A hairpin loop from a pre-mRNA. Highlighted are the nucleobases (green) and the ribose-phosphate backbone (blue).

Ribonucleic acid /raɪbɵ.njuːˌkleɪ.ɨk ˈæsɪd/, or RNA, is part of a group of molecules known as the nucleic acids, which are one of the four major macromolecules (along with lipids, carbohydrates and proteins) essential for all known forms of life. Like DNA, RNA is made up of a long chain of components called nucleotides. Each nucleotide consists of a nucleobase, a ribose sugar, and a phosphate group. The sequence of nucleotides allows RNA to encode genetic information. All cellular organisms use messenger RNA (mRNA) to carry the genetic information that directs the synthesis of proteins. In addition, many viruses use RNA instead of DNA as their genetic material.

Some RNA molecules play an active role in cells by catalyzing biological reactions, controlling gene expression, or sensing and communicating responses to cellular signals. One of these active processes is protein synthesis, a universal function whereby mRNA molecules direct the assembly of proteins on ribosomes. This process uses transfer RNA (tRNA) molecules to deliver amino acids to the ribosome, where ribosomal RNA (rRNA) links amino acids together to form proteins.

The chemical structure of RNA is very similar to that of DNA, with two differences: (a) RNA contains the sugar ribose, while DNA contains the slightly different sugar deoxyribose (a type of ribose that lacks one oxygen atom), and (b) RNA has the nucleobase uracil while DNA contains thymine. Unlike DNA, most RNA molecules are single-stranded and can adopt very complex three-dimensional structures.

Comparison with DNA[link]

Three-dimensional representation of the 50S ribosomal subunit. RNA is in ochre, protein in blue. The active site is in the middle (red).

RNA and DNA are both nucleic acids, but differ in three main ways:

• Unlike double-stranded DNA, RNA is a single-stranded molecule in many of its biological roles and has a much shorter chain of nucleotides.
• While DNA contains deoxyribose, RNA contains ribose (in deoxyribose there is no hydroxyl group attached to the pentose ring in the 2' position). These hydroxyl groups make RNA less stable than DNA because it is more prone to hydrolysis.
• The complementary base to adenine is not thymine, as it is in DNA, but rather uracil, which is an unmethylated form of thymine.^[1]

Like DNA, most biologically active RNAs, including mRNA, tRNA, rRNA, snRNAs, and other non-coding RNAs, contain self-complementary sequences that allow parts of the RNA to fold^[2] and pair with itself to form double helices. Analysis of these RNAs has revealed that they are highly structured. Unlike DNA, their structures do not consist of long double helices but rather collections of short helices packed together into structures akin to proteins. In this fashion, RNAs can achieve chemical catalysis, like enzymes.^[3] For instance, determination of the structure of the ribosome—an enzyme that catalyzes peptide bond formation—revealed that its active site is composed entirely of RNA.^[4]

Structure[link]

Watson-Crick base pairs in a siRNA (hydrogen atoms are not shown)

Each nucleotide in RNA contains a ribose sugar, with carbons numbered 1' through 5'. A base is attached to the 1' position, in general, adenine (A), cytosine (C), guanine (G), or uracil (U). Adenine and guanine are purines, cytosine, and uracil are pyrimidines. A phosphate group is attached to the 3' position of one ribose and the 5' position of the next. The phosphate groups have a negative charge each at physiological pH, making RNA a charged molecule (polyanion). The bases may form hydrogen bonds between cytosine and guanine, between adenine and uracil and between guanine and uracil.^[5] However, other interactions are possible, such as a group of adenine bases binding to each other in a bulge,^[6] or the GNRA tetraloop that has a guanine–adenine base-pair.^[5]

Chemical structure of RNA

An important structural feature of RNA that distinguishes it from DNA is the presence of a hydroxyl group at the 2' position of the ribose sugar. The presence of this functional group causes the helix to adopt the A-form geometry rather than the B-form most commonly observed in DNA.^[7] This results in a very deep and narrow major groove and a shallow and wide minor groove.^[8] A second consequence of the presence of the 2'-hydroxyl group is that in conformationally flexible regions of an RNA molecule (that is, not involved in formation of a double helix), it can chemically attack the adjacent phosphodiester bond to cleave the backbone.^[9]

Secondary structure of a telomerase RNA.

RNA is transcribed with only four bases (adenine, cytosine, guanine and uracil),^[10] but these bases and attached sugars can be modified in numerous ways as the RNAs mature. Pseudouridine (Ψ), in which the linkage between uracil and ribose is changed from a C–N bond to a C–C bond, and ribothymidine (T) are found in various places (the most notable ones being in the TΨC loop of tRNA).^[11] Another notable modified base is hypoxanthine, a deaminated adenine base whose nucleoside is called inosine (I). Inosine plays a key role in the wobble hypothesis of the genetic code.^[12]

There are nearly 100 other naturally occurring modified nucleosides,^[13] of which pseudouridine and nucleosides with 2'-O-methylribose are the most common.^[14] The specific roles of many of these modifications in RNA are not fully understood. However, it is notable that, in ribosomal RNA, many of the post-transcriptional modifications occur in highly functional regions, such as the peptidyl transferase center and the subunit interface, implying that they are important for normal function.^[15]

The functional form of single-stranded RNA molecules, just like proteins, frequently requires a specific tertiary structure. The scaffold for this structure is provided by secondary structural elements that are hydrogen bonds within the molecule. This leads to several recognizable "domains" of secondary structure like hairpin loops, bulges, and internal loops.^[16] Since RNA is charged, metal ions such as Mg²⁺ are needed to stabilise many secondary and tertiary structures.^[17]

Synthesis[link]

Synthesis of RNA is usually catalyzed by an enzyme—RNA polymerase—using DNA as a template, a process known as transcription. Initiation of transcription begins with the binding of the enzyme to a promoter sequence in the DNA (usually found "upstream" of a gene). The DNA double helix is unwound by the helicase activity of the enzyme. The enzyme then progresses along the template strand in the 3’ to 5’ direction, synthesizing a complementary RNA molecule with elongation occurring in the 5’ to 3’ direction. The DNA sequence also dictates where termination of RNA synthesis will occur.^[18]

RNAs are often modified by enzymes after transcription. For example, a poly(A) tail and a 5' cap are added to eukaryotic pre-mRNA and introns are removed by the spliceosome.

There are also a number of RNA-dependent RNA polymerases that use RNA as their template for synthesis of a new strand of RNA. For instance, a number of RNA viruses (such as poliovirus) use this type of enzyme to replicate their genetic material.^[19] Also, RNA-dependent RNA polymerase is part of the RNA interference pathway in many organisms.^[20]

Types of RNA[link]

Overview[link]

Structure of a hammerhead ribozyme, a ribozyme that cuts RNA

Messenger RNA (mRNA) is the RNA that carries information from DNA to the ribosome, the sites of protein synthesis (translation) in the cell. The coding sequence of the mRNA determines the amino acid sequence in the protein that is produced.^[21] Many RNAs do not code for protein however (about 97% of the transcriptional output is non-protein-coding in eukaryotes ^{[22][23][24][25]}).

These so-called non-coding RNAs ("ncRNA") can be encoded by their own genes (RNA genes), but can also derive from mRNA introns.^[26] The most prominent examples of non-coding RNAs are transfer RNA (tRNA) and ribosomal RNA (rRNA), both of which are involved in the process of translation.^[1] There are also non-coding RNAs involved in gene regulation, RNA processing and other roles. Certain RNAs are able to catalyse chemical reactions such as cutting and ligating other RNA molecules,^[27] and the catalysis of peptide bond formation in the ribosome;^[4] these are known as ribozymes.

In translation[link]

Messenger RNA (mRNA) carries information about a protein sequence to the ribosomes, the protein synthesis factories in the cell. It is coded so that every three nucleotides (a codon) correspond to one amino acid. In eukaryotic cells, once precursor mRNA (pre-mRNA) has been transcribed from DNA, it is processed to mature mRNA. This removes its introns—non-coding sections of the pre-mRNA. The mRNA is then exported from the nucleus to the cytoplasm, where it is bound to ribosomes and translated into its corresponding protein form with the help of tRNA. In prokaryotic cells, which do not have nucleus and cytoplasm compartments, mRNA can bind to ribosomes while it is being transcribed from DNA. After a certain amount of time the message degrades into its component nucleotides with the assistance of ribonucleases.^[21]

Transfer RNA (tRNA) is a small RNA chain of about 80 nucleotides that transfers a specific amino acid to a growing polypeptide chain at the ribosomal site of protein synthesis during translation. It has sites for amino acid attachment and an anticodon region for codon recognition that binds to a specific sequence on the messenger RNA chain through hydrogen bonding.^[26]

Ribosomal RNA (rRNA) is the catalytic component of the ribosomes. Eukaryotic ribosomes contain four different rRNA molecules: 18S, 5.8S, 28S and 5S rRNA. Three of the rRNA molecules are synthesized in the nucleolus, and one is synthesized elsewhere. In the cytoplasm, ribosomal RNA and protein combine to form a nucleoprotein called a ribosome. The ribosome binds mRNA and carries out protein synthesis. Several ribosomes may be attached to a single mRNA at any time.^[21] Nearly all the RNA found in a typical eukaryotic cell is rRNA.

Transfer-messenger RNA (tmRNA) is found in many bacteria and plastids. It tags proteins encoded by mRNAs that lack stop codons for degradation and prevents the ribosome from stalling.^[28]

Regulatory RNAs[link]

Several types of RNA can downregulate gene expression by being complementary to a part of an mRNA or a gene's DNA. MicroRNAs (miRNA; 21-22 nt) are found in eukaryotes and act through RNA interference (RNAi), where an effector complex of miRNA and enzymes can cleave complementary mRNA, block the mRNA from being translated, or accelerate its degradation.^[29][30] While small interfering RNAs (siRNA; 20-25 nt) are often produced by breakdown of viral RNA, there are also endogenous sources of siRNAs.^[31][32]

siRNAs act through RNA interference in a fashion similar to miRNAs. Some miRNAs and siRNAs can cause genes they target to be methylated, thereby decreasing or increasing transcription of those genes.^[33][34][35] Animals have Piwi-interacting RNAs (piRNA; 29-30 nt) that are active in germline cells and are thought to be a defense against transposons and play a role in gametogenesis.^[36][37]

Many prokaryotes have CRISPR RNAs, a regulatory system similar to RNA interference.^[38] Antisense RNAs are widespread; most downregulate a gene, but a few are activators of transcription.^[39] One way antisense RNA can act is by binding to an mRNA, forming double-stranded RNA that is enzymatically degraded.^[40] There are many long noncoding RNAs that regulate genes in eukaryotes,^[41] one such RNA is Xist, which coats one X chromosome in female mammals and inactivates it.^[42]

An mRNA may contain regulatory elements itself, such as riboswitches, in the 5' untranslated region or 3' untranslated region; these cis-regulatory elements regulate the activity of that mRNA.^[43] The untranslated regions can also contain elements that regulate other genes.^[44]

In RNA processing[link]

Uridine to pseudouridine is a common RNA modification.

Many RNAs are involved in modifying other RNAs. Introns are spliced out of pre-mRNA by spliceosomes, which contain several small nuclear RNAs (snRNA),^[1] or the introns can be ribozymes that are spliced by themselves.^[45] RNA can also be altered by having its nucleotides modified to other nucleotides than A, C, G and U. In eukaryotes, modifications of RNA nucleotides are in general directed by small nucleolar RNAs (snoRNA; 60-300 nt),^[26] found in the nucleolus and cajal bodies. snoRNAs associate with enzymes and guide them to a spot on an RNA by basepairing to that RNA. These enzymes then perform the nucleotide modification. rRNAs and tRNAs are extensively modified, but snRNAs and mRNAs can also be the target of base modification.^[46][47] RNA can also be methylated.^[48][49]

RNA genomes[link]

Like DNA, RNA can carry genetic information. RNA viruses have genomes composed of RNA that encodes a number of proteins. The viral genome is replicated by some of those proteins, while other proteins protect the genome as the virus particle moves to a new host cell. Viroids are another group of pathogens, but they consist only of RNA, do not encode any protein and are replicated by a host plant cell's polymerase.^[50]

In reverse transcription[link]

Reverse transcribing viruses replicate their genomes by reverse transcribing DNA copies from their RNA; these DNA copies are then transcribed to new RNA. Retrotransposons also spread by copying DNA and RNA from one another,^[51] and telomerase contains an RNA that is used as template for building the ends of eukaryotic chromosomes.^[52]

Double-stranded RNA[link]

Double-stranded RNA (dsRNA) is RNA with two complementary strands, similar to the DNA found in all cells. dsRNA forms the genetic material of some viruses (double-stranded RNA viruses). Double-stranded RNA such as viral RNA or siRNA can trigger RNA interference in eukaryotes, as well as interferon response in vertebrates.^[53][54][55]

Key discoveries in RNA biology[link]

Robert W. Holley, left, poses with his research team.

Research on RNA has led to many important biological discoveries and numerous Nobel Prizes. Nucleic acids were discovered in 1868 by Friedrich Miescher, who called the material 'nuclein' since it was found in the nucleus.^[56] It was later discovered that prokaryotic cells, which do not have a nucleus, also contain nucleic acids. The role of RNA in protein synthesis was suspected already in 1939.^[57] Severo Ochoa won the 1959 Nobel Prize in Medicine (shared with Arthur Kornberg) after he discovered an enzyme that can synthesize RNA in the laboratory.^[58] However, the enzyme discovered by Ochoa (polynucleotide phosphorylase) was later shown to be responsible for RNA degradation, not RNA synthesis.

The sequence of the 77 nucleotides of a yeast tRNA was found by Robert W. Holley in 1965,^[59] winning Holley the 1968 Nobel Prize in Medicine (shared with Har Gobind Khorana and Marshall Nirenberg). In 1967, Carl Woese hypothesized that RNA might be catalytic and suggested that the earliest forms of life (self-replicating molecules) could have relied on RNA both to carry genetic information and to catalyze biochemical reactions—an RNA world.^[60][61]

During the early 1970s, retroviruses and reverse transcriptase were discovered, showing for the first time that enzymes could copy RNA into DNA (the opposite of the usual route for transmission of genetic information). For this work, David Baltimore, Renato Dulbecco and Howard Temin were awarded a Nobel Prize in 1975. In 1976, Walter Fiers and his team determined the first complete nucleotide sequence of an RNA virus genome, that of bacteriophage MS2.^[62]

In 1977, introns and RNA splicing were discovered in both mammalian viruses and in cellular genes, resulting in a 1993 Nobel to Philip Sharp and Richard Roberts. Catalytic RNA molecules (ribozymes) were discovered in the early 1980s, leading to a 1989 Nobel award to Thomas Cech and Sidney Altman. In 1990 it was found in petunia that introduced genes can silence similar genes of the plant's own, now known to be a result of RNA interference.^[63][64]

At about the same time, 22 nt long RNAs, now called microRNAs, were found to have a role in the development of C. elegans.^[65] Studies on RNA interference gleaned a Nobel Prize for Andrew Fire and Craig Mello in 2006, and another Nobel was awarded for studies on transcription of RNA to Roger Kornberg in the same year. The discovery of gene regulatory RNAs has led to attempts to develop drugs made of RNA, such as siRNA, to silence genes.^[66]

See also[link]

References[link]

External links[link]

• RNA World website Link collection (structures, sequences, tools, journals)
• Nucleic Acid Database Images of DNA, RNA and complexes.
• EteRNA a game forming RNA by pairing bases.

This biographical article needs additional citations for verification. Please help by adding reliable sources. Contentious material about living persons that is unsourced or poorly sourced must be removed immediately, especially if potentially libelous or harmful. (October 2009)

Sidney Altman
Born	(1939-05-07) 7 May 1939 (age 73)[1] Montreal, Quebec, Canada
Nationality	Canada & American (since 1984)
Fields	Molecular biology
Alma mater	MIT, University of Colorado at Boulder
Known for	Ribozymes
Notable awards	Nobel Prize in Chemistry (1989)

Sidney Altman (born May 7, 1939) is a Canadian American^[2] molecular biologist, who is currently the Sterling Professor of Molecular, Cellular, and Developmental Biology and Chemistry at Yale University. In 1989 he shared the Nobel Prize in Chemistry with Thomas R. Cech for their work on the catalytic properties of RNA.

Family and Education[link]

Altman was born on May 7, 1939 in Montreal, Quebec, Canada. His parents were immigrants to Canada, each coming from Eastern Europe as a young adult, in the 1920s. Altman's mother was from Bialystok in Poland, and had come to Canada with her sister at the age of eighteen, learning English and working in a textile factory to earn money to bring the rest of their family to Quebec. Altman's father, born in Ukraine, had been a worker on a collective farm in the Soviet Union. He was originally sponsored to come to Canada as a farm worker, but later, as a husband and a father of two sons, he supported the family by running a small grocery store in Montreal.^[2] Sidney Altman was later to look back on his parents' lives as an illustration of the value of the work ethic: "It was from them I learned that hard work in stable surroundings could yield rewards, even if only in infinitesimally small increments."^[3]

As Altman reached adulthood, the family's financial situation had become secure enough that he was able to pursue a college education. He went to the United States to study physics at the Massachusetts Institute of Technology. While at MIT, he was a member of the ice hockey team.^[2] After achieving his bachelor's degree from MIT in 1960, Altman spent 18 months as a graduate student in physics at Columbia University. Due to personal concerns and the lack of opportunity for beginning graduate students to participate in laboratory work, he left the program without completing the degree.^[2] Some months later, he enrolled as a graduate student in biophysics at the University of Colorado Medical Center. His project was a study of the effects of acridines on the replication of bacteriophage T4 DNA. He received his Ph.D. in biophysics from the University of Colorado in 1967.

Altman was married to Ann Korner in 1972. They are the parents of two children, Daniel and Leah.^[3] Having lived primarily in the United States since departing Montreal to attend MIT in 1958, Altman became a U.S. citizen in 1984, maintaining dual citizenship as a Canadian citizen as well.^[2][4]

Career[link]

After receiving his Ph.D., Altman embarked upon the first of two research fellowships. He joined Matthew Meselson's laboratory at Harvard University to study a DNA endonuclease involved in the replication and recombination of T4 DNA. Later, at the MRC Laboratory of Molecular Biology in Cambridge, England, Altman started the work that led to the discovery of RNase P and the enzymatic properties of the RNA subunit of that enzyme. John D. Smith, as well as several post-doctoral colleagues, provided Altman with very good advice that enabled him to test his ideas. "The discovery of the first radiochemically pure precursor to a tRNA molecule enabled me to get a job as an assistant professor at Yale University in 1971, a difficult time to get any job at all."^[1]

Altman's career at Yale followed a standard academic pattern with promotion through the ranks until he became Professor in 1980. He was Chairman of his department from 1983–1985 and in 1985 became the Dean of Yale College for four years. On July 1, 1989 he returned to the post of Professor on a full-time basis.

While at Yale, Altman's Nobel Prize work came with the analysis of the catalytic properties of the ribozyme RNase P. RNase P is a ribonucleoprotein particle consisting of both a structural RNA molecule and one (in prokaryotes) or more (in eukaryotes) proteins. Originally, it was believed that, in the bacterial RNase P complex, the protein subunit was responsible for the catalytic activity of the complex, which is involved in the maturation of tRNAs. During experiments in which the complex was reconstituted in test tubes, Altman and his group discovered that the RNA component, in isolation, was sufficient for the observed catalytic activity of the enzyme, indicating that the RNA itself had catalytic properties, which was the discovery that earned him the Nobel prize.^[4] Although the RNase P complex also exists in eukaryotic organisms, his later work revealed that in those organisms, the protein subunits of the complex are essential to the catalytic activity, in contrast to the bacterial RNase P.

Recognition[link]

Altman was elected a Fellow of the American Academy of Arts and Sciences in 1988.^[5]

Bibliography[link]

• Altman, Sidney (2007). "A view of RNase P.". Mol Biosyst 3 (9): 604–7. 2007 Sep. DOI:10.1039/b707850c. PMID 17700860. 

• Altman, S; Baer, M F; Bartkiewicz, M; Gold, H; Guerrier-Takada, C; Kirsebom, LA; Lumelsky, N; Peck, K (1989). "Catalysis by the RNA subunit of RNase P—a minireview.". Gene 82 (1): 63–4. 1989 Oct 15. DOI:10.1016/0378-1119(89)90030-9. PMID 2479591. 

See also[link]

• History of RNA biology
• List of RNA biologists

References[link]

External links[link]

• Sidney Altman Nobel Lecture: Enzymatic Cleavage of RNA by RNA
• Altman S Author Profile Page on Pubget

Sir David Charles Baulcombe
Born	1952 (age 59–60) Solihull, West Midlands
Residence	Norwich, Norfolk
Nationality	British
Fields	Plant Scientist and Geneticist
Institutions	Sainsbury Laboratory University of East Anglia University of Cambridge
Alma mater	University of Leeds University of Edinburgh
Known for	small interfering RNA
Notable awards	Royal Medal 2006, Knight Bachelor 2009, Harvey Prize 2009

Sir David Charles Baulcombe, FRS (born 1952) is a British plant scientist and geneticist. He is currently Royal Society Research Professor and Regius Professor of Botany at the University of Cambridge.

Biography[link]

David Baulcombe was born in Solihull, West Midlands (then Warwickshire). He received his B.Sc. degree in botany from the University of Leeds in 1973 at the age of 21. He proceeded to the University of Edinburgh, where he received his Ph.D. in botany in 1977. Baulcombe then spent the following three years as a post-doctoral fellow in North America, first at McGill University (Montreal, Canada) from January 1977 to November 1978, and then at the University of Georgia (Athens, Georgia, USA) until December 1980. Dr. Baulcombe returned to the United Kingdom then, where he joined the Plant Breeding Institute (PBI) in Cambridge and started his career as an independent scientist. At the PBI, Dr. Baulcombe initially held the position of Higher Scientific Officer, and was promoted to Principal Scientific Officer in April 1986.^[1] In August 1988 Dr. Baulcombe left Cambridge for Norwich. He joined the Sainsbury Laboratory as a Senior Research Scientist,^[2] and also served as Head of Laboratory between 1990–1993 and 1999-2003. In 1998 he was appointed Honorary Professor at the University of East Anglia, and given a full professorship there in 2002.^[1] In March 2007 it was announced that Prof. Baulcombe would become the next Professor of Botany at Cambridge University as a Royal Society Research Professor, taking up his post in September 2007.^[3] He serves on several committees and study sections,^[4] was elected Member of the European Molecular Biology Organisation in 1997 and was president of the International Society of Plant Molecular Biology 2003-2004. As of 2007^[update], he is also a senior advisor for The Embo Journal.^[5]

In June 2009, Baulcombe was awarded a knighthood by Queen Elizabeth II.^[6] Sir David resides in Norwich. He is married and has four children. His interests include music, sailing and hill walking.^[1]

Contributions to science[link]

Sir David Baulcombe's research interests and contributions to science are mainly in the fields of virus movement, genetic regulation, disease resistance, and gene silencing.

With Andrew Hamilton he discovered the small interfering RNA that is the specificity determinant in RNA-mediated gene silencing.^[7] Baulcombe's group demonstrated that while viruses can induce gene silencing, some viruses encode proteins that suppress gene silencing.^[4] After these initial observations in plants, many laboratories around the world searched for the occurrence of this phenomenon in other organisms. In 1998 Craig Mello and Andrew Fire reported a potent gene silencing effect after injecting double stranded RNA into C. elegans.^[8] This discovery was particularly notable because it represented the first identification of the causative agent for the phenomenon. Fire and Mello were awarded the Nobel Prize in Physiology or Medicine in 2006 for their work. ^[9]

With other members of his research group at the Sainsbury Laboratory, Baulcombe also helped unravel the importance of small interfering RNA in epigenetics and in defence against viruses.

Honours and awards[link]

Sir David Baulcombe has received the following honours and awards:

• 2001 elected Fellow of the Royal Society (FRS)
• 2002 elected Member of the Academia Europaea
• 2002 recipient of the Ruth Allen Award, awarded by the American Phytopathological Society
• 2002 recipient of the Kumho Science International Award in Plant Molecular Biology and Biotechnology, awarded by the Kumho Cultural Foundation, Korea
• 2003 co-recipient (with Craig Mello, Andrew Fire and Thomas Tuschl) of the Wiley Prize in the Biomedical Sciences, awarded by Rockefeller University
• 2004 recipient of the M. W. Beijerinck Virology Prize, awarded by the Royal Netherlands Academy of Arts and Sciences
• 2005 elected Foreign Associate Member of the National Academy of Sciences
• 2005 co-recipient (with Craig Mello and Andrew Fire) of the Massry Prize, awarded by the Massry Foundation and the University of Southern California
• 2006 recipient of the Royal Society's Royal Medal
• 2008 co-recipient of the Benjamin Franklin Medal in Life Science, awarded by the Franklin Institute [1]
• 2008 co-recipient (with Victor Ambros and Gary Ruvkun) of the Albert Lasker Award for Basic Medical Research
• 2008 appointed Fellow of Trinity College, Cambridge
• 2009 knighted by Queen Elizabeth II in the 2009 Birthday Honours List for services to plant science.
• 2009 recipient of the Harvey Prize, granted by the Technion Israeli Institute for Technology.
• 2010 recipient of the Wolf Prize in Agriculture.

Publications[link]

• Hamilton, A.; Voinnet, O.; Chappell, L.; Baulcombe, D. (2002). "Two classes of short interfering RNA in RNA silencing". The EMBO Journal 21 (17): 4671–4679. PMC 125409. PMID 12198169. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=125409.  edit
• Papaefthimiou, I.; Hamilton, A.; Denti, M.; Baulcombe, D.; Tsagris, M.; Tabler, M. (2001). "Replicating potato spindle tuber viroid RNA is accompanied by short RNA fragments that are characteristic of post-transcriptional gene silencing". Nucleic Acids Research 29 (11): 2395–2400. DOI:10.1093/nar/29.11.2395. PMC 55696. PMID 11376158. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=55696.  edit
• Dalmay, T.; Hamilton, A.; Rudd, S.; Angell, S.; Baulcombe, D. (2000). "An RNA-dependent RNA polymerase gene in Arabidopsis is required for posttranscriptional gene silencing mediated by a transgene but not by a virus". Cell 101 (5): 543–553. DOI:10.1016/S0092-8674(00)80864-8. PMID 10850496.  edit
• Burton, R.; Gibeaut, D.; Bacic, A.; Findlay, K.; Roberts, K.; Hamilton, A.; Baulcombe, D.; Fincher, G. (2000). "Virus-induced silencing of a plant cellulose synthase gene". The Plant cell 12 (5): 691–706. PMC 139921. PMID 10810144. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=139921.  edit
• Dalmay, T.; Hamilton, A.; Mueller, E.; Baulcombe, D. (2000). "Potato virus X amplicons in arabidopsis mediate genetic and epigenetic gene silencing". The Plant cell 12 (3): 369–379. PMC 139837. PMID 10715323. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=139837.  edit
• Jones, L.; Hamilton, A.; Voinnet, O.; Thomas, C.; Maule, A.; Baulcombe, D. (1999). "RNA-DNA interactions and DNA methylation in post-transcriptional gene silencing". The Plant cell 11 (12): 2291–2301. PMC 144133. PMID 10590159. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=144133.  edit
• Hamilton, A. J.; Baulcombe, D. C. (1999). "A species of small antisense RNA in posttranscriptional gene silencing in plants". Science 286 (5441): 950–952. DOI:10.1126/science.286.5441.950. PMID 10542148.  edit
• Hamilton, W.; Boccara, M.; Robinson, D.; Baulcombe, D. (1987). "The complete nucleotide sequence of tobacco rattle virus RNA-1". The Journal of general virology 68 (10): 2563–2575. PMID 3668507.  edit
• Boccara, M.; Hamilton, W.; Baulcombe, D. (1986). "The organisation and interviral homologies of genes at the 3' end of tobacco rattle virus RNA1". The EMBO Journal 5 (2): 223–229. PMC 1166722. PMID 16453668. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1166722.  edit

References[link]

External links[link]

• Interview with Professor Baulcombe from in-cites website
• Biography from the American Phytopathological Society 2002 Awards (p.4)