DNA

From Wikipedia, the free encyclopedia.

For other uses, see DNA (disambiguation).

Space-filling model of a section of DNA molecule

Deoxyribonucleic acid (DNA) is a nucleic acid that contains the genetic instructions
specifying the biological development of all cellular forms of life (and many viruses).
DNA is often referred to as the molecule of heredity, as it is responsible for the
genetic propagation of most inherited traits. During reproduction, DNA is replicated
and transmitted to the offspring.

In bacteria (prokaryotes), DNA is not separated from the cytoplasm by a nuclear

envelope. By contrast, in the complex cells that make up other organisms (plants,
animals, fungi and protists), most of the DNA is located in the cell nucleus. The
energy-generating organelles known as chloroplasts and mitochondria also carry
DNA, as do many viruses.

Basically what the description above says is that DNA is a basic building block of life
and that it contains all the information that living things need to function correctly.
For example, in humans this can range from your hair colour to the ability to roll your
tongue and to any herditary diseases.

Your DNA is composed of genes inherited from both your mother and father. DNA
from both of your parents was combined to form the genome of a fertilized egg (that's
you), then that egg divided many times, copying the DNA before every division, until
all the cells of your body were formed. As a result, each cell in your body contains
copies of the DNA from your parents (except red blood cells, which lose their
chromosomal DNA but cannot divide).

Contents
[hide]

• 1 DNA in brief
• 2 DNA in practice
o 2.1 DNA in crime
o 2.2 DNA in computation
• 3 Overview of molecular structure
• 4 The role of the sequence
• 5 DNA replication
• 6 Mechanical properties relevant to biology
o 6.1 Strands association and dissociation
o 6.2 Circular DNA
o 6.3 Great length versus tiny breadth
o 6.4 Different helix geometries
 6.4.1 Supercoiled DNA
 6.4.2 Conditions for formation of
A and Z helices
 6.4.3 Table of comparison of the
properties of different helical
forms
o 6.5 Non-helical forms
• 7 Direction of DNA strands
o 7.1 Chemical nomenclature (5' and 3')
o 7.2 Sense and antisense
o 7.3 An exception: viruses
o 7.4 As viewed by topologists
• 8 Single-stranded DNA (ssDNA) and repair of
mutations
• 9 The history of DNA research
o 9.1 First isolation of DNA
o 9.2 Establishing a link between heritable
traits and chromosomes
o 9.3 Discovery of the structure of DNA
 9.3.1 Discovery that DNA is
helical
 9.3.2 Discovery that
complementary nucleotides occur
in equal proportions
 9.3.3 Watson and Crick's model
 9.3.4 Publishing of the "Central
Dogma"
• 10 Bibliography

• 11 External links
 [edit]

DNA in brief

Unsolved problems in biology: Do all organisms link together to a primary source? Given a DNA
sequence, what shape will the protein fold into? Given a particular desired shape, what DNA sequence
will produce it? What are all the functions of the DNA? The building block of life may be a precursor
to a generation of electronic devices and computers, but what are the electronic properties of DNA? Is
junk DNA only molecular garbage?

DNA Under electron microscope

This section presents a brief and simple overview of DNA.

• Genes can be loosely viewed as the organism's "cookbook" or "blueprint";

• A strand of DNA contains genes, areas that regulate genes, and areas that
either have no function, or a function we do not (yet) know;
• DNA is organized as two complementary strands, head-to-toe, with bonds
between them that can be "unzipped" like a zipper, separating the strands;
• DNA is encoded with four interchangeable "building blocks", called "bases",
which can be abbreviated A, T, C, and G; each base "pairs up" with only one
other base: A+T, T+A, C+G and G+C; that is, an "A" on one strand of double-
stranded DNA will "mate" properly only with a "T" on the other,
complementary strand;
• The order does matter: A+T is not the same as T+A, just as C+G is not the
same as G+C;
• However, since there are just four possible combinations, naming only one
base on the conventionally chosen side of the strand is enough to describe the
sequence;
• The order of the bases along the length of the DNA is what it's all about, the
sequence itself is the description for genes;
• Replication is performed by splitting (unzipping) the double strand down the
middle via relatively trivial chemical reactions, and recreating the "other half"
of each new single strand by drowning each half in a "soup" made of the four
bases. Since each of the "bases" can only combine with one other base, the
base on the old strand dictates which base will be on the new strand. This way,
each split half of the strand plus the bases it collects from the soup will ideally
end up as a complete replica of the original, unless a mutation occurs;
 • Mutations are simply chemical imperfections in this process: a base is
accidentally skipped, inserted, or incorrectly copied, or the chain is trimmed,
or added to; all other basic mutations can be described as combinations of
these accidental "operations".

[edit]

DNA in practice
[edit]

DNA in crime

Forensic scientists can use DNA located in blood, semen, or hair left at the scene of a
crime to identify a possible suspect, a process called DNA profiling or genetic
fingerprinting. In DNA profiling the relative lengths of sections of repetitive DNA,
such as short tandem repeats and minisatellites, are compared. DNA profiling was
developed in 1984 by English geneticist Alec Jeffreys, and was first used in 1986 in
the Enderby murders case in Leicestershire, England. Many jurisdictions require
convicts of certain types of crimes to provide a sample of DNA for inclusion in a
computerized database. This has helped investigators solve old cases where the
perpetrator was unknown and only a DNA sample was obtained from the scene
(particularly in rape cases between strangers). This method is one of the most reliable
techniques for identifying a criminal, but is not always perfect, for example if no
DNA can be retrieved, or if the scene is contaminated with the DNA of several
possible suspects.

[edit]

DNA in computation

Despite its biological origins, DNA plays an important role in computer science, both
as a motivating research problem and as a method of computation in itself, called
DNA computing.

As a simple example, research on string searching algorithms, which find an

occurence of a sequence of letters inside a larger sequence of letters, was motivated
by DNA research, where it is used to find specific sequences of nucleotides in a large
sequence. In other applications like text editors, even simple algorithms for this
problem usually suffice, but DNA sequences cause these algorithms to exhibit near-
worst-case behavior due to their small number of distinct characters.

Databases have also been strongly motivated by DNA research, which requires special
tools for storing and manipulating DNA sequences. Databases specialized for this
purpose are called genomic databases, and have a number of unique technical
challenges associated with the operations of approximate matching, sequence
comparison, finding repeating patterns, and homology searching.
In 1994, Leonard Adleman of the University of Southern California made headlines
when he discovered a way of solving the directed Hamiltonian path problem, an NP-
complete problem, using tools from molecular biology, in particular DNA. The new
approach, dubbed DNA computing, has practical advantages over traditional
computers in power use, space use, and efficiency, due to its ability to highly
parallelize the computation (see parallel computing). A number of other problems,
including simulation of various abstract machines, the boolean satisfiability problem,
and the bounded version of the Post correspondence problem, have since been tackled
using DNA computing.

Due to its compactness, DNA also has an important role in cryptography, where in
particular it allows unbreakable one-time pads to be efficiently constructed and
used.[1]

[edit]

Overview of molecular structure

Schematic representation of the DNA which illustrates its double helix structure

Although sometimes called "the molecule of heredity", pieces of DNA as people

typically think of them are not single molecules. Rather, they are pairs of molecules,
which entwine like vines to form a double helix (see the illustration at the right).

Each vine-like molecule is a strand of DNA: a chemically linked chain of

nucleotides, each of which consists of a sugar, a phosphate and one of four kinds
of nucleobases ("bases"). Because DNA strands are composed of these nucleotide
subunits, they are polymers.

The diversity of the bases means that there are four kinds of nucleotides, which are
commonly referred to by the identity of their bases. These are adenine (A), thymine
(T), cytosine (C), and guanine (G).

In a DNA double helix, two polynucleotide strands can associate through the
hydrophobic effect and pi stacking. Specificity of which strands stay associated is
determined by complementary pairing. Each base forms hydrogen bonds readily to
only one other -- A to T and C to G -- so that the identity of the base on one strand
dictates the strength of the association; the more complementary bases exist, the
stronger and longer-lasting the association.

The cell's machinery is capable of melting or disassociating a DNA double helix, and
using each DNA strand as a template for synthesizing a new strand which is nearly
identical to the previous strand. Errors that occur in the synthesis are known as
mutations. The process known as PCR mimics this process in vitro in a nonliving
system.

Because pairing causes the nucleotide bases to face the helical axis, the sugar and
phosphate groups of the nucleotides run along the outside; the two chains they form
are sometimes called the "backbones" of the helix. In fact, it is chemical bonds
between the phosphates and the sugars that link one nucleotide to the next in the DNA
strand.

Rotating DNA stick model (info)

Animation of a section of DNA rotating. (1.00 MB, animated GIF format).
Problems seeing the videos? Media help.

[edit]

The role of the sequence

Within a gene, the sequence of nucleotides along a DNA strand defines a messenger
RNA sequence which then defines a protein, that an organism is liable to manufacture
or "express" at one or several points in its life using the information of the sequence.
The relationship between the nucleotide sequence and the amino-acid sequence of the
protein is determined by simple cellular rules of translation, known collectively as the
genetic code. The genetic code is made up of three-letter 'words' (termed a codon)
formed from a sequence of three nucleotides (e.g. ACT, CAG, TTT). These codons
can then be translated with messenger RNA and then transfer RNA, with a codon
corresponding to a particular amino acid. Since there are 64 possible codons, most
amino acids have more than one possible codon. There are also three 'stop' or
'nonsense' codons signifying the end of the coding region, namely the UAA, UGA and
UAG codons.

In many species, only a small fraction of the total sequence of the genome appears to
encode protein. For example, only about 1.5% of the human genome consists of
protein-coding exons. The function of the rest is a matter of speculation. It is known
that certain nucleotide sequences specify affinity for DNA binding proteins, which
play a wide variety of vital roles, in particular through control of replication and
transcription. These sequences are frequently called regulatory sequences, and
researchers assume that so far they have identified only a tiny fraction of the total that
exist. "Junk DNA" represents sequences that do not yet appear to contain genes or to
have a function. The reasons for the presence of so much non-coding DNA in
eukaryotic genomes and the extraordinary differences in genome size ("C-value")
among species represent a long-standing puzzle in DNA research known as the "C-
value enigma".

Sequence also determines a DNA segment's susceptibility to cleavage by restriction

enzymes, the quintessential tools of genetic engineering. The position of cleavage
sites throughout an individual's genome determines one kind of an individual's "DNA
fingerprint".

[edit]

DNA replication
Main article: DNA replication

DNA replication

DNA replication or DNA synthesis is the process of copying the double-stranded

DNA prior to cell division. The two resulting double strands are generally almost
perfectly identical, but occasionally errors in replication can result in a less than
perfect copy (see mutation), and each of them consists of one original and one newly
synthesized strand. This is called semiconservative replication. The process of
replication consists of three steps: initiation, replication and termination.

[edit]

Mechanical properties relevant to biology

Main article: Mechanical properties of DNA.

[edit]

Strands association and dissociation

The hydrogen bonds between the strands of the double helix are weak enough that
they can be easily separated by enzymes. Enzymes known as helicases unwind the
strands to facilitate the advance of sequence-reading enzymes such as DNA
polymerase. The unwinding requires that helicases chemically cleave the phosphate
backbone of one of the strands so that it can swivel around the other. The strands can
also be separated by gentle heating, as used in PCR, provided they have fewer than
about 10,000 base pairs (10 kilobase pairs, or 10 kbp). The intertwining of the DNA
strands makes long segments difficult to separate.

[edit]

Circular DNA

When the ends of a piece of double-helical DNA are joined so that it forms a circle, as
in plasmid DNA, the strands are topologically knotted. This means they cannot be
separated by gentle heating or by any process that does not involve breaking a strand.
The task of unknotting topologically linked strands of DNA falls to enzymes known
as topoisomerases. Some of these enzymes unknot circular DNA by cleaving two
strands so that another double-stranded segment can pass through. Unknotting is
required for the replication of circular DNA as well as for various types of
recombination in linear DNA.

[edit]

Great length versus tiny breadth

The narrow breadth of the double helix makes it impossible to detect by conventional
electron microscopy, except by heavy staining. At the same time, the DNA found in
many cells can be macroscopic in length -- approximately 5 centimetres long for
strands in a human chromosome. Consequently, cells must compact or "package"
DNA to carry it within them. This is one of the functions of the chromosomes, which
contain spool-like proteins known as histones, around which DNA winds.

[edit]

Different helix geometries

The DNA helix can assume one of three slightly different geometries, of which the
"B" form described by James D. Watson and Francis Crick is believed to predominate
in cells. It is 2 nanometres wide and extends 3.4 nanometres per 10 bp of sequence.
This is also the approximate length of sequence in which the double helix makes one
complete turn about its axis. This frequency of twist (known as the helical pitch)
depends largely on stacking forces that each base exerts on its neighbors in the chain.
[edit]

Supercoiled DNA

The B form of the DNA helix twists 360° per 10.6 bp in the absence of strain. But
many molecular biological processes can induce strain. A DNA segment with excess
or insufficient helical twisting is referred to, respectively, as positively or negatively
"supercoiled". DNA in vivo is typically negatively supercoiled, which facilitates the
unwinding of the double-helix required for RNA transcription.

[edit]

Conditions for formation of A and Z helices

The two other known double-helical forms of DNA, called A and Z, differ modestly in
their geometry and dimensions. The A form appears likely to occur only in dehydrated
samples of DNA, such as those used in crystallographic experiments, and possibly in
hybrid pairings of DNA and RNA strands. Segments of DNA that cells have
methylated for regulatory purposes may adopt the Z geometry, in which the strands
turn about the helical axis like a mirror image of the B form.

[edit]

Table of comparison of the properties of different helical forms

Geometry attribute A-form B-form Z-form

Helix sense right-handed right-handed left-handed
Repeating unit 1 bp 1 bp 2 bp
Rotation/bp 33.6° 35.9° 60°/2
Mean bp/turn 10.7 10.0 12
Inclination of bp to axis +19° -1.2° -9°
Rise/bp along axis 0.23 nm 0.332 nm 0.38 nm
Pitch/turn of helix 2.46 nm 3.32 nm 4.56 nm
Mean propeller twist +18° +16° 0°
C: anti,
Glycosyl angle anti anti
G: syn
C: C2'-endo,
Sugar pucker C3'-endo C2'-endo
G: C2'-exo
Diameter 26 nm 20 nm 18 nm
[edit]

Non-helical forms

Other, including non-helical, forms of DNA have been described, for example a side-
by-side (SBS) configuration. Indeed, it is far from certain that the B-form double
helix is the dominant form in living cells.

[edit]
Direction of DNA strands
The asymmetric shape and linkage of nucleotides means that a DNA strand always
has a discernible orientation or directionality. Because of this directionality, close
inspection of a double helix reveals that nucleotides are heading one way along one
strand (the "ascending strand"), and the other way along the other strand (the
"descending strand"). This arrangement of the strands is called antiparallel.

[edit]

Chemical nomenclature (5' and 3')

For reasons of chemical nomenclature, people who work with DNA refer to the
asymmetric ends of each strand as the 5' and 3' ends (pronounced "five prime" and
"three prime"). DNA workers and enzymes alike always read nucleotide sequences in
the "5' to 3' direction". In a vertically oriented double helix, the 3' strand is said to be
ascending while the 5' strand is said to be descending.

[edit]

Sense and antisense

As a result of their antiparallel arrangement and the sequence-reading preferences of

enzymes, even if both strands carried identical instead of complementary sequences,
cells could properly translate only one of them. The other strand a cell can only read
backwards. Molecular biologists call a sequence "sense" if it is translated or
translatable, and they call its complement "antisense". It follows then, somewhat
paradoxically, that the template for transcription is the antisense strand. The resulting
transcript is an RNA replica of the sense strand and is itself sense.

[edit]

An exception: viruses

Some viruses blur the distinction between sense and antisense, because certain
sequences of their genomes do double duty, encoding one protein when read 5' to 3'
along one strand, and a second protein when read in the opposite direction along the
other strand. As a result, the genomes of these viruses are unusually compact for the
number of genes they contain, which biologists view as an adaptation.

[edit]

As viewed by topologists

Topologists like to note that the juxtaposition of the 3′ end of one DNA strand beside
the 5′ end of the other at both ends of a double-helical segment makes the
arrangement a "crab canon".

[edit]
Single-stranded DNA (ssDNA) and repair of
mutations
In some viruses DNA appears in a non-helical, single-stranded form. Because many of
the DNA repair mechanisms of cells work only on paired bases, viruses that carry
single-stranded DNA genomes mutate more frequently than they would otherwise. As
a result, such species may adapt more rapidly to avoid extinction. The result would
not be so favorable in more complicated and more slowly replicating organisms,
however, which may explain why only viruses carry single-stranded DNA. These
viruses presumably also benefit from the lower cost of replicating one strand versus
two.

[edit]

The history of DNA research

James Watson in the Cavendish Laboratory at the University of Cambridge

The discovery that DNA was the carrier of genetic information was a process that
required many earlier discoveries. The existence of DNA was discovered in the mid
19th century. However, it was only in the early 20th century that researchers began
suggesting that it might store genetic information. This was only accepted after the
structure of DNA was elucidated by Watson and Crick in their 1953 Nature
publication. Watson and Crick proposed the central dogma of molecular biology in
1957, describing the process whereby proteins are produced from nucleic DNA.

[edit]

First isolation of DNA

Working in the 19th century, biochemists initially isolated DNA and RNA (mixed
together) from cell nuclei. They were relatively quick to appreciate the polymeric
nature of their "nucleic acid" isolates, but realized only later that nucleotides were of
two types--one containing ribose and the other deoxyribose. It was this subsequent
discovery that led to the identification and naming of DNA as a substance distinct
from RNA.

Friedrich Miescher (1844-1895) discovered a substance he called "nuclein" in 1869.

Somewhat later, he isolated a pure sample of the material now known as DNA from
the sperm of salmon, and in 1889 his pupil, Richard Altmann, named it "nucleic acid".
This substance was found to exist only in the chromosomes.

[edit]

Establishing a link between heritable traits and chromosomes

Max Delbrück, Nikolai V. Timofeeff-Ressovsky, and Karl G. Zimmer published

results in 1935 suggesting that chromosomes are very large molecules the structure of
which can be changed by treatment with X-rays, and that by so changing their
structure it was possible to change the heritable characteristics governed by those
chromosomes. (Delbrück and Salvador Luria were awarded the Nobel Prize in 1969
for their work on the genetic structure of viruses.) In 1943, Oswald Theodore Avery
discovered that traits proper to the "smooth" form of the Pneumococcus could be
transferred to the "rough" form of the same bacteria merely by making the killed
"smooth" (S) form available to the live "rough" (R) form. Quite unexpectedly, the
living R Pneumococcus bacteria were transformed into a new strain of the S form, and
the transferred S characteristics turned out to be heritable. Avery called the medium of
transfer of traits the transforming principle; he identified DNA as the transforming
principle, and not protein as previously thought. In 1953, Alfred Hershey and Martha
Chase did an experiment (Hershey-Chase experiment) that showed, in T2 phage, that
DNA is the genetic material (Hershey shared the Nobel prize with Luria).

Francis Crick's first sketch of the deoxyribonucleic acid double-helix pattern

In 1944, the renowned physicist, Erwin Schrödinger, published a brief book entitled
What is Life?, where he maintained that chromosomes contained what he called the
"hereditary code-script" of life. He added: "But the term code-script is, of course, too
narrow. The chromosome structures are at the same time instrumental in bringing
about the development they foreshadow. They are law-code and executive power -- or,
to use another simile, they are architect's plan and builder's craft -- in one." He
conceived of these dual functional elements as being woven into the molecular
structure of chromosomes. By understanding the exact molecular structure of the
chromosomes one could hope to understand both the "architect's plan" and also how
that plan was carried out through the "builder's craft." Francis Crick, James D.
Watson, Maurice Wilkins, Rosalind Franklin, Seymour Benzer, et al., took up the
physicist's challenge to work out the structure of the chromosomes and the question of
how the segments of the chromosomes that were conceived to relate to specific traits
could possibly do their jobs.

Just how the presence of specific features in the molecular structure of chromosomes
could produce traits and behaviors in living organisms was unimaginable at the time.
Because chemical dissection of DNA samples always yielded the same four
nucleotides, the chemical composition of DNA appeared simple, perhaps even
uniform. Organisms, on the other hand, are fantastically complex individually and
widely diverse collectively. Geneticists did not speak of genes as conveyors of
"information" in such words, but if they had, they would not have hesitated to
quantify the amount of information that genes need to convey as vast. The idea that
information might reside in a chemical in the same way that it exists in text--as a
finite alphabet of letters arranged in a sequence of unlimited length--had not yet been
conceived. It would emerge upon the discovery of DNA's structure, but few
researchers imagined that DNA's structure had much to say about genetics.

[edit]

Discovery of the structure of DNA

In the 1950s, only a few groups made it their goal to determine the structure of DNA.
These included an American group led by Linus Pauling, and two groups in Britain.
At the University of Cambridge, Crick and Watson were building physical models
using metal rods and balls, in which they incorporated the known chemical structures
of the nucleotides, as well as the known position of the linkages joining one
nucleotide to the next along the polymer. At King's College, London, Maurice Wilkins
and Rosalind Franklin were examining X-ray diffraction patterns of DNA fibers. Of
the three groups, only the London group was able to produce good quality diffraction
patterns and thus produce sufficient quantitative data about the structure
The chemical structure of DNA
[edit]

Discovery that DNA is helical

A key inspiration in the work of all of these teams was the discovery in 1948 by
Pauling that many proteins included helical (see alpha helix) shapes. Pauling had
deduced this structure from X-ray patterns. Even in the initial diffraction data from
DNA by Maurice Wilkins, it was evident that the structure involved helices. But this
insight was only a beginning. There remained the questions of how many strands
came together, whether this number was the same for every helix, whether the bases
pointed toward the helical axis or away, and ultimately what were the explicit angles
and coordinates of all the bonds and atoms. Such questions motivated the modeling
efforts of Watson and Crick.

[edit]

Discovery that complementary nucleotides occur in equal proportions

In their modeling, Watson and Crick restricted themselves to what they saw as
chemically and biologically reasonable. Still, the breadth of possibilities was very
wide. A breakthrough occurred in 1952, when Erwin Chargaff visited Cambridge and
inspired Crick with a description of experiments Chargaff had published in 1947.
Chargaff had observed that the proportions of the four nucleotides vary between one
DNA sample and the next, but that for particular pairs of nucleotides -- adenine and
thymine, guanine and cytosine -- the two nucleotides are always present in equal
proportions.

[edit]

Watson and Crick's model

Crick and Watson DNA model built in 1953, currently on display at the National
Science Museum in London.

Watson and Crick had begun to contemplate double helical arrangements, but they
lacked information about the amount of twist (pitch) and the distance between the two
strands. Fortunately Rosalind Franklin had to disclose some of her findings for the
Medical Research Council and Crick saw this material through MaxPerutz's links to
the MRC. Franklin's work confirmed a double helix that was on the outside of the
molecule and also gave an insight into its symmetry, in particular that the two helical
strands ran in opposite directions.

Watson and Crick were again greatly assisted by more of Franklin's data. This is
controversial because Franklin's critical X-ray pattern was shown to Watson and Crick
without Franklin's knowledge or permission. Wilkins showed the famous Photo 51 to
Watson at his lab immediately after Watson had been unsucessful in asking Franklin
to collaborate to beat Pauling in finding the structure.

From the data in photograph 51 Watson and Crick were able to discern that not only
was the distance between the two strands was constant, but also to measure its exact
value of 2 nanometres. The same photograph also gave them the 3.4 nanometre-per-
10 bp "pitch" of the helix.

The final insight came when Crick and Watson saw that a complementary pairing of
the bases could provide an explanation for Chargaff's puzzling finding. However the
structure of the bases had been incorrectly guessed in the textbooks as the enol
tautomer when they were more likely to be in the keto form. When Jerry Donohue
pointed this fallacy out to Watson, Watson quickly realised that the pairs of adenine
and thymine, and guanine and cytosine were almost identical in shape and so would
provide equally sized 'rungs' between the two strands. With the base-pairing, the
Watson and Crick quickly converged upon a model, which they announced before
Franklin herself had published any of her work.

Franklin was two steps away from the solution. She had not guessed the base-pairing
and had not appreciated the implications of the symmetry that she had described.
However she had been working almost alone and did not have regular contact with a
partner like Crick and Watson, and with other experts such as Jerry Donohoe. Her
notebooks show that she was aware both of Jerry Donohue's work concerning
tautomeric forms of bases (she used the keto forms for three of the bases) and of
Chargaff's work.

The disclosure of Franklin's data to Watson has angered some people who believe
Franklin did not receive the credit due to her at the time and that she might have
discovered the structure on her own before Crick and Watson. In Crick and Watson's
famous paper in Nature in 1953, they said that their work had been stimulated by the
work of Wilkins and Franklin, whereas it had been the basis of their work. However
they had agreed with Wilkins and Franklin that they all should publish papers in the
same issue of Nature in support of the proposed structure.

[edit]

Publishing of the "Central Dogma"

Watson and Crick's model attracted great interest immediately upon its presentation.
Arriving at their conclusion on February 21, 1953, Watson and Crick made their first
announcement on February 28. Their paper 'A Structure for Deoxyribose Nucleic
Acid' was published on April 25. In an influential presentation in 1957, Crick laid out
the "Central Dogma", which foretold the relationship between DNA, RNA, and
proteins, and articulated the "sequence hypothesis." A critical confirmation of the
replication mechanism that was implied by the double-helical structure followed in
1958 in the form of the Meselson-Stahl experiment. Work by Crick and coworkers
showed that the genetic code was based on non-overlapping triplets of codons, and
Har Gobind Khorana and others deciphered the genetic code not long afterward.
These findings represent the birth of molecular biology.

Watson, Crick, and Wilkins were awarded the 1962 Nobel Prize for Medicine for
discovering the molecular structure of DNA, by which time Franklin had died. Nobel
prizes are not awarded posthumously.

[edit]

Bibliography
• DNA: The Secret of Life, by James D. Watson. ISBN 0-375-41546-7
• The Double Helix: A Personal Account of the Discovery of the Structure of
DNA (Norton Critical Editions), by James D. Watson. ISBN 0393950751

[edit]

External links
• Defending Franklin's Legacy
• 17 April, 2003, BBC News: Most ancient DNA ever?
• DNA Interactive (requires Macromedia Flash)
• Free DNA Images
 • DNA: PDB molecule of the month
• DNA under electron microscope
• My First Book About DNA Designed for children to learn more about DNA.
• Nucleic Acids at the Open Directory Project
• Rotten Library Article on DNA
• Watson, James, and Francis Crick, "Molecular structure of nucleic acids, A
structure for Deoxyribose Nucleic Acid". April 2, 1953. (paper on the structure
of DNA)

Nucleic acids edit

Nucleobases: Adenine - Thymine - Uracil - Guanine - Cytosine - Purine - Pyrimidine
Nucleosides: Adenosine - 5-Methyluridine - Uridine - Guanosine - Cytidine - Deoxyadenosine -
Thymidine - Deoxyuridine - Deoxyguanosine - Deoxycytidine - Ribose - Deoxyribose

Nucleotides: AMP - m5UMP - UMP - GMP - CMP - ADP - m5UDP - UDP - GDP - CDP - ATP -
m5UTP - UTP - GTP - CTP - cAMP - cGMP

Deoxynucleotides: dAMP - dTMP - dUMP - dGMP - dCMP - dADP - dTDP - dUDP - dGDP - dCDP -
dATP - dTTP - dUTP - dGTP - dCTP

Nucleic acids: DNA - RNA - LNA - PNA - mRNA - ncRNA - miRNA - rRNA - shRNA - siRNA -
tRNA - mtDNA - Oligonucleotide