ADIPOSE TISSUE
Adipose tissue is specialized connective tissue that
functions as the major storage site for fat in the form of triglycerides. Adipose tissue is found in mammals in two
different forms: white adipose tissue and brown adipose tissue. The presence, amount, and distribution of
each varies depending upon the species.
Most adipose tissue is white, the focus of this review.
White adipose tissue serves three functions: heat
insulation, mechanical cushion, and most importantly, a source of energy. Subcutaneous adipose tissue, found directly
below the skin, is an especially important heat insulator in the body, because
it conducts heat only one third as readily as other tissues. The degree of insulation is dependent upon
the thickness of this fat layer. For
example, a person with a 2-mm layer of subcutaneous fat will feel as
comfortable at 15°C as a person with a 1-mm layer at 16°C. Adipose tissue also surrounds internal organs
and provides some protection for these organs from jarring.
As the major form of energy storage, fat provides a buffer
for energy imbalances when energy intake is not equal to energy output. It is an efficient way to store excess
energy, because it is stored with very little water. Consequently, more energy can be derived per
gram of fat (9 kcal.gm-1) than per gram of carbohydrate
(4 kcal.gm-1) or protein (4 kcal.gm-1). In addition, if terrestrial animals stored
their excess energy as carbohydrate, the increased mass would interfere with
mobility. The average woman with 20%
body fat has about one month of energy stored as fat.
There are some constraints on the use of fat as fuel. Most animals cannot convert lipid into
carbohydrate. Tissues that function
predominantly anaerobically (e.g.. erythrocytes) must rely on carbohydrate for
energy and need to have an ample supply available. Additionally, under normal conditions the
brain is dependent upon glucose for energy and does not use fatty acids. In unusual metabolic circumstances, the brain
can use ketone bodies (a by-product of incomplete fat metabolism) when they are
present in sufficiently high quantities.
Finally, a typical diet contains a high proportion of carbohydrate, and
the transport of insoluble lipids through blood requires a specific mechanism,
so it may be "easier" metabolically for tissues to use glucose under
typical dietary conditions.
Brown adipose tissue, which derives its color from rich
vascularization and densely packed mitochondria, is found in various locations,
depending upon the species and/or age of the animal. In the rat, brown adipose tissue is found
primarily in the interscapular region and the axillae, minor amounts are found
near the thymus and in the dorsal midline region of the thorax and
abdomen. During maturation, in
non-hibernating animals, brown adipose tissue is metabolically less active,
although cold exposure can activate it.
In hibernating animals and neonates, brown adipose tissue is important
for regulating body temperature via non-shivering thermogenesis.
Instead of serving as a substrate, the lipid in brown
adipose tissue releases energy directly as heat and is, therefore used in heat
production for non-shivering thermogenesis and for utilization of excess
caloric intake via diet-induced-thermogenesis.
The mechanism of heat generation is related to the metabolism of the
mitochondria. Mitochondria from brown
adipose tissue have a specific carrier called uncoupling protein that transfers
protons from outside to inside without subsequent production of ATP.
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