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Glycogen
Glycogen is a multibranched polysaccharide of glucose that
serves as a form of energy storage in animals[2] and fungi.
The polysaccharide structure represents the main storage
form of glucose in the body.
In humans, glycogen is made and stored primarily in the cells
of the liver and the muscles hydrated with three or four parts of
water.[3] Glycogen functions as the secondary long-term
energy storage, with the primary energy stores being fats held
in adipose tissue. Muscle glycogen is converted into glucose
by muscle cells, and liver glycogen converts to glucose for use
throughout the body including the central nervous system.
Glycogen is the analogue of starch, a glucose polymer that
functions as energy storage in plants. It has a similar structure
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Schematic two-dimensional cross-sectional

view of glycogen: A core protein of
glycogenin is surrounded by branches of
glucose units. The entire globular granule
may contain around 30,000 glucose units. [1]
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to amylopectin (a component of starch), but is more

extensively branched and compact than starch. Glycogen is
found in the form of granules in the cytosol/cytoplasm in many
cell types, and plays an important role in the glucose cycle.
Glycogen forms an energy reserve that can be quickly
mobilized to meet a sudden need for glucose, but one that is
less compact than the energy reserves of triglycerides
(lipids).
In the liver cells (hepatocytes), glycogen can compose up to
8% of its fresh weight (100120 g in an adult) soon after a
meal.[4] Only the glycogen stored in the liver can be made
accessible to other organs. In the muscles, glycogen is found
in a low concentration (1-2% of the muscle mass). The

A view of the atomic structure of a single

branched strand of glucose units in a
glycogen molecule.

amount of glycogen stored in the bodyespecially within the

muscles, liver, and red blood cells[5][6][7]mostly depends on
physical training, basal metabolic rate, and eating habits. Small
amounts of glycogen are found in the kidneys, and even smaller
amounts in certain glial cells in the brain and white blood cells. The
uterus also stores glycogen during pregnancy to nourish the
embryo.[8]

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Glycogen (black granules) in

spermatozoa of a flatworm;
transmission electron microscopy,
scale: 0.3 m
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Structure
Glycogen is a branched biopolymer consisting of linear chains of
glucose residues with further chains branching off every 8 to 12
glucoses or so. Glucoses are linked together linearly by (14)
glycosidic bonds from one glucose to the next. Branches are linked
to the chains from which they are branching off by (16)
glycosidic bonds between the first glucose of the new branch and a
glucose on the stem chain.[9]
Due to the way glycogen is synthesised, every glycogen granule
has at its core a glycogenin protein.[10]
Glycogen in muscle, liver, and fat cells is stored in a hydrated form,
composed of three or four parts of water per part of glycogen
associated with 0.45 millimoles of potassium per gram of

1,4--glycosidic linkages in the

glycogen oligomer

glycogen.[3]

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glycogen.

1,4--glycosidic and 1,6-glycosidic

linkages in the glycogen oligomer

Function
Liver
As a meal containing carbohydrates or protein is eaten and digested, blood glucose levels rise, and
the pancreas secretes insulin. Blood glucose from the portal vein enters liver cells (hepatocytes). Insulin
acts on the hepatocytes to stimulate the action of several enzymes, including glycogen synthase.
Glucose molecules are added to the chains of glycogen as long as both insulin and glucose remain
plentiful. In this postprandial or "fed" state, the liver takes in more glucose from the blood than it
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releases.
After a meal has been digested and glucose levels begin to fall, insulin secretion is reduced, and
glycogen synthesis stops. When it is needed for energy, glycogen is broken down and converted again
to glucose. Glycogen phosphorylase is the primary enzyme of glycogen breakdown. For the next 812
hours, glucose derived from liver glycogen is the primary source of blood glucose used by the rest of
the body for fuel.
Glucagon, another hormone produced by the pancreas, in many respects serves as a countersignal to
insulin. In response to insulin levels being above normal (when blood levels of glucose begin to fall
below the normal range), glucagon is secreted in increasing amounts and stimulates both
glycogenolysis (the breakdown of glycogen) and gluconeogenesis.

Muscle
Muscle cell glycogen appears to function as an immediate reserve source of available glucose for
muscle cells. Other cells that contain small amounts use it locally, as well. As muscle cells lack glucose6-phosphatase, which is required to pass glucose into the blood, the glycogen they store is available
solely for internal use and is not shared with other cells. This is in contrast to liver cells, which, on
demand, readily do break down their stored glycogen into glucose and send it through the blood stream
as fuel for other organs. Glycogen is also a suitable storage substance due to its insolubility in water,
which means it does not affect the osmotic pressure of a cell.
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History
Glycogen was discovered by Claude Bernard. His experiments showed that the liver contained a
substance that could give rise to reducing sugar by the action of a "ferment" in the liver. By 1857, he
described the isolation of a substance he called "la matire glycogne", or "sugar-forming substance".
Soon after the discovery of glycogen in the liver, A. Sanson found that muscular tissue also contains
glycogen. The empirical formula for glycogen of (C6H10O5)n was established by Kekule in 1858.[11]

Metabolism
Synthesis
Main article: Glycogenesis

Glycogen synthesis is, unlike its breakdown, endergonic - it requires the input of energy. Energy for
glycogen synthesis comes from uridine triphosphate (UTP), which reacts with glucose-1-phosphate,
forming UDP-glucose, in a reaction catalysed by UTPglucose-1-phosphate uridylyltransferase.
Glycogen is synthesized from monomers of UDP-glucose initially by the protein glycogenin, which has
two tyrosine anchors for the reducing end of glycogen, since glycogenin is a homodimer. After about
eight glucose molecules have been added to a tyrosine residue, the enzyme glycogen synthase
progressively lengthens the glycogen chain using UDP-glucose, adding (14)-bonded glucose. The
glycogen branching enzyme catalyzes the transfer of a terminal fragment of six or seven glucose
residues from a nonreducing end to the C-6 hydroxyl group of a glucose residue deeper into the interior
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of the glycogen molecule. The branching enzyme can act upon only a branch having at least 11
residues, and the enzyme may transfer to the same glucose chain or adjacent glucose chains.

Breakdown
Main article: Glycogenolysis

Glycogen is cleaved from the nonreducing ends of the

chain by the enzyme glycogen phosphorylase to
produce monomers of glucose-1-phosphate, which is
then converted to glucose 6-phosphate by

Action of glycogen phosphorylase on glycogen

phosphoglucomutase. A special debranching enzyme is

needed to remove the (1-6) branches in branched glycogen and reshape the chain into linear polymer.
The G6P monomers produced have three possible fates:
G6P can continue on the glycolysis pathway and be used as fuel.
G6P can enter the pentose phosphate pathway via the enzyme glucose-6-phosphate dehydrogenase
to produce NADPH and 5-carbon sugars.
In the liver and kidney, G6P can be dephosphorylated back to glucose by the enzyme glucose 6phosphatase. This is the final step in the gluconeogenesis pathway.

Clinical relevance
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Disorders of glycogen metabolism

The most common disease in which glycogen metabolism becomes abnormal is diabetes, in which,
because of abnormal amounts of insulin, liver glycogen can be abnormally accumulated or depleted.
Restoration of normal glucose metabolism usually normalizes glycogen metabolism, as well.
In hypoglycemia caused by excessive insulin, liver glycogen levels are high, but the high insulin levels
prevent the glycogenolysis necessary to maintain normal blood sugar levels. Glucagon is a common
treatment for this type of hypoglycemia.
Various inborn errors of metabolism are caused by deficiencies of enzymes necessary for glycogen
synthesis or breakdown. These are collectively referred to as glycogen storage diseases.

Glycogen depletion and endurance exercise

Long-distance athletes, such as marathon runners, cross-country skiers, and cyclists, often experience
glycogen depletion, where almost all of the athlete's glycogen stores are depleted after long periods of
exertion without enough energy consumption. This phenomenon is referred to as "hitting the wall".
Glycogen depletion can be forestalled in three possible ways. First, during exercise, carbohydrates with
the highest possible rate of conversion to blood glucose (high glycemic index) are ingested
continuously. The best possible outcome of this strategy replaces about 35% of glucose consumed at
heart rates above about 80% of maximum. Second, through endurance training adaptations and
specialized regimens (e.g. fasted low-intensity endurance training), the body can condition type I
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muscle fibers to improve both fuel use efficiency and workload capacity to increase the percentage of
fatty acids used as fuel,[12][13][citation needed] sparing carbohydrate use from all sources. Third, by
consuming large quantities of carbohydrates after depleting glycogen stores as a result of exercise or
diet, the body can increase storage capacity of intramuscular glycogen stores.[14][15][16] This process is
known as carbohydrate loading. In general, glycemic index of carbohydrate source does not matter
since muscular insulin sensitivity is increased as a result of temporary glycogen depletion.[17][18]
When experiencing glycogen debt, athletes often experience extreme fatigue to the point that it is
difficult to move. As a reference, the very best professional cyclists in the world will usually finish a 4- to
5-hr stage race right at the limit of glycogen depletion using the first three strategies.
When athletes ingest both carbohydrate and caffeine following exhaustive exercise, their glycogen is
replenished more rapidly.[19][20][unreliab le medical source?][21]

See also
Chitin
Peptidoglycan
Triglyceride

References
1. ^ William D. McArdle, Frank I. Katch, Victor L. Katch (2006). Exercise physiology: energy, nutrition,
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and human performance

(6 ed.). Lippincott Williams & Wilkins. p. 12. ISBN 978-0-7817-4990-9.

2. ^ Sadava et al. (2011). Life (9th, International ed.). W. H. Freeman. ISBN 9781429254311.
3. ^ a b Kreitzman SN, Coxon AY, Szaz KF (1992). "Glycogen storage: illusions of easy weight loss,
excessive weight regain, and distortions in estimates of body composition"

(PDF). The American

Journal of Clinical Nutrition 56 (1 Suppl): 292s293s. PMID 1615908 .

4. ^ Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life . Boston,
Massachusetts: Pearson Prentice Hall. ISBN 0-13-250882-6.
5. ^ Moses SW, Bashan N, Gutman A (December 1972). "Glycogen metabolism in the normal red
blood cell" . Blood 40 (6): 83643. PMID 5083874 .
6. ^ Ingermann RL, Virgin GL (1987). "Glycogen content and release of glucose from red blood cells of
the sipunculan worm themiste dyscrita"

(PDF). J Exp Biol 129: 1419.

7. ^ Miwa I, Suzuki S (November 2002). "An improved quantitative assay of glycogen in erythrocytes".
Annals of Clinical Biochemistry 39 (Pt 6): 6123. doi:10.1258/000456302760413432 .
PMID 12564847 .
8. ^ Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life . Boston:
Pearson Prentice Hall. ISBN 0-13-250882-6.
9. ^ Berg, Tymoczko & Stryer (2012). Biochemistry (7th, International ed.). W. H. Freeman. p. 338.
ISBN 1429203145.
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10. ^ Berg et al. (2012). Biochemistry (7th, International ed.). W. H. Freeman. p. 650.
11. ^ F. G. Young (1957). "Claude Bernard and the Discovery of Glycogen". British Medical Journal 1
(5033 (Jun. 22, 1957)): 14317. doi:10.1136/bmj.1.5033.1431 . JSTOR 25382898 .
12. ^ http://www.bodyrecomposition.com/training/methods-of-endurance-training-part-1.html
13. ^ http://www.bodyrecomposition.com/fat-loss/qa-steady-state-vs-tempo-training-and-fat-loss.html
14. ^ http://www.simplyshredded.com/research-review-an-in-depth-look-into-carbing-up-on-the-cyclicalketogenic-diet-with-lyle-mcdonald.html
15. ^ McDonald, Lyle. The Ketogenic Diet: A Complete Guide for the Dieter and the Practitioner. Lyle
McDonald, 1998
16. ^ "Costill DL et. al. Muscle glycogen utilization during prolonged exercise on successive days. J
Appl Physiol (1971) 31: 834-838."
17. ^ Glycogen depletion and increased insulin sensitivity and responsiveness in muscle after
exerciseAm J Physiol Endocrinol MetabDecember 1, 1986 251:(6) E664-E669
18. ^ McDonald, Lyle. The Ultimate Diet 2.0. Lyle McDonald, 2003
19. ^ Pedersen DJ, Lessard SJ, Coffey VG et al. (July 2008). "High rates of muscle glycogen
resynthesis after exhaustive exercise when carbohydrate is coingested with caffeine". Journal of
Applied Physiology (Original article) 105 (1): 713. doi:10.1152/japplphysiol.01121.2007 .
PMID 18467543 .
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20. ^ "Post-exercise Caffeine Helps Muscles Refuel"

(Press release). American Physiological

Society. Newswise. Retrieved July 6, 2008.

21. ^ Gaudet, Laura; Jackson, Allen; Streitz, Carmyn; McIntire, Kyle; McDaniel, Larry. "The Effects Of
Caffeine On Athletic Performance" .
http://journals.cluteonline.com/index.php/CTMS/article/view/5518 . Clute Institute. Retrieved 17 June
2014.

External links
Glycogen detection using Periodic Acid Schiff Staining
Glycogen storage disease - McArdle's Disease Website
Glycogen

at the US National Library of Medicine Medical Subject Headings (MeSH)

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