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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.<ref name=BrJNutr>{{cite journal|author1=E. Connolly |author2=R. D. Morriseyt |author3=J. A. Carnie |journal=The British Journal of Nutrition|year=1982|volume=47|pages=653–658|title=The effect of interscapular brown adipose tissue removal on body-weight and cold response in the mouse|doi=10.1079/BJN19820077|pmid=6282304|issue=3}}</ref>
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.<ref name=BrJNutr>{{cite journal|author1=E. Connolly |author2=R. D. Morriseyt |author3=J. A. Carnie |journal=The British Journal of Nutrition|year=1982|volume=47|pages=653–658|title=The effect of interscapular brown adipose tissue removal on body-weight and cold response in the mouse|doi=10.1079/BJN19820077|pmid=6282304|issue=3}}</ref>

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<ref>{{Cite journal|last=Keil|first=Gerald|last2=Cummings|first2=Elizabeth|last3=de Magalhães|first3=João Pedro|date=2015-08-01|title=Being cool: how body temperature influences ageing and longevity|url=https://www.ncbi.nlm.nih.gov/pubmed/25832892|journal=Biogerontology|volume=16|issue=4|pages=383–397|doi=10.1007/s10522-015-9571-2|issn=1573-6768|pmc=4486781|pmid=25832892}}</ref><ref>{{Cite journal|last=Ducharme|first=Michel B.|last2=Larochelle|first2=Jacques|last3=Richard|first3=Denis|date=1989-01-01|title=Thermogenic Capacity in Gray and Black Morphs of the Gray Squirrel, Sciurus carolinensis|url=http://www.jstor.org/stable/30156213|journal=Physiological Zoology|volume=62|issue=6|pages=1273–1292}}</ref><ref>{{Cite journal|last=Woodley|first=Ryan|last2=Buffenstein|first2=Rochelle|date=2002-11-01|title=Thermogenic changes with chronic cold exposure in the naked mole-rat (Heterocephalus glaber)|url=https://www.ncbi.nlm.nih.gov/pubmed/12443938|journal=Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology|volume=133|issue=3|pages=827–834|issn=1095-6433|pmid=12443938}}</ref><ref>{{Cite journal|last=DALY|first=T. JOSEPH M.|last2=WILLIAMS|first2=LAURA A.|last3=BUFFENSTEIN|first3=ROCHELLE|date=1997-04-01|title=Catecholaminergic innervation of interscapular brown adipose tissue in the naked mole-rat (Heterocephalus glaber)|url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1467613/|journal=Journal of Anatomy|volume=190|issue=Pt 3|pages=321–326|doi=10.1046/j.1469-7580.1997.19030321.x|issn=0021-8782|pmc=1467613|pmid=9147219}}</ref><ref>{{Cite journal|last=Kim|first=Eun Bae|last2=Fang|first2=Xiaodong|last3=Fushan|first3=Alexey A.|last4=Huang|first4=Zhiyong|last5=Lobanov|first5=Alexei V.|last6=Han|first6=Lijuan|last7=Marino|first7=Stefano M.|last8=Sun|first8=Xiaoqing|last9=Turanov|first9=Anton A.|date=2011-10-12|title=Genome sequencing reveals insights into physiology and longevity of the naked mole rat|url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3319411/|journal=Nature|volume=479|issue=7372|pages=223–227|doi=10.1038/nature10533|issn=0028-0836|pmc=3319411|pmid=21993625}}</ref><ref>{{Cite journal|last=KUNZ|first=Thomas H.|last2=WRAZEN|first2=John A.|last3=BURNETT|first3=Christopher D.|date=1998-01-01|title=Changes in body mass and fat reserves in prehibernating little brown bats (Myotis lucifugus)|url=http://www.jstor.org/stable/42900766|journal=Écoscience|volume=5|issue=1|pages=8–17}}</ref>. 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<ref>{{Cite journal|last=Munch|first=Stephan B.|last2=Salinas|first2=Santiago|date=2009-08-18|title=Latitudinal variation in lifespan within species is explained by the metabolic theory of ecology|url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2728985/|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=106|issue=33|pages=13860–13864|doi=10.1073/pnas.0900300106|issn=0027-8424|pmc=2728985|pmid=19666552}}</ref> suggesting that cooler environments lead to increased BAT activation and increased lifespan across a wide range of species.


==See also==
==See also==

Revision as of 21:29, 19 August 2016

Brown adipose tissue
Brown adipose tissue in a woman shown in a PET/CT exam
Details
Identifiers
Latintextus adiposus fuscus
MeSHD002001
THH2.00.03.4.00004
FMA20118
Anatomical terminology

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, 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]

BAT activation has been shown to improve 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, brain, tendons, endothelial/cardiovascular system and brain[33][34][35], and results in higher levels of damaging advanced glycation end products.

Other animals

The interscapular brown adipose tissue is commonly and inappropriately referred to as the hibernating gland.[36] 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.[37] Composed of brown adipose tissue and divided into two lobes, it resembles a primitive gland, regulating the output of a variety of hormones.[38][39][40] 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.[40]

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[41][42][43][44][45][46]. 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[47] suggesting that cooler environments lead to increased BAT activation and increased lifespan across a wide range of species.

See also

References

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External links

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