Biochemistry, or biological chemistry (distinct from chemical biology), is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology, and metabolism. Over the last decades of the 20th century, biochemistry has become successful at explaining living processes through these three disciplines. Almost all areas of the life sciences are being uncovered and developed through biochemical methodology and research.Voet (2005), p. 3.

Biochemistry focuses on understanding the chemical basis that allows biological molecules to give rise to the processes that occur within living cells and between cells,Karp (2009), p. 2. in turn relating greatly to the understanding of tissues and organs as well as organism structure and function.Miller (2012). p. 62. Biochemistry is closely related to molecular biology, the study of the molecular mechanisms of biological phenomena.Astbury (1961), p. 1124.

Much of biochemistry deals with the structures, functions, and interactions of biological macromolecules such as proteins, nucleic acids, carbohydrates, and lipids. They provide the structure of cells and perform many of the functions associated with life.Eldra (2007), p. 45. The chemistry of the cell also depends upon the reactions of small molecules and ions. These can be inorganic (for example, water and metal ions) or organic (for example, the amino acids, which are used to synthesize proteins).Marks (2012), Chapter 14. The mechanisms used by cells to harness energy from their environment via chemical reactions are known as metabolism.

The findings of biochemistry are applied primarily in medicine, nutrition, and agriculture. In medicine, biochemists investigate the causes and cures of diseases.Finkel (2009), pp. 1–4. Nutrition studies how to maintain health and wellness and also the effects of nutritional deficiencies.UNICEF (2010), pp. 61, 75. In agriculture, biochemists investigate soil and fertilizers with the goal of improving crop cultivation, crop storage, and pest control. In recent decades, biochemical principles and methods have been combined with problem-solving approaches from engineering to manipulate living systems in order to produce useful tools for research, industrial processes, and diagnosis and control of diseasethe discipline of biotechnology.

History

At its most comprehensive definition, biochemistry can be seen as a study of the components and composition of living things and how they come together to become life. In this sense, the history of biochemistry may therefore go back as far as the ancient Greeks.Helvoort (2000), p. 81. However, biochemistry as a specific scientific discipline began sometime in the 19th century, or a little earlier, depending on which aspect of biochemistry is being focused on. Some argued that the beginning of biochemistry may have been the discovery of the first enzyme, diastase (now called amylase), in 1833 by Anselme Payen,Hunter (2000), p. 75. while others considered Eduard Buchner's first demonstration of a complex biochemical process alcoholic fermentation in cell-free extracts in 1897 to be the birth of biochemistry.Hamblin (2005), p. 26.Hunter (2000), pp. 96–98. Some might also point as its beginning to the influential 1842 work by Justus von Liebig, Animal chemistry, or, Organic chemistry in its applications to physiology and pathology, which presented a chemical theory of metabolism, or even earlier to the 18th century studies on fermentation and respiration by Antoine Lavoisier.Berg (1980), pp. 1–2.Holmes (1987), p. xv. Many other pioneers in the field who helped to uncover the layers of complexity of biochemistry have been proclaimed founders of modern biochemistry. Emil Fischer, who studied the chemistry of proteins,Feldman (2001), p. 206. and F. Gowland Hopkins, who studied enzymes and the dynamic nature of biochemistry, represent two examples of early biochemists.Rayner-Canham (2005), p. 136.

The term "biochemistry" was first used when Vinzenz Kletzinsky (1826–1882) had his "Compendium der Biochemie" printed in Vienna in 1858; it derived from a combination of biology and chemistry. In 1877, Felix Hoppe-Seyler used the term ( in German) as a synonym for physiological chemistry in the foreword to the first issue of Zeitschrift für Physiologische Chemie (Journal of Physiological Chemistry) where he argued for the setting up of institutes dedicated to this field of study.Ziesak (1999), p. 169.Kleinkauf (1988), p. 116. The German chemist Carl Neuberg however is often cited to have coined the word in 1903,Ben-Menahem (2009), p. 2982.Amsler (1986), p. 55.Horton (2013), p. 36. while some credited it to Franz Hofmeister.Kleinkauf (1988), p. 43.

It was once generally believed that life and its materials had some essential property or substance (often referred to as the "vital principle") distinct from any found in non-living matter, and it was thought that only living beings could produce the molecules of life.Fiske (1890), pp. 419–20. In 1828, Friedrich Wöhler published a paper on his serendipitous urea synthesis from potassium cyanate and ammonium sulfate; some regarded that as a direct overthrow of vitalism and the establishment of organic chemistry.Kauffman (2001), pp. 121–133. However, the Wöhler synthesis has sparked controversy as some reject the death of vitalism at his hands. Since then, biochemistry has advanced, especially since the mid-20th century, with the development of new techniques such as chromatography, X-ray diffraction, dual polarisation interferometry, NMR spectroscopy, radioisotopic labeling, electron microscopy and molecular dynamics simulations. These techniques allowed for the discovery and detailed analysis of many molecules and metabolic pathways of the cell, such as glycolysis and the Krebs cycle (citric acid cycle), and led to an understanding of biochemistry on a molecular level.

Another significant historic event in biochemistry is the discovery of the gene, and its role in the transfer of information in the cell. In the 1950s, James D. Watson, Francis Crick, Rosalind Franklin and Maurice Wilkins were instrumental in solving DNA structure and suggesting its relationship with the genetic transfer of information.Tropp (2012), pp. 19–20. In 1958, George Beadle and Edward Tatum received the Nobel Prize for work in fungi showing that one gene produces one enzyme.Krebs (2012), p. 32. In 1988, Colin Pitchfork was the first person convicted of murder with DNA evidence, which led to the growth of forensic science.Butler (2009), p. 5. More recently, Andrew Z. Fire and Craig C. Mello received the 2006 Nobel Prize for discovering the role of RNA interference (RNAi) in the silencing of gene expression.Chandan (2007), pp. 193–194.

The chemical elements of life

Around two dozen chemical elements are essential to various kinds of biological life. Most rare elements on Earth are not needed by life (exceptions being selenium and iodine), while a few common ones (aluminium and titanium) are not used. Most organisms share element needs, but there are a few differences between plants and animals. For example, ocean algae use bromine, and land animals incorporate ionic bromide in their connective tissue, but it is non-essential for land plants and can even be problematic at high quantities. All animals require sodium, but it is only an essential element for a small subset of plants, although many others do grow better in its presence . Plants need boron and silicon, but animals may not (or may need ultra-small amounts).

Biomolecules are constructed from just six bulk macronutrients, abbreviated CHNOPS (carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur), and these six make up >99% of the mass of all of our cells. Humans further require significant quantities of four key ions (magnesium, potassium, sodium, and calcium) and smaller amounts of some other elements and ions, termed micronutrients. In addition to the six major elements that compose most of the human body, humans require smaller amounts of possibly 18 more .Nielsen (1999), pp. 283–303.

Biomolecules

The four main classes of molecules in biochemistry (often called biomolecules) are carbohydrates, lipids, proteins, and nucleic acids.Slabaugh (2007), pp. 3–6. Many biological molecules are polymers: chains of smaller repeating units called monomers. When monomers are linked together to synthesize a biological polymer, they undergo a process called dehydration synthesis. Macromolecules can assemble in larger complexes with other molecules of the same or different type, often required for biological activity.

Carbohydrates

Two of the main functions of carbohydrates are energy storage and providing structure. The term 'sugar' typically refers to mono- and disaccharides (defined below), which fall into the former category and taste sweet. There are more carbohydrates on Earth than any other known type of biomolecule; they are used to store energy and genetic information, as well as play important roles in cell to cell interactions and communications. Carbohydrates generally have ratios of carbon:hydrogen:oxygen of 1:2:1, meaning a 1:1 ratio of carbon atoms to water (H2O), from which the term "carbohydrate" derives.

The simplest type of carbohydrate is a monosaccharide, which have the chemical formula C6H12O6 but vary in their precise structures. Glucose, fructose, and galactose are hexoses, i.e. 6-carbon sugar, sharing the formula C6H12O6, but varying in precise structure. A well-known pentose, or 5-carbon sugar, is deoxyribose (C5H10O4), a component of DNA. A monosaccharide can switch between acyclic (open-chain) form and cyclic form. This occurs through reaction of one of the hydroxyl groups with the carbonyl group on the end of the linear form, bridging them with an oxygen atom and releasing a water molecule in the process. The cyclic molecule has a hemiacetal or hemiketal group, depending on whether the linear form was an aldose or a ketose, which describe the position of the carbonyl group to begin with.Voet (2005), pp. 358–359.

In these cyclic forms, the ring usually has 5 or 6 atoms. These forms are called furanoses and pyranoses, respectively—by analogy with furan and pyran, the simplest compounds with the same carbon-oxygen ring (although they lack the carbon-carbon double bonds of these two molecules). For example, the aldohexose glucose may form a hemiacetal linkage between the hydroxyl on carbon 1 and the oxygen on carbon 4, yielding a molecule with a 5-membered ring, called glucofuranose. The same reaction can take place between carbons 1 and 5 to form a molecule with a 6-membered ring, called glucopyranose. Notice that this means that pentoses/furanoses, and hexoses/pyranoses, are not interchangeable pairs: the number of carbons in the molecule can be different from the number of atoms in the ring, as not all carbons are necessarily included in the ring backbone.

Two monosaccharides can be joined by a glycosidic or ester bond into a disaccharide through a dehydration reaction during which a molecule of water is released. The reverse reaction in which the glycosidic bond of a disaccharide is broken into two monosaccharides is termed hydrolysis. The best-known disaccharide is sucrose or ordinary sugar, which consists of a glucose molecule and a fructose molecule joined. Another important disaccharide is lactose, found in milk, consisting of a glucose molecule and a galactose molecule. Lactose may be hydrolysed by lactase, and deficiency in this enzyme results in lactose intolerance.