e.g., subcutaneous fat and fine body hair (especially in prematurely born children)
The inability to move away from cold areas, air currents or heat-draining materials
The inability to use additional ways of keeping warm (e.g., drying their skin, putting on clothing, moving into warmer areas, or performing physical exercise)
The nervous system is not fully developed and does not respond quickly and/or properly to cold (e.g., by contracting blood vessels in and just below the skin: vasoconstriction).

Heat production in brown fat provides an infant with an alternative means of heat regulation.

Adults

Micrograph of a hibernoma, a benign tumour thought to arise from brown fat (haematoxylin and eosin stain).

It was believed that after infants grow up, most of the mitochondria (which are responsible for the brown color) in brown adipose tissue disappear, and the tissue becomes similar in function and appearance to white fat. In rare cases, brown fat continues to grow, rather than involuting; this leads to a tumour known as a hibernoma. More recent research has shown that brown fat is related not to white fat, but to skeletal muscle.^[18]^[19]^[20]

Studies using positron emission tomography scanning of adult humans have shown that BAT is still present in most adults in the upper chest and neck (especially paravertebrally). The remaining deposits become more visible (increasing tracer uptake, that is, more metabolically active) with cold exposure, and less visible if an adrenergic beta blocker is given before the scan. These discoveries could lead to new methods of weight loss, since brown fat takes calories from normal fat and burns it. Scientists have been able to stimulate brown fat growth in mice.^[21]^[22]^[23]^[24] One study of APOE knock out mice showed cold exposure could promote atherosclerotic plaque growth and instability^[25] when study mice were subjected to sustained low temperatures of 4 °C for 8 weeks, which may cause a stress condition that shows rapid forced change rather than a safe acclimatisation that can be used to understand the potential in adult humans during modest reductions of ambient temperature by just 5 to 10 °C. Furthermore, several newer studies have documented the substantial benefits of cold exposure in multiple species including humans, for example researchers concluded that "activation of BAT is a powerful therapeutic avenue to ameliorate hyperlipidaemia and protect from atherosclerosis"^[26] and that brown fat activation reduces plasma triglyceride and cholesterol levels and attenuates diet-induced atherosclerosis development.^[27]

Long term studies of adult humans are needed to establish a balance of benefit and risk, in combination with historical research of living conditions of recent human generations prior to the current increase of poor health related to excessive accumulation of white fat. Pharmacological approaches using β3-adrenoceptor agonists have been shown to enhance glucose metabolic activity of brown adipose tissue in rodents.^[28]^[29]^[30]

Additionally research has shown:

BAT activation improves glucose homeostasis^[31] and insulin sensitivity in humans^[32] suggesting that anyone with impaired insulin function might benefit from BAT activation, however there is broader application given research showing even mildly elevated blood glucose in healthy non-diabetic humans is associated with damage over time of many organs such as eyes, tendons, endothelial/cardiovascular system and brain,^[33]^[34]^[35] and results in higher levels of damaging advanced glycation end products.
BAT activation may play an important role in bone health and bone density.^[36]^[37]
BAT activation though cold exposure increases adiponectin levels, just two hours of cold exposure resulted in a 70% increase in circulating adiponectin in adult men.^[38] Centenarians (both men and women) and their offspring have been found to have genetics that boost adiponectin, and they have higher circulating adiponectin, suggesting a link between longevity and adiponectin production^[39] In addition, high concentrations of plasma adiponectin in centenarians was associated with favorable metabolic indicators, and with lower levels of C-reactive protein and E-selectin.^[40]
Fibroblast Growth Factor 21 Production (FGF-21) has been documented as a pathway to longevity.^[41] BAT activation though cold exposure up-regulates circulating fibroblast growth factor 21 (FGF21) in humans by 37%.^[42] FGF21 improves insulin sensitivity and glucose metabolism^[43] which may partially explain its longevity promoting benefits.
Cold exposure increases circulating irisin.^[42] Irisin improves insulin sensitivity, increases bone quality and quantity, is involved in the building of lean muscle mass, and helps reduce obesity by converting white fat to brown fat,^[44] providing many of the same benefits of exercise.^[45] Healthy centenarians are characterized by increased serum irisin levels, whereas levels of this hormone were found to be significantly lower in young patients with myocardial infarction. These findings may prompt further research into the role played by irisin not only in vascular disorders but also in life span modulation.^[46]
Cold exposure increases SIRT1 phosphorylation/activity in both skeletal muscle and BAT, increasing thermogenesis and insulin sensitivity.^[47] Elevated SIRT1 levels in people are associated with increased human longevity.^[48] SIRT1 (and the other sirtuins) have many metabolic effects, but an important one for improving health and longevity is the fact that SIRT1 increases insulin sensitivity and glucose control in skeletal muscles,^[49] triggers the browning of white fat^[50] and increases BAT activity.^[51]

Other animals

The interscapular brown adipose tissue is commonly and inappropriately referred to as the hibernating gland.^[52] Whilst believed by many to be a type of gland, it is actually a collection of adipose tissues lying between the scapulae of rodentine mammals.^[53] Composed of brown adipose tissue and divided into two lobes, it resembles a primitive gland, regulating the output of a variety of hormones.^[54]^[55]^[56] The function of the tissue appears to be involved in the storage of medium to small lipid chains for consumption during hibernation. The smaller lipid structure allowing for a more rapid path of energy production than glycolysis.

In studies where the interscapular brown adipose tissue of rats were lesioned, it was demonstrated that the rats had difficulty regulating their normal body-weight.^[56]

The longest-lived small mammals: grey squirrels (24 yrs), bats (30 yrs), and naked mole rats (32 yrs), all have remarkably high levels of BAT and BAT activity.^[57]^[58]^[59]^[60]^[61]^[62] Furthermore, in animal species that span a wide range of latitudes, the within-species longevity is correlated with how far north (or south, in the southern hemisphere) an individual lives^[63] suggesting that cooler environments lead to increased BAT activation and increased lifespan across a wide range of species.

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External links

Histology image: 04901lob – Histology Learning System at Boston University – "Connective Tissue: multilocular (brown) adipocytes"

UpToDate Contents

全文を閲覧するには購読必要です。 To read the full text you will need to subscribe.

1. 肥満の病因 pathogenesis of obesity
2. 後期早産児 late preterm infants
3. 胸部PET thoracic positron emission tomography
4. 胎児（子宮内）発育遅延が認められる胎児 infants with fetal intrauterine growth restriction
5. まれな脳腫瘍 uncommon brain tumors

English Journal

Assessment of human brown adipose tissue density during daily ingestion of thermogenic capsinoids using near-infrared time-resolved spectroscopy.

Nirengi S1, Homma T2, Inoue N3, Sato H4, Yoneshiro T5, Matsushita M6, Kameya T7, Sugie H7, Tsuzaki K8, Saito M9, Sakane N8, Kurosawa Y10, Hamaoka T10.
Journal of biomedical optics.J Biomed Opt.2016 Sep 1;21(9):91305. doi: 10.1117/1.JBO.21.9.091305.
PMID 27135066

Lipid Profiling of In Vitro Cell Models of Adipogenic Differentiation: Relationships With Mouse Adipose Tissues.

Liaw L1,2, Prudovsky I1,2, Koza RA1,2, Anunciado-Koza RV1, Siviski ME1,2, Lindner V1,2, Friesel RE1,2, Rosen CJ1,2, Baker PR3, Simons B3, Vary CP1,2.
Journal of cellular biochemistry.J Cell Biochem.2016 Sep;117(9):2182-93. doi: 10.1002/jcb.25522. Epub 2016 Mar 16.
Our objective was to characterize lipid profiles in cell models of adipocyte differentiation in comparison to mouse adipose tissues in vivo. A novel lipid extraction strategy was combined with global lipid profiling using direct infusion and sequential precursor ion fragmentation, termed MS/MS(ALL)
PMID 26910604

Cell-Autonomous Brown-Like Adipogenesis of Preadipocytes From Retinoblastoma Haploinsufficient Mice.

Petrov PD1, Palou A1, Bonet ML1, Ribot J1.
Journal of cellular physiology.J Cell Physiol.2016 Sep;231(9):1941-52. doi: 10.1002/jcp.25299. Epub 2016 Jan 14.
Mechanisms behind the emergence of brown adipocyte-like (brite or beige) adipocytes within white adipose tissue (WAT) are of interest. Retinoblastoma protein gene (Rb) haploinsufficiency associates in mice with improved metabolic regulation linked to a greater capacity for fatty acid oxidation and t
PMID 26727985

Japanese Journal

香辛料などの食品成分によるヒト褐色脂肪組織の活性化と肥満予防

天使大学紀要 = Bulletin of Tenshi College 15(2), 31-40, 2015-08-31
NAID 120005657528

GLP-1の受容体活性化作用は視床下部AMPキナーゼを介して褐色脂肪組織の熱産生と脂肪組織の褐色化を刺激する

Diabetes : a journal of the American Diabetes Association 8(3), 26-37, 2015-06
NAID 40020531179

褐色脂肪組織と運動習慣との関係および褐色脂肪組織増加のための栄養介入

デサントスポーツ科学 36, 13-21, 2015-06
NAID 40020507440

Related Pictures

★リンクテーブル★

リンク元	「褐色脂肪組織」「brown fat」
関連記事	「adipose」「brown」

「褐色脂肪組織」

Brown adipose tissue
	Brown adipose tissue in a woman shown in a PET/CT exam
Details
Identifiers
Latin	textus adiposus fuscus
TH	H2.00.04.0.0004
FMA	20118
	Anatomical terminology [edit on Wikidata]

　　[★]

英: brown adipose tissue
関: 褐色脂肪

同: 825
関: BAT

「brown fat」

　　[★]

褐色脂肪

関: brown adipose tissue

「adipose」

　　[★]

adj.

脂肪の、脂肪性の、脂質の

関: aliphatic、fat、fatty、lipid、lipidic、lipo、lipogenous

「brown」

　　[★]

n.

褐色、茶色

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brown adipose tissue