Adipose tissue

Adipose redirects here. For the ctional creature from

Doctor Who, see List of Doctor Who universe creatures
and aliens (09, AG) Adipose.
See also: Fat
In biology, adipose tissue i /dpos/, body fat, or

providing insulation from heat and cold. Around organs,

it provides protective padding. However, its main function is to be a reserve of lipids, which can be burned to
meet the energy needs of the body and to protect it from
excess glucose by storing triglycerides produced by the
liver from sugars, although some evidence suggests that
most lipid synthesis from carbohydrates occurs in the adipose tissue itself.[4] Adipose depots in dierent parts of
the body have dierent biochemical proles. Under normal conditions, it provides feedback for hunger and diet
to the brain.

Adipose tissue is one of the main types of connective tissue.

1.1 Mice

simply fat is loose connective tissue composed mostly

of adipocytes.[1] In addition to adipocytes, adipose tissue contains the stromal vascular fraction (SVF) of cells
including preadipocytes, broblasts, vascular endothelial
cells and a variety of immune cells (i.e., adipose tissue
macrophages [ATMs]). Adipose tissue is derived from
preadipocytes. Its main role is to store energy in the form
of lipids, although it also cushions and insulates the body.
Far from hormonally inert, adipose tissue has, in recent
years, been recognized as a major endocrine organ,[2] as it
produces hormones such as leptin, estrogen, resistin, and
the cytokine TNF. Moreover, adipose tissue can aect
other organ systems of the body and may lead to disease.
The two types of adipose tissue are white adipose tissue
(WAT), which stores energy, and brown adipose tissue
(BAT), which generates body heat. The formation of adipose tissue appears to be controlled in part by the adipose
gene. Adipose tissue more specically brown adipose
tissue was rst identied by the Swiss naturalist Conrad
Gessner in 1551.[3]

The obese mouse on the left has large stores of adipose tissue. For
comparison, a mouse with a normal amount of adipose tissue is
shown on the right.

Mice have eight major adipose depots, four of which are

within the abdominal cavity.[1] The paired gonadal depots
are attached to the uterus and ovaries in females and the
epididymis and testes in males; the paired retroperitoneal
depots are found along the dorsal wall of the abdomen,
surrounding the kidney, and, when massive, extend into
the pelvis. The mesenteric depot forms a glue-like web
that supports the intestines and the omental depot (which
originates near the stomach and spleen) and - when massive - extends into the ventral abdomen. Both the mesenteric and omental depots incorporate much lymphoid tissue as lymph nodes and milky spots, respectively. The
two supercial depots are the paired inguinal depots,
which are found anterior to the upper segment of the hind
limbs (underneath the skin) and the subscapular depots,
paired medial mixtures of brown adipose tissue adjacent
to regions of white adipose tissue, which are found under
the skin between the dorsal crests of the scapulae. The

Anatomical features

In humans, adipose tissue is located beneath the skin

(subcutaneous fat), around internal organs (visceral fat),
in bone marrow (yellow bone marrow), intermuscular
(Muscular system) and in the breast tissue. Adipose tissue is found in specic locations, which are referred to as
adipose depots. Apart from adipocytes, which comprise
the highest percentage of cells within adipose tissue, other
cell types are present, collectively termed stromal vascular fraction (SVF) of cells. SVF includes preadipocytes,
broblasts, adipose tissue macrophages, and endothelial
cells. Adipose tissue contains many small blood vessels.
In the integumentary system, which includes the skin, it
accumulates in the deepest level, the subcutaneous layer,
1

layer of brown adipose tissue in this depot is often covered by a frosting of white adipose tissue; sometimes
these two types of fat (brown and white) are hard to distinguish. The inguinal depots enclose the inguinal group
of lymph nodes. Minor depots include the pericardial,
which surrounds the heart, and the paired popliteal depots, between the major muscles behind the knees, each
containing one large lymph node.[5] Of all the depots in
the mouse, the gonadal depots are the largest and the most
easily dissected,[6] comprising about 30% of dissectible
fat.[7]

1.2

Obesity

ANATOMICAL FEATURES

total abdominal fat.[18][19] One study suggests at least 10

MET-hours per week of aerobic exercise is required for
visceral fat reduction.[20]

1.3 Epicardial fat

Epicardial adipose tissue (EAT) is a particular form of
visceral fat deposited around the heart and found to be a
metabolically active organ that generates various bioactive molecules, which might signicantly aect cardiac
function.[21] Marked component dierences have been
observed in comparing EAT with subcutaneous fat, suggesting a depot specic impact of stored fatty acids on
adipocyte function and metabolism.[22]

In an obese person, excess adipose tissue hanging downward from the abdomen is referred to as a panniculus 1.4 Subcutaneous fat
(or pannus). A panniculus complicates surgery of the
morbidly obese individual. It may remain as a literal See also: Body fat percentage
apron of skin if a severely obese person quickly loses Most of the remaining nonvisceral fat is found just belarge amounts of fat (a common result of gastric bypass
surgery). This condition cannot be eectively corrected
through diet and exercise alone, as the panniculus consists
of adipocytes and other supporting cell types shrunken
to their minimum volume and diameter. Reconstructive
surgery is one method of treatment.
See also: Abdominal obesity
Visceral fat or abdominal fat[8] (also known as organ fat
or intra-abdominal fat) is located inside the abdominal
cavity, packed between the organs (stomach, liver, intestines, kidneys, etc.). Visceral fat is dierent from
subcutaneous fat underneath the skin, and intramuscular
fat interspersed in skeletal muscles. Fat in the lower
body, as in thighs and buttocks, is subcutaneous and is
not consistently spaced tissue, whereas fat in the abdomen
is mostly visceral and semi-uid.[9] Visceral fat is composed of several adipose depots, including mesenteric,
epididymal white adipose tissue (EWAT), and perirenal
depots.

Micro-anatomy of subcutaneous fat.

low the skin in a region called the hypodermis.[23] This

subcutaneous fat is not related to many of the classic
obesity-related pathologies, such as heart disease, cancer,
and stroke, and some evidence even suggests it might be
protective.[24] The typically female (or gynecoid) pattern
of body fat distribution around the hips, thighs, and buttocks is subcutaneous fat, and therefore poses less of a
health risk compared to visceral fat.[25]

An excess of visceral fat is known as central obesity,

or belly fat, in which the abdomen protrudes excessively and new developments such as the Body Volume Index (BVI) are specically designed to measure
abdominal volume and abdominal fat. Excess visceral
fat is also linked to type 2 diabetes,[10] insulin resis- Like all other fat organs, subcutaneous fat is an active part
tance,[11] inammatory diseases,[12] and other obesity- of the endocrine system, secreting the hormones leptin
related diseases.[13]
and resistin.[23]
Men are more likely to have fat stored in the abdomen The relationship between the subcutaneous adipose layer
due to sex hormone dierences. Female sex hormone and total body fat in a person is often modelled by using
causes fat to be stored in the buttocks, thighs, and hips regression equations. The most popular of these equain women.[14][15] When women reach menopause and the tions was formed by Durnin and Wormersley, who rigorestrogen produced by the ovaries declines, fat migrates ously tested many types of skinfold, and, as a result, crefrom the buttocks, hips and thighs to the waist;[16] later ated two formulae to calculate the body density of both
fat is stored in the abdomen.[17]
men and women. These equations present an inverse corHigh-intensity exercise is one way to eectively reduce relation between skinfolds and body densityas the sum

2.1

Brown fat

of skinfolds increases, the body density decreases.[26]

3
This suggests a possible cause-and-eect link between the
two, wherein stress promotes the accumulation of visceral fat, which in turn causes hormonal and metabolic
changes that contribute to heart disease and other health
problems.[29]

Factors such as sex, age, population size or other variables may make the equations invalid and unusable, and,
as of 2012, Durnin and Wormersleys equations remain
only estimates of a persons true level of fatness. New
formulae are still being created.[26]
Recent advances in biotechnology have allowed for the
harvesting of adult stem cells from adipose tissue, allowing stimulation of tissue regrowth using a patients own
1.5 Ectopic fat
cells. In addition, adipose-derived stem cells from both
human and animals reportedly can be eciently reproEctopic fat is the storage of triglycerides in tissues other grammed into induced pluripotent stem cells without the
than adipose tissue, that are supposed to contain only need for feeder cells.[30] The use of a patients own cells
small amounts of fat, such as the liver, skeletal muscle, reduces the chance of tissue rejection and avoids ethiheart, and pancreas.[1] This can interfere with cellular cal issues associated with the use of human embryonic
functions and hence organ function and is associated with stem cells.[31] A growing body of evidence also suginsulin resistance in type-2 diabetes.[27] It is stored in rel- gests that dierent fat depots (i.e. abdominal, omenatively high amounts around the organs of the abdominal tal, pericardial) yield adipose-derived stem cells with
cavity, but is not to be confused as visceral fat.
dierent characteristics.[31][32] These depot-dependent
The specic cause for the accumulation of ectopic fat is features include proliferation rate, immunophenotype,
as well as senunknown. The cause is likely a combination of genetic, dierentiation potential, gene expression,
[33]
sitivity
to
hypoxic
culture
conditions.
environmental, and behavioral factors that are involved
in excess energy intake and decreased physical activity. Adipose tissue is the greatest peripheral source of
Substantial weight loss can reduce ectopic fat stores in all aromatase in both males and females, contributing to the
organs and this is associated with an improvement of the production of estradiol.
function of that organ.[27]
Adipose derived hormones include:

Physiology

Adiponectin
Resistin

Free fatty acids are liberated from lipoproteins by

lipoprotein lipase (LPL) and enter the adipocyte, where
they are reassembled into triglycerides by esterifying it
onto glycerol. Human fat tissue contains about 87%
lipids.

Plasminogen activator inhibitor-1 (PAI-1)

TNF
IL-6

Leptin
There is a constant ux of FFA (Free Fatty Acids) entering and leaving adipose tissue. The net direction of this
Estradiol (E2)
ux is controlled by insulin and leptinif insulin is elevated, then there is a net inward ux of FFA, and only
when insulin is low can FFA leave adipose tissue. Insulin Adipose tissues also secrete a type of cytokines (cellsecretion is stimulated by high blood sugar, which results to-cell signalling proteins) called adipokines (adipocytokines), which play a role in obesity-associated complifrom consuming carbohydrates.
cations. Perivascular adipose tissue releases adipokines
In humans, lipolysis (hydrolysis of triglycerides into free such as adiponectin that aect the contractile function of
fatty acids) is controlled through the balanced control the vessels that they surround.[1][34]
of lipolytic B-adrenergic receptors and a2A-adrenergic
receptor-mediated antilipolysis.
Fat cells have an important physiological role in maintaining triglyceride and free fatty acid levels, as well as determining insulin resistance. Abdominal fat has a dierent
metabolic prolebeing more prone to induce insulin resistance. This explains to a large degree why central obesity is a marker of impaired glucose tolerance and is an
independent risk factor for cardiovascular disease (even
in the absence of diabetes mellitus and hypertension).[28]
Studies of female monkeys at Wake Forest University
(2009) discovered that individuals suering from higher
stress have higher levels of visceral fat in their bodies.

2.1 Brown fat

Main article: Brown adipose tissue
A specialized form of adipose tissue in humans, most
rodents and small mammals, and some hibernating animals, is brown fat or brown adipose tissue. It is located
mainly around the neck and large blood vessels of the
thorax. This specialized tissue can generate heat by uncoupling the respiratory chain of oxidative phosphorylation within mitochondria. The process of uncoupling

3 BODY FAT METER

means that when protons transit down the electrochemical gradient across the inner mitochondrial membrane,
the energy from this process is released as heat rather
than being used to generate ATP. This thermogenic process may be vital in neonates exposed to cold, which then
require this thermogenesis to keep warm, as they are unable to shiver, or take other actions to keep themselves
warm.[35]

Gene defects in the leptin gene (ob) are rare in human obesity.[47] As of July, 2010, only 14 individuals
from ve families have been identied worldwide who
carry a mutated ob gene (one of which was the rst
ever identied cause of genetic obesity in humans)
two families of Pakistani origin living in the UK,
one family living in Turkey, one in Egypt, and one
in Austria[48][49][50][51][52] and two other families have
been found that carry a mutated ob receptor.[53][54] Others have been identied as genetically partially decient
in leptin, and, in these individuals, leptin levels on the low
end of the normal range can predict obesity.[55]

Attempts to simulate this process pharmacologically have

so far been unsuccessful. Techniques to manipulate the
dierentiation of brown fat could become a mechanism for weight loss therapy in the future, encouraging the
growth of tissue with this specialized metabolism without Several mutations of genes involving the melanocortins
inducing it in other organs.
(used in brain signaling associated with appetite) and their
a
Until recently, brown adipose tissue was thought to be receptors have also been identied as causing obesity in
[56]
larger
portion
of
the
population
than
leptin
mutations.
primarily limited to infants in humans, but new evidence
has now overturned that belief. Metabolically active tissue with temperature responses similar to brown adipose
was rst reported in the neck and trunk of some human
adults in 2007,[36] and the presence of brown adipose in
human adults was later veried histologically in the same
anatomical regions.[37][38][39]

In 2007, researchers isolated the adipose gene, which

those researchers hypothesize serves to keep animals lean
during times of plenty. In that study, increased adipose
gene activity was associated with slimmer animals.[57] Although its discoverers dubbed this gene the adipose gene,
it is not a gene responsible for creating adipose tissue.

2.2

Pre-adipocytes are undierentiated broblasts that can

be stimulated to form adipocytes. Recent studies shed
light into potential molecular mechanisms in the fate determination of pre-asipocytes although the exact lineage
of adipocyte is still unclear.[58][59]

Genetics

The thrifty gene hypothesis (also called the famine hypothesis) states that in some populations the body would
be more ecient at retaining fat in times of plenty,
thereby endowing greater resistance to starvation in times
of food scarcity. This hypothesis, originally advanced in
the context of glucose metabolism and insulin resistance,
has been discredited by physical anthropologists, physiologists, and the original proponent of the idea himself with
respect to that context, although according to its developer it remains as viable as when [it was] rst advanced
in other contexts.[40][41]

2.3 Physical properties

Adipose tissue has a density of ~0.9 g/ml.[60] Thus, a person with more adipose tissue will oat more easily than
a person of the same weight with more muscular tissue,
since muscular tissue has a density of 1.06 g/ml.[61]

3 Body fat meter

In 1995, Jerey Friedman, in his residency at the

Rockefeller University, together with Rudolph Leibel,
Douglas Coleman et al. discovered the protein leptin that
the genetically obese mouse lacked.[42][43][44] Leptin is
produced in the white adipose tissue and signals to the
hypothalamus. When leptin levels drop, the body interprets this as a loss of energy, and hunger increases. Mice
lacking this protein eat until they are four times their normal size.

A body fat meter is a widely available tool used to measure the percentage of fat in the human body. Dierent meters use various methods to determine the body
fat to weight ratio. They tend to under-read body fat
percentage.[62]

Leptin, however, plays a dierent role in diet-induced

obesity in rodents and humans. Because adipocytes produce leptin, leptin levels are elevated in the obese. However, hunger remains, and - when leptin levels drop due
to weight loss - hunger increases. The drop of leptin is
better viewed as a starvation signal than the rise of leptin
as a satiety signal.[45] However, elevated leptin in obesity
is known as leptin resistance. The changes that occur in
the hypothalamus to result in leptin resistance in obesity
are currently the focus of obesity research.[46]

In contrast with clinical tools, one relatively inexpensive

type of body fat meter uses the principle of bioelectrical
impedance analysis (BIA) in order to determine an individuals body fat percentage. To achieve this, the meter passes a small, harmless, electric current through the
body and measures the resistance, then uses information
on the persons weight, height, age, and sex to calculate an
approximate value for the persons body fat percentage.
The calculation measures the total volume of water in the
body (lean tissue and muscle contain a higher percent-

See also: Bioelectrical impedance analysis

5
age of water than fat), and estimates the percentage of fat
based on this information. The result can uctuate several percentage points depending on what has been eaten
and how much water has been drunk before the analysis.

Additional images
Diagrammatic sectional view of the skin (magnied).
White adipose tissue in paran section
Electronic instrument of body fat meter

See also
Adipose dierentiation-related protein
Adiposopathy
Apelin
Bioelectrical impedance analysis a method to measure body fat percentage.
Body Volume Index - a method to measure abdominal volume and abdominal fat.
Blubber an extra thick form of adipose tissue
found in some marine mammals.

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Further reading
MeSH A10.165.114
Stock, M. J.; Cinti, S. (2003). Adipose Tissue /
Structure and Function of Brown Adipose Tissue.
Encyclopedia of Food Sciences and Nutrition. p.
29. doi:10.1016/B0-12-227055-X/00008-0. ISBN
9780122270550.
Vernon, R. G.; Flint, D. J. (2003). Adipose Tissue
/ Structure and Function of White Adipose Tissue.
Encyclopedia of Food Sciences and Nutrition. p.
23. doi:10.1016/B0-12-227055-X/00007-9. ISBN
9780122270550.

External links
Adipose tissue photomicrographs

EXTERNAL LINKS

Text and image sources, contributors, and licenses

9.1

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9.3

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