Carbohydrate metabolism

Carbohydrate metabolism denotes the various biochemical processes responsible for the formation, breakdown and interconversion of carbohydrates in living organisms.

The most important carbohydrate is glucose, a simple sugar (monosaccharide) that is metabolized by nearly all known organisms. Glucose and other carbohydrates are part of a wide variety of metabolic pathways across species: plants synthesize carbohydrates from carbon dioxide and water by photosynthesis, storing the absorbed energy internally, often in the form of starch or lipids. Plant components are consumed by animals and fungi, and used as fuel for cellular respiration. Oxidation of one gram of carbohydrate yields approximately 4 kcal of energy, while the oxidation of one gram of lipids yields about 9 kcal. Energy obtained from metabolism (e.g., oxidation of glucose) is usually stored temporarily within cells in the form of ATP.^[1] Organisms capable of aerobic respiration metabolize glucose and oxygen to release energy with carbon dioxide and water as byproducts.

Carbohydrates can be chemically divided into two types: complex and simple. Simple carbohydrates consist of single or double sugar units (monosaccharides and disaccharides, respectively). Sucrose or table sugar (a disaccharide) is a common example of a simple carbohydrate. Complex carbohydrates contain three or more sugar units linked in a chain, with most containing hundreds to thousands of sugar units. They are digested by enzymes to release the simple sugars. Starch, for example, is a polymer of glucose units and is typically broken down to glucose. Cellulose is also a polymer of glucose but it cannot be digested by most organisms. Bacteria that produce enzymes to digest cellulose live inside the gut of some mammals, such as cows, and when these mammals eat plants, the cellulose is broken down by the bacteria and some of it is released into the gut.

Doctors and scientists once believed that eating complex carbohydrates instead of sugars would help maintain lower blood glucose. Numerous studies suggest, however, that both sugars and starch produce an unpredictable range of glycemic and insulinemic responses.^[2] While some studies support a more rapid absorption of sugars relative to starches^[3] other studies reveal that many carbohydrates such as those found in white bread, some types of white rice, and potatoes have glycemic indices similar to simple carbohydrates such as sucrose.^[4] Sucrose, for example, has a glycemic index (83) lower than expected because the sucrose molecule is half fructose, which has little effect on blood glucose.^[5] The value of classifying carbohydrates as simple or complex is questionable. The glycemic index is a better predictor of a carbohydrate's effect on blood glucose.^[6]

Carbohydrates are a superior short-term fuel for organisms because they are simpler to metabolize than fats or those amino acids (components of proteins) that can be used for fuel. In animals, the most important carbohydrate is glucose. The concentration of glucose in the blood is used as the main control for the central metabolic hormone, insulin. Starch, and cellulose in a few organisms (e.g., some animals (such as termites^[7]) and some microorganisms (such as protists and bacteria)), both being glucose polymers, are disassembled during digestion and absorbed as glucose. Some simple carbohydrates have their own enzymatic oxidation pathways, as do only a few of the more complex carbohydrates. The disaccharide lactose, for instance, requires the enzyme lactase to be broken into its monosaccharide components; many animals lack this enzyme in adulthood.

Carbohydrates are typically stored as long polymers of glucose molecules with glycosidic bonds for structural support (e.g. chitin, cellulose) or for energy storage (e.g. glycogen, starch). However, the strong affinity of most carbohydrates for water makes storage of large quantities of carbohydrates inefficient due to the large molecular weight of the solvated water-carbohydrate complex. In most organisms, excess carbohydrates are regularly catabolised to form acetyl-CoA, which is a feed stock for the fatty acid synthesis pathway; fatty acids, triglycerides, and other lipids are commonly used for long-term energy storage. The hydrophobic character of lipids makes them a much more compact form of energy storage than hydrophilic carbohydrates. However, animals, including humans, lack the necessary enzymatic machinery and so do not synthesize glucose from lipids (with a few exceptions, e.g. glycerol).^[8]

All carbohydrates share a general formula of approximately C_nH_2nO_n; glucose is C₆H₁₂O₆. Monosaccharides may be chemically bonded together to form disaccharides such as sucrose and longer polysaccharides such as starch and cellulose.

Catabolism

Oligosaccharides and/or polysaccharides are typically cleaved into smaller monosaccharides by enzymes called glycoside hydrolases. The monosaccharide units then enter monosaccharide catabolism. Organisms vary in the range of monosaccharides they can absorb and use and they can also vary in the range of more complex carbohydrates they are capable of disassembling.

Metabolic pathways

• Carbon fixation, or photosynthesis, in which CO₂ is reduced to carbohydrate.
• Glycolysis - the oxidation metabolism of glucose molecules to obtain ATP and pyruvate^[9]
  • Pyruvate from glycolysis enters the Krebs cycle, also known as the citric acid cycle, in aerobic organisms after moving through pyruvate dehydrogenase complex.^[10]
• The pentose phosphate pathway, which acts in the conversion of hexoses into pentoses and in NADPH regeneration.^[11] NADPH is an essential antioxidant in cells which prevents oxidative damage and acts as precursor for production of many biomolecules.
• Glycogenesis - the conversion of excess glucose into glycogen as a cellular storage mechanism; this prevents excessive osmotic pressure buildup inside the cell
• Glycogenolysis - the breakdown of glycogen into glucose, which provides a glucose supply for glucose-dependent tissues.
• Gluconeogenesis - de novo synthesis of glucose molecules from simple organic compounds. An example in humans is the conversion of a few amino acids in cellular protein to glucose.

Metabolic use of glucose is highly important as an energy source for muscle cells and in the brain, and red blood cells.

Energy production

Typically, a breakdown of one molecule of glucose by aerobic respiration (i.e. involving both glycolysis and Krebs cycle) is about 33-35 ATP.^[1] This is categorized as:

• Anaerobic breakdown by glycolysis - yielding 8-10 ATP
• Aerobic respiration by Krebs cycle - yielding 25 ATP

Glucoregulation

Glucoregulation is the maintenance of steady levels of glucose in the body; it is part of homeostasis, and so keeps a constant internal environment around cells in the body.

The hormone insulin is the primary regulatory signal in animals, suggesting that the basic mechanism is very old and very central to animal life. When present, it causes many tissue cells to take up glucose from the circulation, causes some cells to store glucose internally in the form of glycogen, causes some cells to take in and hold lipids, and in many cases controls cellular electrolyte balances and amino acid uptake as well. Its absence turns off glucose uptake into cells, reverses electrolyte adjustments, begins glycogen breakdown and glucose release into the circulation by some cells, begins lipid release from lipid storage cells, etc. The level of circulatory glucose (known informally as "blood sugar") is the most important signal to the insulin-producing cells. Because the level of circulatory glucose is largely determined by the intake of dietary carbohydrates, diet controls major aspects of metabolism via insulin. In humans, insulin is made by beta cells in the pancreas, fat is stored in adipose tissue cells, and glycogen is both stored and released as needed by liver cells. Regardless of insulin levels, no glucose is released to the blood from internal glycogen stores from muscle cells.

The hormone glucagon, on the other hand, has an effect opposite to that of insulin, forcing the conversion of glycogen in liver cells to glucose, which is then released into the blood. Muscle cells, however, lack the ability to export glucose into the blood. The release of glucagon is precipitated by low levels of blood glucose. Other hormones, notably growth hormone, cortisol, and certain catecholamines (such as epinepherine) have glucoregulatory actions similar to glucagon.

Human diseases

• Diabetes mellitus
• Lactose intolerance
• Fructose intolerance
• Galactosemia
• Glycogen storage disease

References

External links

• Carbohydrate metabolism at the US National Library of Medicine Medical Subject Headings (MeSH)
• BBC - GCSE Bitesize - Biology | Humans | Glucoregulation
• Sugar4Kids

Metabolism map

Carbon Fixation Photo- respiration Pentose Phosphate Pathway Citric Acid Cycle Glyoxylate Cycle Urea Cycle Fatty Acid Synthesis Fatty Acid Elongation Beta Oxidation Peroxisomal Beta Oxidation Glyco- genolysis Glyco- genesis Glyco- lysis Gluconeo- genesis Decarb- oxylation Fermentation Keto- lysis Keto- genesis feeders to Gluconeo- genesis Direct / C4 / CAM Carbon Intake Light Reaction Oxidative Phosphorylation Amino Acid Deamination Citrate Shuttle Lipogenesis Lipolysis Steroidogenesis MVA Pathway MEP Pathway Shikimate Pathway Transcription & Replication Translation Proteolysis Glycosy- lation Sugar Acids Double/Multiple Sugars & Glycans Simple Sugars Inositol-P Amino Sugars & Sialic Acids Nucleotide Sugars Hexose-P Triose-P Glycerol P-glycerates Pentose-P Tetrose-P Propionyl -CoA Succinate Acetyl -CoA Pentose-P P-glycerates Glyoxylate Photosystems Pyruvate Lactate Acetyl -CoA Citrate Oxalo- acetate Malate Succinyl -CoA α-Keto- glutarate Ketone Bodies Respiratory Chain Serine Group Alanine Branched-chain Amino Acids Aspartate Group Homoserine Group & Lysine Glutamate Group & Proline Arginine Creatine & Polyamines Ketogenic & Glucogenic Amino Acids Amino Acids Shikimate Aromatic Amino Acids & Histidine Ascorbate (Vitamin C) δ-ALA Bile Pigments Hemes Cobalamins (Vitamin B12) Various Vitamin B's Calciferols (Vitamin D) Retinoids (Vitamin A) Quinones (Vitamin K) & Carotenoids (Vitamin E) Cofactors Vitamins & Minerals Antioxidants PRPP Nucleotides Nucleic Acids Proteins Glycoproteins & Proteoglycans Chlorophylls MEP MVA Acetyl -CoA Polyketides Terpenoid Backbones Terpenoids & Carotenoids (Vitamin A) Cholesterol Bile Acids Glycero- phospholipids Glycerolipids Acyl-CoA Fatty Acids Glyco- sphingolipids Sphingolipids Waxes Polyunsaturated Fatty Acids Neurotransmitters & Thyroid Hormones Steroids Endo- cannabinoids Eicosanoids Major metabolic pathways in metro-style map. Click any text (name of pathway or metabolites) to link to the corresponding article. Single lines: pathways common to most lifeforms. Double lines: pathways not in humans (occurs in e.g. plants, fungi, prokaryotes). Orange nodes: carbohydrate metabolism. Violet nodes: photosynthesis. Red nodes: cellular respiration. Pink nodes: cell signaling. Blue nodes: amino acid metabolism. Grey nodes: vitamin and cofactor metabolism. Brown nodes: nucleotide and protein metabolism. Green nodes: lipid metabolism.