Biochemistry

For the journal, see Biochemistry (journal).

1 History
Biological Chemistry redirects here. For the journal
formerly named Biological Chemistry Hoppe-Seyler, see Main article: History of biochemistry
Biological Chemistry (journal).
It once was generally believed that life and its materiBiochemistry, sometimes called biological chemistry,
is the study of chemical processes within and relating
to living organisms.[1] By controlling information ow
through biochemical signaling and the ow of chemical
energy through metabolism, biochemical processes give
rise to the complexity of life. Over the last 40 years,
biochemistry has become so successful at explaining living processes that now almost all areas of the life sciences from botany to medicine are engaged in biochemical research.[2] Today, the main focus of pure biochemistry is in understanding how biological molecules give
rise to the processes that occur within living cells, which
in turn relates greatly to the study and understanding of
whole organisms.
Biochemistry is closely related to molecular biology, the
study of the molecular mechanisms by which genetic information encoded in DNA is able to result in the processes of life. Depending on the exact denition of the
terms used, molecular biology can be thought of as a
branch of biochemistry, or biochemistry as a tool with
which to investigate and study molecular biology.
Much of biochemistry deals with the structures, functions
and interactions of biological macromolecules, such as
proteins, nucleic acids, carbohydrates and lipids, which
provide the structure of cells and perform many of the
functions associated with life. The chemistry of the cell
also depends on the reactions of smaller 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. The mechanisms by
which cells harness energy from their environment via
chemical reactions are known as metabolism. The ndings of biochemistry are applied primarily in medicine,
nutrition, and agriculture. In medicine, biochemists investigate the causes and cures of disease. In nutrition,
they study how to maintain health and study the eects
of nutritional deciencies. In agriculture, biochemists investigate soil and fertilizers, and try to discover ways to
improve crop cultivation, crop storage and pest control.

Gerty Cori and Carl Cori jointly won the Nobel Prize in 1947 for
their discovery of the Cori cycle at RPMI.

als 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.[3] Then, in
1828, Friedrich Whler published a paper on the synthesis of urea, proving that organic compounds can be created articially.[4]
The beginning of biochemistry may have been the discovery of the rst enzyme, diastase (today called amylase),
in 1833 by Anselme Payen.[5] Eduard Buchner contributed the rst demonstration of a complex biochemical process outside a cell in 1896: alcoholic fermentation in cell extracts of yeast.[6] Although the term biochemistry seems to have been rst used in 1882, it
is generally accepted that the formal coinage of biochemistry occurred in 1903 by Carl Neuberg, a German
chemist.[7] Since then, biochemistry has advanced, especially since the mid-20th century, with the development of new techniques such as chromatography, X-ray
1

3 BIOMOLECULES

diraction, 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).

tively small micromolecules that are linked together to

create large macromolecules known as polymers. When
monomers are linked together to synthesize a biological
polymer, they undergo a process called dehydration synthesis. Dierent macromolecules can assemble in larger
complexes, often needed for biological activity.

Another signicant historic event in biochemistry is the

discovery of the gene and its role in the transfer of in3.1 Carbohydrates
formation in the cell. This part of biochemistry is of[8]
ten called molecular biology. In the 1950s, James D.
Carbohydrate, Monosaccharide,
Watson, Francis Crick, Rosalind Franklin, and Maurice Main articles:
Disaccharide
and
Polysaccharide
Wilkins were instrumental in solving DNA structure
Carbohydrates are made from monomers called
and suggesting its relationship with genetic transfer of
[9]
information. In 1958, George Beadle and Edward
Tatum received the Nobel Prize for work in fungi showing that one gene produces one enzyme.[10] In 1988,
Colin Pitchfork was the rst person convicted of murder with DNA evidence, which led to growth of forensic
science.[11] 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.[12]
A molecule of sucrose (glucose + fructose), a disaccharide.

Starting materials: the chemical

elements of life

monosaccharides.
Some of these monosaccharides
include glucose (C6 H12 O6 ), fructose (C6 H12 O6 ), and
deoxyribose (C5 H10 O4 ). When two monosaccharides
undergo dehydration synthesis, water is produced, as two
Main articles: Composition of the human body and
hydrogen atoms and one oxygen atom are lost from the
Dietary mineral
two monosaccharides hydroxyl group.
Around two dozen of the 92 naturally occurring chemical
elements are essential to various kinds of biological life. 3.2 Lipids
Most rare elements on Earth are not needed by life (exceptions being selenium and iodine), while a few com- Main articles: Lipid, Glycerol and Fatty acid
mon ones (aluminum and titanium) are not used. Most Lipids are usually made from one molecule of glycerol
organisms share element needs, but there are a few differences between plants and animals. For example ocean
algae use bromine but land plants and animals seem to
need none. All animals require sodium, but some plants
do not. Plants need boron and silicon, but animals may
not (or may need ultra-small amounts).
Just six elementscarbon, hydrogen, nitrogen, oxygen,
calcium, and phosphorusmake up almost 99% of the
mass of a human body (see composition of the human
body for a complete list). In addition to the six major
A triglyceride with a glycerol molecule on the left and three fatty
elements that compose most of the human body, humans
acids coming o it.
[13]
require smaller amounts of possibly 18 more.
combined with other molecules. In triglycerides, the main
group of bulk lipids, there is one molecule of glycerol and
three fatty acids. Fatty acids are considered the monomer
3 Biomolecules
in that case, and may be saturated (no double bonds in the
The four main classes of molecules in biochemistry carbon chain) or unsaturated (one or more double bonds
(often called biomolecules) are carbohydrates, lipids, in the carbon chain).
proteins, and nucleic acids. Many biological molecules Lipids, especially phospholipids, are also used in various
are polymers: in this terminology, monomers are rela- pharmaceutical products, either as co-solubilisers (e.g., in

3
parenteral infusions) or else as drug carrier components
(e.g., in a liposome or transfersome).
5 end
O

3.3

Proteins

Thymine

Adenine
O

NH 2

3 end

N
O

Main articles: Proteins and Amino Acids

Proteins are very large molecules macro-biopolymers

OH

HN

O
O

O
O
O
O

NH 2

P
O
N

HN

N
N

O H2N

PhosphateO
deoxyribose O P

backbone

O
O

O H2N

NH

N
N

O
O
O
O

NH
O

NH 2

H2N

N
O

O
O

O
OH

Cytosine
3 end
Guanine
5 end
O

The general structure of an -amino acid, with the amino group

on the left and the carboxyl group on the right.

The structure of deoxyribonucleic acid (DNA), the picture shows

the monomers being put together.

a ribose sugar, and a nitrogenous base. The phosphate

group and the sugar of each nucleotide bond with each
other to form the backbone of the nucleic acid, while
the sequence of nitrogenous bases stores the information. The most common nitrogenous bases are adenine,
cytosine, guanine, thymine, and uracil. The nitrogenous
bases of each strand of a nucleic acid will form hydrogen
bonds with certain other nitrogenous bases in a complementary strand of nucleic acid (similar to a zipper). Adenine binds with thymine and uracil; Thymine binds only
In order to determine whether two proteins are related, or with adenine; and cytosine and guanine can bind only with
in other words to decide whether they are homologous or one another.
not, scientists use sequence-comparison methods. Methods like Sequence Alignments and Structural Alignments
are powerful tools that help scientists identify homologies 4 Carbohydrates
between related molecules.[14]
made from monomers called amino acids. There are 20
standard amino acids, each containing a carboxyl group,
an amino group, and a side-chain (known as an R
group). The R group is what makes each amino acid
dierent, and the properties of the side-chains greatly inuence the overall three-dimensional conformation of a
protein. When amino acids combine, they form a special
bond called a peptide bond through dehydration synthesis, and become a polypeptide, or protein.

The relevance of nding homologies among proteins goes Main article: Carbohydrate
beyond forming an evolutionary pattern of protein families. By nding how similar two protein sequences are,
we acquire knowledge about their structure and therefore The function of carbohydrates includes energy storage
and providing structure. Sugars are carbohydrates, but
their function.
not all carbohydrates are sugars. There are more carbohydrates on Earth than any other known type of
biomolecule; they are used to store energy and genetic
3.4 Nucleic acids
information, as well as play important roles in cell to cell
Main articles: Nucleic acid, DNA, RNA and Nucleotides interactions and communications.
Nucleic acids are the molecules that make up DNA, an
extremely important substance that all cellular organisms use to store their genetic information. The most
common nucleic acids are deoxyribonucleic acid (DNA)
and ribonucleic acid (RNA). Their monomers are called
nucleotides. A nucleotide consists of a phosphate group,

4.1 Monosaccharides
The simplest type of carbohydrate is a monosaccharide,
which between other properties contains carbon, hydrogen, and oxygen, mostly in a ratio of 1:2:1 (generalized
formula CnHnOn, where n is at least 3). Glucose, one

4 CARBOHYDRATES
tic contexts, called table sugar or cane sugar to dierentiate it from other sugars). Sucrose consists of a glucose
molecule and a fructose molecule joined together. Another important disaccharide is lactose, consisting of a
glucose molecule and a galactose molecule. As most humans age, the production of lactase, the enzyme that hydrolyzes lactose back into glucose and galactose, typically
decreases. This results in lactase deciency, also called
lactose intolerance.
Sugar polymers are characterized by having reducing or
non-reducing ends. A reducing end of a carbohydrate is
a carbon atom that can be in equilibrium with the openchain aldehyde or keto form. If the joining of monomers
takes place at such a carbon atom, the free hydroxy group
of the pyranose or furanose form is exchanged with an
OH-side-chain of another sugar, yielding a full acetal.
This prevents opening of the chain to the aldehyde or
keto form and renders the modied residue non-reducing.
Lactose contains a reducing end at its glucose moiety,
whereas the galactose moiety form a full acetal with the
C4-OH group of glucose. Saccharose does not have a reducing end because of full acetal formation between the
aldehyde carbon of glucose (C1) and the keto carbon of
fructose (C2).

Glucose

of the most important carbohydrates, is an example of

a monosaccharide. So is fructose, the sugar commonly
associated with the sweet taste of fruits.[15][a] Some carbohydrates (especially after condensation to oligo- and
polysaccharides) contain less carbon relative to H and O,
which still are present in 2:1 (H:O) ratio. Monosaccharides can be grouped into aldoses (having an aldehyde
group at the end of the chain, e.g. glucose) and ketoses
(having a keto group in their chain; e.g. fructose). Both
aldoses and ketoses occur in an equilibrium (starting with
chain lengths of C4) cyclic forms. These are generated by
bond formation between one of the hydroxyl groups of
the sugar chain with the carbon of the aldehyde or keto
group to form a hemiacetal bond. This leads to saturated
ve-membered (in furanoses) or six-membered (in pyra- 4.3
noses) heterocyclic rings containing one O as heteroatom.

4.2

Oligosaccharides and polysaccharides

Disaccharides
CH2OH
H

HO

H
OH
H

CH2OH
O

O H
H
OH

H
OH

HO
H

CH2OH

Cellulose as polymer of -D-glucose

When a few (around three to six) monosaccharides are

joined, it is called an oligosaccharide (oligo- meaning
Sucrose: ordinary table sugar and probably the most familiar few). These molecules tend to be used as markers
carbohydrate.
and signals, as well as having some other uses. Many
monosaccharides joined together make a polysaccharide.
Two monosaccharides can be joined using dehydration They can be joined together in one long linear chain,
synthesis, in which a hydrogen atom is removed from or they may be branched. Two of the most common
the end of one molecule and a hydroxyl group (OH) polysaccharides are cellulose and glycogen, both consistis removed from the other; the remaining residues are ing of repeating glucose monomers.
then attached at the sites from which the atoms were
removed. The HOH or H2 O is then released as a
Cellulose is made by plants and is an important strucmolecule of water, hence the term dehydration. The new
tural component of their cell walls. Humans can neimolecule, consisting of two monosaccharides, is called a
ther
manufacture nor digest it.
disaccharide and is conjoined together by a glycosidic or
ether bond. The reverse reaction can also occur, using a
molecule of water to split up a disaccharide and break the
Glycogen, on the other hand, is an animal carbohyglycosidic bond; this is termed hydrolysis. The most welldrate; humans and other animals use it as a form of
known disaccharide is sucrose, ordinary sugar (in scienenergy storage.

4.4

Use of carbohydrates as an energy ism with far more energy than any oxygen-independent
metabolic feature, and this is thought to be the reason why
source

Main article: Carbohydrate metabolism

complex life appeared only after Earths atmosphere accumulated large amounts of oxygen.

Glucose is the major energy source in most life forms. 4.4.3 Gluconeogenesis
For instance, polysaccharides are broken down into
their monomers (glycogen phosphorylase removes glu- Main article: Gluconeogenesis
cose residues from glycogen). Disaccharides like lactose
or sucrose are cleaved into their two component monosacIn vertebrates, vigorously contracting skeletal muscles
charides.
(during weightlifting or sprinting, for example) do not receive enough oxygen to meet the energy demand, and so
they shift to anaerobic metabolism, converting glucose to
4.4.1 Glycolysis (anaerobic)
lactate. The liver regenerates the glucose, using a process
Glucose is mainly metabolized by a very important ten- called gluconeogenesis. This process is not quite the opstep pathway called glycolysis, the net result of which is to posite of glycolysis, and actually requires three times the
break down one molecule of glucose into two molecules amount of energy gained from glycolysis (six molecules
of pyruvate; this also produces a net two molecules of of ATP are used, compared to the two gained in glyATP, the energy currency of cells, along with two re- colysis). Analogous to the above reactions, the glucose
ducing equivalents as converting NAD+ to NADH. This produced can then undergo glycolysis in tissues that need
does not require oxygen; if no oxygen is available (or the energy, be stored as glycogen (or starch in plants), or be
cell cannot use oxygen), the NAD is restored by convert- converted to other monosaccharides or joined into di- or
ing the pyruvate to lactate (lactic acid) (e.g., in humans) oligosaccharides. The combined pathways of glycolysis
or to ethanol plus carbon dioxide (e.g., in yeast). Other during exercise, lactates crossing via the bloodstream to
of glumonosaccharides like galactose and fructose can be con- the liver, subsequent gluconeogenesis and release [17]
cose into the bloodstream is called the Cori cycle.
verted into intermediates of the glycolytic pathway.[16]

4.4.2

Aerobic

In aerobic cells with sucient oxygen, as in most human

cells, the pyruvate is further metabolized. It is irreversibly
converted to acetyl-CoA, giving o one carbon atom as
the waste product carbon dioxide, generating another reducing equivalent as NADH. The two molecules acetylCoA (from one molecule of glucose) then enter the citric
acid cycle, producing two more molecules of ATP, six
more NADH molecules and two reduced (ubi)quinones
(via FADH2 as enzyme-bound cofactor), and releasing
the remaining carbon atoms as carbon dioxide. The produced NADH and quinol molecules then feed into the
enzyme complexes of the respiratory chain, an electron
transport system transferring the electrons ultimately to
oxygen and conserving the released energy in the form
of a proton gradient over a membrane (inner mitochondrial membrane in eukaryotes). Thus, oxygen is reduced
to water and the original electron acceptors NAD+ and
quinone are regenerated. This is why humans breathe in
oxygen and breathe out carbon dioxide. The energy released from transferring the electrons from high-energy
states in NADH and quinol is conserved rst as proton
gradient and converted to ATP via ATP synthase. This
generates an additional 28 molecules of ATP (24 from the
8 NADH + 4 from the 2 quinols), totaling to 32 molecules
of ATP conserved per degraded glucose (two from glycolysis + two from the citrate cycle). It is clear that using
oxygen to completely oxidize glucose provides an organ-

5 Proteins
Main article: Protein
Like carbohydrates, some proteins perform largely struc-

A schematic of hemoglobin. The red and blue ribbons represent

the protein globin; the green structures are the heme groups.

tural roles. For instance, movements of the proteins actin

and myosin ultimately are responsible for the contraction

5 PROTEINS

of skeletal muscle. One property many proteins have is

that they specically bind to a certain molecule or class of
moleculesthey may be extremely selective in what they
bind. Antibodies are an example of proteins that attach to
one specic type of molecule. In fact, the enzyme-linked
immunosorbent assay (ELISA), which uses antibodies, is
one of the most sensitive tests modern medicine uses to
detect various biomolecules. Probably the most important proteins, however, are the enzymes. These molecules
recognize specic reactant molecules called substrates;
they then catalyze the reaction between them. By lowering the activation energy, the enzyme speeds up that
reaction by a rate of 1011 or more: a reaction that would
normally take over 3,000 years to complete spontaneously
might take less than a second with an enzyme. The enzyme itself is not used up in the process, and is free to
catalyze the same reaction with a new set of substrates.
Using various modiers, the activity of the enzyme can
be regulated, enabling control of the biochemistry of the
cell as a whole.
In essence, proteins are chains of amino acids. An amino
acid consists of a carbon atom bound to four groups. One
is an amino group, NH2 , and one is a carboxylic acid
group, COOH (although these exist as NH3 + and
COO under physiologic conditions). The third is a
simple hydrogen atom. The fourth is commonly denoted
"R and is dierent for each amino acid. There are
20 standard amino acids. Some of these have functions
by themselves or in a modied form; for instance, glutamate functions as an important neurotransmitter. Also if
a glycine amino acid undergoes methylation to a pseudo
alanine amino acid, it is an indication of cancer metastasis.

concerned with local morphology (morphology being the

study of structure). Some combinations of amino acids
will tend to curl up in a coil called an -helix or into a
sheet called a -sheet; some -helixes can be seen in the
hemoglobin schematic above. Tertiary structure is the entire three-dimensional shape of the protein. This shape is
determined by the sequence of amino acids. In fact, a
single change can change the entire structure. The alpha
chain of hemoglobin contains 146 amino acid residues;
substitution of the glutamate residue at position 6 with
a valine residue changes the behavior of hemoglobin
so much that it results in sickle-cell disease. Finally,
quaternary structure is concerned with the structure of
a protein with multiple peptide subunits, like hemoglobin
with its four subunits. Not all proteins have more than
one subunit.[19]
Ingested proteins are usually broken up into single amino
acids or dipeptides in the small intestine, and then absorbed. They can then be joined to make new proteins.
Intermediate products of glycolysis, the citric acid cycle,
and the pentose phosphate pathway can be used to make
all twenty amino acids, and most bacteria and plants possess all the necessary enzymes to synthesize them. Humans and other mammals, however, can synthesize only
half of them. They cannot synthesize isoleucine, leucine,
lysine, methionine, phenylalanine, threonine, tryptophan,
and valine. These are the essential amino acids, since
it is essential to ingest them. Mammals do possess
the enzymes to synthesize alanine, asparagine, aspartate,
cysteine, glutamate, glutamine, glycine, proline, serine,
and tyrosine, the nonessential amino acids. While they
can synthesize arginine and histidine, they cannot produce it in sucient amounts for young, growing animals,
and so these are often considered essential amino acids.

If the amino group is removed from an amino acid, it

leaves behind a carbon skeleton called an -keto acid. Enzymes called transaminases can easily transfer the amino
group from one amino acid (making it an -keto acid) to
another -keto acid (making it an amino acid). This is
Generic amino acids (1) in neutral form, (2) as they exist physi- important in the biosynthesis of amino acids, as for many
of the pathways, intermediates from other biochemical
ologically, and (3) joined together as a dipeptide.
pathways are converted to the -keto acid skeleton, and
Amino acids can be joined via a peptide bond. In this de- then an amino group is added, often via transamination.
acids may then be linked together to make a
hydration synthesis, a water molecule is removed and the The amino
[20]
protein.
peptide bond connects the nitrogen of one amino acids
amino group to the carbon of the others carboxylic acid A similar process is used to break down proteins. It is
group. The resulting molecule is called a dipeptide, and rst hydrolyzed into its component amino acids. Free
short stretches of amino acids (usually, fewer than thirty) ammonia (NH3 ), existing as the ammonium ion (NH4 + )
are called peptides or polypeptides. Longer stretches in blood, is toxic to life forms. A suitable method for
merit the title proteins. As an example, the important excreting it must therefore exist. Dierent tactics have
blood serum protein albumin contains 585 amino acid evolved in dierent animals, depending on the animals
residues.[18]
needs. Unicellular organisms, of course, simply release
The structure of proteins is traditionally described in a hi- the ammonia into the environment. Likewise, bony sh
erarchy of four levels. The primary structure of a protein can release the ammonia into the water where it is quickly
convert the ammonia into
simply consists of its linear sequence of amino acids; for diluted. In general, mammals
[21]
urea
cycle.
urea,
via
the
instance, alanine-glycine-tryptophan-serine-glutamateasparagine-glycine-lysine-".

Secondary structure is

Lipids

fer in the specic sugar found in their chain (e.g., DNA

or deoxyribonucleic acid contains 2-deoxyriboses). Also,
the nitrogenous bases possible in the two nucleic acids are
Main article: Lipid
dierent: adenine, cytosine, and guanine occur in both
RNA and DNA, while thymine occurs only in DNA and
The term lipid composes a diverse range of molecules and uracil occurs in RNA.[24]
to some extent is a catchall for relatively water-insoluble
or nonpolar compounds of biological origin, including waxes, fatty acids, fatty-acid derived phospholipids,
8 Relationship
to
other
sphingolipids, glycolipids, and terpenoids (e.g., retinoids
molecular-scale
biological
and steroids). Some lipids are linear aliphatic molecules,
while others have ring structures. Some are aromatic,
sciences
while others are not. Some are exible, while others are
[22]
rigid.
Most lipids have some polar character in addition to being
largely nonpolar. In general, the bulk of their structure
is nonpolar or hydrophobic (water-fearing), meaning
that it does not interact well with polar solvents like water. Another part of their structure is polar or hydrophilic
(water-loving) and will tend to associate with polar solvents like water. This makes them amphiphilic molecules
(having both hydrophobic and hydrophilic portions). In
the case of cholesterol, the polar group is a mere -OH
(hydroxyl or alcohol). In the case of phospholipids, the
polar groups are considerably larger and more polar, as
described below.
Lipids are an integral part of our daily diet. Most oils
and milk products that we use for cooking and eating
like butter, cheese, ghee etc., are composed of fats.
Vegetable oils are rich in various polyunsaturated fatty
acids (PUFA). Lipid-containing foods undergo digestion
within the body and are broken into fatty acids and glyc- Schematic relationship between biochemistry, genetics, and
erol, which are the nal degradation products of fats and molecular biology
lipids.
Researchers in biochemistry use specic techniques native to biochemistry, but increasingly combine these with
techniques and ideas developed in the elds of genetics,
7 Nucleic acids
molecular biology and biophysics. There has never been
a hard-line among these disciplines in terms of content
Main article: Nucleic acid
and technique. Today, the terms molecular biology and
biochemistry are nearly interchangeable. The following
A nucleic acid is a complex, high-molecular-weight bio- gure is a schematic that depicts one possible view of the
chemical macromolecule composed of nucleotide chains relationship between the elds:
that convey genetic information.[23] The most common
nucleic acids are deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). Nucleic acids are found in all living cells and viruses. Aside from the genetic material of
the cell, nucleic acids often play a role as second messengers, as well as forming the base molecule for adenosine
triphosphate (ATP), the primary energy-carrier molecule
found in all living organisms.

Biochemistry is the study of the chemical substances

and vital processes occurring in living organisms.
Biochemists focus heavily on the role, function, and
structure of biomolecules. The study of the chemistry behind biological processes and the synthesis
of biologically active molecules are examples of biochemistry.

Nucleic acid, so called because of its prevalence in cellular nuclei, is the generic name of the family of biopolymers. The monomers are called nucleotides, and each
consists of three components: a nitrogenous heterocyclic
base (either a purine or a pyrimidine), a pentose sugar,
and a phosphate group. Dierent nucleic acid types dif-

Genetics is the study of the eect of genetic differences on organisms. Often this can be inferred
by the absence of a normal component (e.g., one
gene). The study of "mutants" organisms with a
changed gene that leads to the organism being different with respect to the so-called "wild type" or

11
normal phenotype. Genetic interactions (epistasis)
can often confound simple interpretations of such
knock-out or knock-in studies.
Molecular biology is the study of molecular underpinnings of the process of replication, transcription and translation of the genetic material. The
central dogma of molecular biology where genetic
material is transcribed into RNA and then translated
into protein, despite being an oversimplied picture
of molecular biology, still provides a good starting point for understanding the eld. This picture,
however, is undergoing revision in light of emerging
novel roles for RNA.[25]

REFERENCES

Proteolysis
Small molecule
Structural biology

10 Notes

a. ^ Fructose is not the only sugar found in fruits. Glucose and sucrose are also found in varying quantities in
various fruits, and indeed sometimes exceed the fructose
present. For example, 32% of the edible portion of date is
glucose, compared with 23.70% fructose and 8.20% su Chemical biology seeks to develop new tools based crose. However, peaches contain more sucrose (6.66%)
[26]
on small molecules that allow minimal perturbation than they do fructose (0.93%) or glucose (1.47%).
of biological systems while providing detailed information about their function. Further, chemical
biology employs biological systems to create non- 11 References
natural hybrids between biomolecules and synthetic
devices (for example emptied viral capsids that can [1] Biochemistry. acs.org.
deliver gene therapy or drug molecules).
[2] scientic term 'biochemistry'".

See also

Main article: Outline of biochemistry

9.1

Lists

Important publications in biochemistry (chemistry)

List of biochemistry topics
List of biochemists
List of biomolecules

9.2

See also

[3] Fiske, John (1890). Outlines of Cosmic Philosophy Based

on the Doctrines of Evolution, with Criticisms on the Positive Philosophy, Volume 1. Boston and New York:
Houghton, Miin. pp. 41920. Retrieved 16 February
2015.
[4] Kauman, G.B.; Chooljian, S.H. (2001). Friedrich
Whler (18001882), on the bicentennial of his
birth.
The Chemical Educator 6 (2): 121133.
doi:10.1007/s00897010444a.
[5] Hunter (2000), p. 75.
[6] Hunter (2000), pp. 9698.
[7] Ben-Menahem, Ari (2009). Historical Encyclopedia of
Natural and Mathematical Sciences. Springer. p. 2982.
ISBN 978-3-540-68831-0.
[8] Tropp (2012), p. 2.
[9] Tropp (2012), pp. 1920.

Chemical ecology
Computational biomodeling
EC number
Hypothetical types of biochemistry
International Union of Biochemistry and Molecular
Biology
Metabolome
Metabolomics
Molecular medicine
Plant biochemistry

[10] Krebs, Jocelyn E.; Goldstein, Elliott S.; Lewin, Benjamin;

Kilpatrick, Stephen T. (2012). Essential Genes. Jones &
Bartlett Publishers. p. 32. ISBN 978-1-4496-1265-8.
[11] Butler, John M. (2009). Fundamentals of Forensic DNA
Typing. Academic Press. p. 5. ISBN 978-0-08-0961767.
[12] Sen, Chandan K.; Roy, Sashwati (2007). miRNA:
Licensed to kill the messenger. DNA Cell Biology
26 (4): 193194. doi:10.1089/dna.2006.0567. PMID
17465885.
[13] Ultratrace minerals. Authors: Nielsen, Forrest H. USDA,
ARS Source: Modern nutrition in health and disease / editors, Maurice E. Shils ... et al.. Baltimore : Williams &
Wilkins, c1999., p. 283-303. Issue Date: 1999 URI:

[14] Fariselli, Piero; Rossi, Ivan; Capriotti, Emidio; Casadio,

Rita (2007). The WWWH of remote homolog detection:
the state of the art. Briengs in Bioinformatics 8 (2): 78
87. doi:10.1093/bib/bbl032. PMID 17003074.
[15] Whiting, G.C (1970). Sugars. In A.C. Hulme. The Biochemistry of Fruits and their Products. Volume 1. London
& New York: Academic Press. pp. 131.
[16] Fromm and Hargrove (2012), pp. 163180.
[17] Fromm and Hargrove (2012), pp. 183194.
[18] Metzler, David Everett; Metzler, Carol M. (2001).
Biochemistry: The Chemical Reactions of Living Cells 1.
Academic Press. p. 58. ISBN 978-0-12-492540-3.
[19] Fromm and Hargrove (2012), pp. 3551.
[20] Fromm and Hargrove (2012), pp. 279292.
[21] Sherwood, Lauralee; Klandorf, Hillar; Yancey, Paul H.
(2012). Animal Physiology: From Genes to Organisms.
Cengage Learning. p. 558. ISBN 978-0-8400-6865-1.
[22] Fromm and Hargrove (2012), pp. 2227.
[23] Voet, D. and Voet, J, G. (2011). Biochemistry (4th ed.).
John Wiley & Sons Inc.: Hoboken, NJ
[24] Tropp (2012), pp. 59.
[25] Ulveling, Damien; Francastel, Claire; Hub, Florent (2011). When one is better than two: RNA
with dual functions. Biochimie 93 (4): 633644.
doi:10.1016/j.biochi.2010.11.004. PMID 21111023.
[26] Whiting, G.C. (1970), p.5

11.1

Cited literature

Fromm, Herbert J.; Hargrove, Mark (2012). Essentials of Biochemistry. Springer. ISBN 978-3-64219623-2.
Hunter, Graeme K. (2000). Vital Forces: The Discovery of the Molecular Basis of Life. Academic
Press. ISBN 978-0-12-361811-5.
Tropp, Burton E. (2012). Molecular Biology (4th
ed.). Jones & Bartlett Learning. ISBN 978-1-44960091-4.

12

External links

The Virtual Library of Biochemistry and Cell Biology

Biochemistry, 5th ed. Full text of Berg, Tymoczko,
and Stryer, courtesy of NCBI.
Biochemistry, 2nd ed. Full text of Garrett and Grisham.

Biochemistry Animation (Narrated Flash animations.)

SystemsX.ch - The Swiss Initiative in Systems Biology
Biochemistry Online Resources Lists of Biochemistry departments, websites, journals, books and reviews, employment opportunities and events.
Full text of Biochemistry by Kevin and Indira, an
introductory biochemistry textbook.

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