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Nucleotide
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This nucleotide contains the five-carbon sugar deoxyribose (at center), a nitrogenous base called adenine (upper right), and one phosphate group (left). The deoxyribose sugar joined only to the nitrogenous base forms a Deoxyribonucleoside called deoxyadenosine, whereas the whole structure along with the phosphate group is a nucleotide, a constituent of DNA with the name deoxyadenosine monophosphate.
Nucleotides are organic molecules that serve as the monomer units for forming the nucleic acid polymers deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecules in all life-forms on Earth. Nucleotides are the building blocks of nucleic acids; they are composed of three subunit molecules: a nitrogenous base, a five-carbon sugar (ribose or deoxyribose), and at least one phosphate group. They are also known as phosphate nucleotides.
A nucleoside is a nitrogenous base and a 5-carbon sugar. Thus a nucleoside plus a phosphate group yields a nucleotide.
Nucleotides also play a central role in life-form metabolism at the fundamental, cellular level. They carry packets of chemical energy—in the form of the nucleoside triphosphates ATP, GTP, CTP and UTP—throughout the cell to the many cellular functions that demand energy, which include synthesizing amino acids, proteins and cell membranes and parts; moving the cell and moving cell parts, both internally and intercellularly; dividing the cell, etc.[1] In addition, nucleotides participate in cell signaling (cGMP and cAMP), and are incorporated into important cofactors of enzymatic reactions (e.g. coenzyme A, FAD, FMN, NAD, and NADP+).
In experimental biochemistry, nucleotides can be radiolabeled with radionuclides to yield radionucleotides.
Contents
[hide]
1 Structure
2 Synthesis
2.1 Pyrimidine ribonucleotide synthesis
2.2 Purine ribonucleotide synthesis
2.3 Pyrimidine and purine degradation
3 Unnatural base pair (UBP)
4 Length unit
5 Nucleotide supplements
6 Abbreviation codes for degenerate bases
7 See also
8 References
9 Further reading
10 External links
Structure[edit]
Showing the arrangement of nucleotides within the structure of nucleic acids: At lower left, a monophosphate nucleotide; its nitrogenous base represents one side of a base-pair. At upper right, four nucleotides form two base-pairs: thymine and adenine (connected by double hydrogen bonds) and guanine and cytosine (connected by triple hydrogen bonds). The individual nucleotide monomers are chain-joined at their sugar and phosphate molecules, forming two 'backbones' (a double helix) of a nucleic acid, shown at upper left.
A nucleotide is composed of three distinctive chemical sub-units: a five-carbon sugar molecule, a nitrogenous base—which two together are called a nucleoside—and one phosphate group. With all three joined, a nucleotide is also termed a "nucleoside monophosphate". The chemistry sources ACS Style Guide[2] and IUPAC Gold Book[3] prescribe that a nucleotide should contain only one phosphate group, but common usage in molecular biology textbooks often extends the definition to include molecules with two, or with three, phosphates.[1][4][5][6] Thus, the terms "nucleoside diphosphate" or "nucleoside triphosphate" may also indicate nucleotides.
Nucleotides contain either a purine or a pyrimidine base—i.e., the nitrogenous base molecule, also known as a nucleobase—and are termed ribonucleotides if the sugar is ribose, or deoxyribonucleotides if the sugar is deoxyribose. Individual phosphate molecules repetitively connect the sugar-ring molecules in two adjacent nucleotide monomers, thereby connecting the nucleotide monomers of a nucleic acid end-to-end into a long chain. These chain-joins of sugar and phosphate molecules create a 'backbone' strand for a single- or double helix. In any one strand, the chemical orientation (directionality) of the chain-joins runs from the 5'-end to the 3'-end (read: 5 prime-end to 3 prime-end)—referring to the five carbon sites on sugar molecules in adjacent nucleotides. In a double helix, the two strands are oriented in opposite directions, which permits base pairing and complementarity between the base-pairs, all which is essential for replicating or transcribing the encoded information found in DNA.
Unlike in nucleic acid nucleotides, singular cyclic nucleotides are formed when the phosphate group is bound twice to the same sugar molecule, i.e., at the corners of the sugar hydroxyl groups.[1] These individual nucleotides function in cell metabolism rather than the nucleic acid structures of long-chain molecules.
Nucleic acids then are polymeric macromolecules assembled from nucleotides, the monomer-units of nucleic acids. The purine bases adenine and guanine and pyrimidine base cytosine occur in both DNA and RNA, while the pyrimidine bases thymine (in DNA) and uracil (in RNA) in just one. Adenine forms a base pair with thymine with two hydrogen bonds, while guanine pairs with cytosine with three hydrogen bonds.
Structural elements of three nucleotides—where one-, two- or three-phosphates are attached to the nucleoside (in yellow, blue, green) at center: 1st, the nucleotide termed as a nucleoside monophosphate is formed by adding a phosphate group (in red); 2nd, adding a second phosphate group forms a nucleoside diphosphate; 3rd, adding a third phosphate group results in a nucleoside triphosphate. + The nitrogenous base (nucleobase) is indicated by "Base" and "glycosidic bond" (sugar bond). All five primary, or canonical, bases—the purines and pyrimidines—are sketched at right (in blue).
Synthesis[edit]
Nucleotides can be synthesized by a variety of means both in vitro and in vivo.
In vivo, nucleotides can be synthesized de novo or recycled through salvage pathways.[7] The components used in de novo nucleotide synthesis are derived from biosynthetic precursors of carbohydrate and amino acid metabolism, and from ammonia and carbon dioxide. The liver is the major organ of de novo synthesis of all four nucleotides. De novo synthesis of pyrimidines and purines follows two different pathways. Pyrimidines are synthesized first from aspartate and carbamoyl-phosphate in the cytoplasm to the common precursor ring structure orotic acid, onto which a phosphorylated ribosyl unit is covalently linked. Purines, however, are first synthesized from the sugar template onto which the ring synthesis occurs. For reference, the syntheses of the purine and pyrimidine nucleotides are carried out by several enzymes in the cytoplasm of the cell, not within a specific organelle. Nucleotides undergo breakdown such that useful parts can be reused in synthesis reactions to create new nucleotides.
In vitro, protecting groups may be used during laboratory production of nucleotides. A purified nucleoside is protected to create a phosphoramidite, which can then be used to obtain analogues not found in nature and/or to synthesize an oligonucleotide.
Pyrimidine ribonucleotide synthesis[edit]
The synthesis of UMP.
The color scheme is as follows: enzymes, coenzymes, substrate names, inorganic molecules
Main article: Pyrimidine metabolism
The synthesis of the pyrimidines CTP and UTP occurs in the cytoplasm and starts with the formation of carbamoyl phosphate from glutamine and CO2. Next, aspartate carbamoyltransferase catalyzes a condensation reaction between aspartate and carbamoyl phosphate to form carbamoyl aspartic acid, which is cyclized into 4,5-dihydroorotic acid by dihydroorotase. The latter is converted to orotate by dihydroorotate oxidase. The net reaction is:
(S)-Dihydroorotate + O2 → Orotate + H2O2
Orotate is covalently linked with a phosphorylated ribosyl unit. The covalent linkage between the ribose and pyrimidine occurs at position C1[8] of the ribose unit, which contains a pyrophosphate, and N1 of the pyrimidine ring. Orotate phosphoribosyltransferase (PRPP transferase) catalyzes the net reaction yielding orotidine monophosphate (OMP):
Orotate + 5-Phospho-α-D-ribose 1-diphosphate (PRPP) → Orotidine 5'-phosphate + Pyrophosphate
Orotidine 5'-monophosphate is decarboxylated by orotidine-5'-phosphate decarboxylase to form uridine monophosphate (UMP). PRPP transferase catalyzes both the ribosylation and decarboxylation reactions, forming UMP from orotic acid in the presence of PRPP. It is from UMP that other pyrimidine nucleotides are derived. UMP is phosphorylated by two kinases to uridine triphosphate (UTP) via two sequential reactions with ATP. First the diphosphate form UDP is produced, which in turn is phosphorylated to UTP. Both steps are fueled by ATP hydrolysis:
ATP + UMP → ADP + UDP
UDP + ATP → UTP + ADP
CTP is subsequently formed by amination of UTP by the catalytic activity of CTP synthetase. Glutamine is the NH3 donor and the reaction is fueled by ATP hydrolysis, too:
UTP + Glutamine + ATP + H2O → CTP + ADP + Pi
Cytidine monophosphate (CMP) is derived from cytidine triphosphate (CTP) with subsequent loss of two phosphates.[9] [10]
Purine ribonucleotide synthesis[edit]
Main article: Purine metabolism
The atoms that are used to build the purine nucleotides come from a variety of sources:
The synthesis of IMP. The color scheme is as follows: enzymes, coenzymes, substrate names, metal ions, inorganic molecules
The biosynthetic origins of purine ring atoms

N1 arises from the amine group of Asp
C2 and C8 originate from formate
N3 and N9 are contributed by the amide group of Gln
C4, C5 and N7 are derived from Gly
C6 comes from HCO3− (CO2)
The de novo synthesis of purine nucleotides by which these precursors are incorporated into the purine ring proceeds by a 10-step pathway to the branch-point intermediate IMP, the nucleotide of the base hypoxanthine. AMP and GMP are subsequently synthesized from this intermediate via separate, two-step pathways. Thus, purine moieties are initially formed as part of the ribonucleotides rather than as free bases.
Six enzymes take part in IMP synthesis. Three of them are multifunctional:
GART (reactions 2, 3, and 5)
PAICS (reactions 6, and 7)
ATIC (reactions 9, and 10)
The pathway starts with the formation of PRPP. PRPS1 is the enzyme that activates R5P, which is formed primarily by the pentose phosphate pathway, to PRPP by reacting it with ATP. The reaction is unusual in that a pyrophosphoryl group is directly transferred from ATP to C1 of R5P and that the product has the α configuration about C1. This reaction is also shared with the pathways for the synthesis of Trp, His, and the pyrimidine nucleotides. Being on a major metabolic crossroad and requiring much energy, this reaction is highly regulated.
In the first reaction unique to purine nucleotide biosynthesis, PPAT catalyzes the displacement of PRPP's pyrophosphate group (PPi) by an amide nitrogen donated from either glutamine (N), glycine (N&C), aspartate (N), folic acid (C1), or CO2. This is the committed step in purine synthesis. The reaction occurs with the inversion of configuration about ribose C1, thereby forming β-5-phosphorybosylamine (5-PRA) and establishing the anomeric form of the future nucleotide.
Next, a glycine is incorporated fueled by ATP hydrolysis and the carboxyl group forms an amine bond to the NH2 previously introduced. A one-carbon unit from folic acid coenzyme N10-formyl-THF is then added to the amino group of the substituted glycine followed by the closure of the imidazole ring. Next, a second NH2 group is transferred from a glutamine to the first carbon of the glycine unit. A carboxylation of the second carbon of the glycin unit is concomittantly added. This new carbon is modified by the additional of a third NH2 unit, this time transferred from an aspartate residue. Finally, a second one-carbon unit from formyl-THF is added to the nitrogen group and the ring covalently closed to form the common purine precursor inosine monophosphate (IMP).
Inosine monophosphate is converted to adenosine monophosphate in two steps. First, GTP hydrolysis fuels the addition of aspartate to IMP by adenylosuccinate synthase, substituting the carbonyl oxygen for a nitrogen and forming the intermediate adenylosuccinate. Fumarate is then cleaved off forming adenosine monophosphate. This step is catalyzed by adenylosuccinate lyase.
Inosine monophosphate is converted to guanosine monophosphate by the oxidation of IMP forming xanthylate, followed by the insertion of an amino group at C2. NAD+ is the electron acceptor in the oxidation reaction. The amide group transfer from glutamine is fueled by ATP hydrolysis.
Pyrimidine and purine degradation[edit]
In humans, pyrimidine rings (C, T, U) can be degraded completely to CO2 and NH3 (urea excretion). That having been said, purine rings (G, A) cannot. Instead they are degraded to the metabolically inert uric acid which is then excreted from the body. Uric acid is formed when GMP is split into the base guanine and ribose. Guanine is deaminated to xanthine which in turn is oxidized to uric acid. This last reaction is irreversible. Similarly, uric acid can be formed when AMP is deaminated to IMP from which the ribose unit is removed to form hypoxanthine. Hypoxanthine is oxidized to xanthine and finally to uric acid. Instead of uric acid secretion, guanine and IMP can be used for recycling purposes and nucleic acid synthesis in the presence of PRPP and aspartate (NH3 donor).
Unnatural base pair (UBP)[edit]
Main article: Base pair § Unnatural base pair (UBP)
An unnatural base pair (UBP) is a designed subunit (or nucleobase) of DNA which is created in a laboratory and does not occur in nature. In 2012, a group of American scientists led by Floyd Romesberg, a chemical biologist at the Scripps Research Institute in San Diego, California, published that his team designed an unnatural base pair (UBP).[11] The two new artificial nucleotides or Unnatural Base Pair (UBP) were named d5SICS and dNaM. More technically, these artificial nucleotides bearing hydrophobic nucleobases, feature two fused aromatic rings that form a (d5SICS–dNaM) complex or base pair in DNA.[12][13] In 2014 the same team from the Scripps Research Institute reported that they synthesized a stretch of circular DNA known as a plasmid containing natural T-A and C-G base pairs along with the best-performing UBP Romesberg's laboratory had designed, and inserted it into cells of the common bacterium E. coli that successfully replicated the unnatural base pairs through multiple generations.[14] This is the first known example of a living organism passing along an expanded genetic code to subsequent generations.[12][15] This was in part achieved by the addition of a supportive algal gene that expresses a nucleotide triphosphate transporter which efficiently imports the triphosphates of both d5SICSTP and dNaMTP into E. coli bacteria.[12] Then, the natural bacterial replication pathways use them to accurately replicate the plasmid containing d5SICS–dNaM.
The successful incorporation of a third base pair is a significant breakthrough toward the goal of greatly expanding the number of amino acids which can be encoded by DNA, from the existing 21 amino acids to a theoretically possible 172, thereby expanding the potential for living organisms to produce novel proteins.[14] The artificial strings of DNA do not encode for anything yet, but scientists speculate they could be designed to manufacture new proteins which could have industrial or pharmaceutical uses.[16]
Length unit[edit]
Nucleotide (abbreviated "nt") is a common unit of length for single-stranded nucleic acids, similar to how base pair is a unit of length for double-stranded nucleic acids.
Nucleotide supplements[edit]
A study done by the Department of Sports Science at the University of Hull in Hull, UK has shown that nucleotides have significant impact on cortisol levels in saliva. Post exercise, the experimental nucleotide group had lower cortisol levels in their blood than the control or the placebo. Additionally, post supplement values of Immunoglobulin A were significantly higher than either the placebo or the control. The study concluded, "nucleotide supplementation blunts the response of the hormones associated with physiological stress."[17]
Another study conducted in 2013 looked at the impact nucleotide supplementation had on the immune system in athletes. In the study, all athletes were male and were highly skilled in taekwondo. Out of the twenty athletes tested, half received a placebo and half received 480 mg per day of nucleotide supplement. After thirty days, the study concluded that nucleotide supplementation may counteract the impairment of the body's immune function after heavy exercise.[18]
Abbreviation codes for degenerate bases[edit]
Main article: Nucleic acid notation
The IUPAC has designated the symbols for nucleotides.[19] Apart from the five (A, G, C, T/U) bases, often degenerate bases are used especially for designing PCR primers. These nucleotide codes are listed here. Some primer sequences may also include the character "I", which codes for the non-standard nucleotide inosine. Inosine occurs in tRNAs, and will pair with adenine, cytosine, or thymine. This character does not appear in the following table however, because it does not represent a degeneracy. While inosine can serve a similar function as the degeneracy "D", it is an actual nucleotide, rather than a representation of a mix of nucleotides that covers each possible pairing needed.
Symbol[19]
Description
Bases represented
A adenine A 1
C cytosine C
G guanine G
T thymine T
U uracil U
W weak A T 2
S strong C G
M amino A C
K keto G T
R purine A G
Y pyrimidine C T
B not A (B comes after A) C G T 3
D not C (D comes after C) A G T
H not G (H comes after G) A C T
V not T (V comes after T and U) A C G
N any base (not a gap) A C G T 4
See also[edit]
Molecular and cellular biology portal
Biology
Chromosome
Gene
Genetics
Nucleic acid analogues
Nucleic acid sequence
Nucleobase
References[edit]
^ Jump up to: a b c Alberts B, Johnson A, Lewis J, Raff M, Roberts K & Walter P (2002). Molecular Biology of the Cell (4th ed.). Garland Science. ISBN 0-8153-3218-1. pp. 120–121.
Jump up ^ Coghill, Anne M.; Garson, Lorrin R., eds. (2006). The ACS style guide: effective communication of scientific information (3rd ed.). Washington, D.C.: American Chemical Society. p. 244. ISBN 978-0-8412-3999-9.
Jump up ^ "Nucleotides". IUPAC Gold Book. International Union of Pure and Applied Chemists. doi:10.1351/goldbook.N04255. Retrieved 30 June 2014.
Jump up ^ Lehninger, Albert L. (1975). Biochemistry: the molecular basis of cell structure and function. New York: Worth Publishers Inc. doi:10.1002/jobm.19770170116.
Jump up ^ Stryer, Lubert (1988). Biochemistry (3rd ed.). New York: W. H. Freeman. ISBN 9780716719205.
Jump up ^ Garrett, Reginald H.; Grisham, Charles M. (2007). Biochemistry (4th ed.). Belmont, California: Brooks/Cole, Cengage Learning.
Jump up ^ Zaharevitz, DW; Anerson, LW; Manlinowski, NM; Hyman, R; Strong, JM; Cysyk, RL. "Contribution of de-novo and salvage synthesis to the uracil nucleotide pool in mouse tissues and tumors in vivo".
Jump up ^ See IUPAC nomenclature of organic chemistry for details on carbon residue numbering
Jump up ^ Jones, M. E. (1980). "Pyrimidine nucleotide biosynthesis in animals: Genes, enzymes, and regulation of UMP biosynthesis". Annu. Rev. Biochem. 49 (1): 253–79. doi:10.1146/annurev.bi.49.070180.001345. PMID 6105839.
Jump up ^ McMurry, JE; Begley, TP (2005). The organic chemistry of biological pathways. Roberts & Company. ISBN 978-0-9747077-1-6.
Jump up ^ Malyshev, Denis A.; Dhami, Kirandeep; Quach, Henry T.; Lavergne, Thomas; Ordoukhanian, Phillip (24 July 2012). "Efficient and sequence-independent replication of DNA containing a third base pair establishes a functional six-letter genetic alphabet". Proceedings of the National Academy of Sciences of the United States of America. 109 (30): 12005–12010. Bibcode:2012PNAS..10912005M. doi:10.1073/pnas.1205176109. PMC 3409741  . PMID 22773812. Retrieved 2014-05-11.
^ Jump up to: a b c Malyshev, Denis A.; Dhami, Kirandeep; Lavergne, Thomas; Chen, Tingjian; Dai, Nan; Foster, Jeremy M.; Corrêa, Ivan R.; Romesberg, Floyd E. (May 7, 2014). "A semi-synthetic organism with an expanded genetic alphabet". Nature. 509: 385–8. doi:10.1038/nature13314. PMC 4058825  . PMID 24805238. Retrieved May 7, 2014.
Jump up ^ Callaway, Ewan (May 7, 2014). "Scientists Create First Living Organism With 'Artificial' DNA". Nature News. Huffington Post. Retrieved 8 May 2014.
^ Jump up to: a b Fikes, Bradley J. (May 8, 2014). "Life engineered with expanded genetic code". San Diego Union Tribune. Retrieved 8 May 2014.
Jump up ^ Sample, Ian (May 7, 2014). "First life forms to pass on artificial DNA engineered by US scientists". The Guardian. Retrieved 8 May 2014.
Jump up ^ Pollack, Andrew (May 7, 2014). "Scientists Add Letters to DNA's Alphabet, Raising Hope and Fear". New York Times. Retrieved 8 May 2014.
Jump up ^ Mc Naughton, L.; Bentley, D.; Koeppel, P. (2007-03-01). "The effects of a nucleotide supplement on the immune and metabolic response to short term, high intensity exercise performance in trained male subjects". The Journal of Sports Medicine and Physical Fitness. 47 (1): 112–118. ISSN 0022-4707. PMID 17369807.
Jump up ^ Riera, Joan; Pons, Victoria; Martinez-Puig, Daniel; Chetrit, Carlos; Tur, Josep A.; Pons, Antoni; Drobnic, Franchek (2013-04-08). "Dietary nucleotide improves markers of immune response to strenuous exercise under a cold environment". Journal of the International Society of Sports Nutrition. 10 (1): 20. doi:10.1186/1550-2783-10-20. PMC 3626726  . PMID 23566489.
^ Jump up to: a b Nomenclature Committee of the International Union of Biochemistry (NC-IUB) (1984). "Nomenclature for Incompletely Specified Bases in Nucleic Acid Sequences". Retrieved 2008-02-04.
Further reading[edit]
Sigel, Astrid; Operschall, Bert P.; Sigel, Helmut (2017). "Chapter 11. Complex Formation of Lead(II) with Nucleotides and Their Constituents". In Astrid, S.; Helmut, S.; Sigel, R. K. O. Lead: Its Effects on Environment and Health. Metal Ions in Life Sciences. 17. de Gruyter. pp. 319–402. doi:10.1515/9783110434330-011.
External links[edit]
Abbreviations and Symbols for Nucleic Acids, Polynucleotides and their Constituents (IUPAC)
Provisional Recommendations 2004 (IUPAC)
Chemistry explanation of nucleotide structure
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Nucleic acid constituents
Nucleobase
Purine
Adenine
Guanine
Hypoxanthine
Xanthine
Purine analogue
Pyrimidine
Uracil
Thymine
Cytosine
Pyrimidine analogue
Unnatural base pair (UBP)
Nucleoside
Ribonucleoside
Adenosine
Guanosine
5-Methyluridine
Uridine
5-Methylcytidine
Cytidine
Inosine
Xanthosine
Wybutosine
Deoxyribonucleoside
Deoxyadenosine
Deoxyguanosine
Thymidine
Deoxyuridine
Deoxycytidine
Deoxyinosine
Deoxyxanthosine
Nucleotide
(Nucleoside monophosphate)
Ribonucleotide
AMP
GMP
m5UMP
UMP
CMP
IMP
XMP
Deoxyribonucleotide
dAMP
dGMP
dTMP
dUMP
dCMP
dIMP
dXMP
Cyclic nucleotide
cAMP
cGMP
c-di-GMP
c-di-AMP
cADPR
Nucleoside diphosphate
ADP
GDP
m5UDP
UDP
CDP
dADP
dGDP
dTDP
dUDP
dCDP
Nucleoside triphosphate
ATP
GTP
m5UTP
UTP
CTP
ITP
XTP
dATP
dGTP
dTTP
dUTP
dCTP
dITP
dXTP
[show]
v
t
e
Purine receptor modulators
Receptor
(ligands)
P0 (adenine)
Agonists: 8-Aminoadenine
Adenine
P1
(adenosine)
Agonists: 2-(1-Hexynyl)-N-methyladenosine
2-Cl-IB-MECA
2'-MeCCPA
4'-O-β-D-Glucosyl-9-O-(6''-deoxysaccharosyl)olivil
5'-N-ethylcarboxamidoadenosine
Adenosine
ADP
AMP
Apadenoson
ATL-146e
ATP
BAY 60–6583
Binodenoson
Capadenoson
CCPA
CGS-21680
CP-532,903
Evodenoson
GR 79236
LUF-5835
LUF-5845
N6-Cyclopentyladenosine
Namodenoson
Neladenoson dalanate
Piclidenoson
Regadenoson
SDZ WAG 994
Selodenoson
Sonedenoson
Tecadenoson
UK-432,097
Antagonists: 8-Chlorotheophylline
8-Phenyl-1,3-dipropylxanthine
8-Phenyltheophylline
Acefylline
Aminophylline
ATL-444
Bamifylline
Cafedrine
Caffeine
Caffeine citrate
Cartazolate
CGH-2466
CGS-15943
Choline theophyllinate
CPX
CVT-6883
Dimethazan
DMPX
DPCPX
Dyphylline
Enprofylline
Etazolate
Fenethylline
IBMX
Isovaleric acid
Istradefylline
KF-26777
MRE3008F20
MRS-1220
MRS-1334
MRS-1706
MRS-1754
MRS-3777
Paraxanthine
Pentoxifylline
Preladenant
Propentofylline
Proxyphylline
PSB-10
PSB-11
PSB-36
PSB-603
PSB-788
PSB-1115
Reversine
Rolofylline
SCH-442,416
SCH-58261
Theacrine
Theobromine
Theodrenaline
Theophylline
Tozadenant
Tracazolate
VUF-5574
ZM-241,385
P2
(nucleotide)
P2X
(ATP)
Agonists: 2-Me-SATP
α,β-Me-ATP
Adenosine
ADP
AMP
Ap4A
Ap5A
ATP
ATPγS
BzATP
Cibacron blue
CTP
D-β,γ-Me-ATP
GTP
HT-AMP
Ivermectin
L-β,γ-Me-ATP
MRS-2219
PAPET-ATP
UTP
Zinc
Antagonists: 5-BDBD
A-317491
A-438079
A-740003
A-804598
A-839977
AF-353
AZ-10606120
AZ-11645373
BBG
Calcium
Calmidazolium
Chelerythrine
Copper
Emodin (Rheum officinale)
Evans Blue
GW-791343
HMA
Ip5I
isoPPADS
JNJ-47965567
KN-04
KN-62
Magnesium
MRS-2159
NF-023
NF-110
NF-157
NF-279
NF-449
Opiranserin (VVZ-149)
Oxidized-ATP
Phenol Red
Phenolphthalein
PPADS
PPNDS
PSB-12062
Puerarin (Radix puerariae)
Purotoxin 1
RB-2
Ro 0437626
Ro 51
RO-3
Sodium ferulate (Angelica sinensis, Ligusticum wallichii)
Suramin
TC-P 262
Tetramethylpyrazine (ligustrazine) (Ligusticum wallichii)
TNP-ATP
Zinc
P2Y
Agonists: 2-Me-SADP
2-Me-SATP
2-Thio-UTP
5-Br-UDP
5-OMe-UDP
α,β-Me-ATP
Adenosine
ADP
ADPβS
Ap3A
AR-C 67085MX
ATP
ATPγS
CTP
dATP
Denufosol
Diquafosol
IDP
ITP
INS-365
INS-37217
MRS-2365
MRS-2690
MRS-2693
MRS-2768
MRS-2957
MRS-4062
NF-546
PAPET-ATP
PSB-0474
PSB-1114
UDP
UDPβS
UDP-galactose
UDP-glucose
UDP-N-acetylglucosamine
Up3U
UTP
UTPγS
Antagonists: 2-Me-SAMP
A3P5PS
AMPαS
Ap4A
AR-C 66096
AR-C 67085MX
AR-C 69931MX
AR-C 118925XX
ATP
BzATP
C1330-7
Cangrelor
Clopidogrel
Elinogrel
Ip5I
MRS-2179
MRS-2211
MRS-2279
MRS-2395
MRS-2500
MRS-2578
NF-157
NF-340
PIT
PPADS
Prasugrel
PSB-0739
RB-2
Regrelor
Suramin
Ticagrelor
Ticlopidine
UDP
Transporter
(blockers)
CNTs
6-Hydroxy-7-methoxyflavone
Adenosine
dMeThPmR
Estradiol
KGO-2142
KGO-2173
MeThPmR
Phloridzin
Progesterone
ENTs
Barbiturates
Benzodiazepines
Cilostazol
Dilazep
Dipyridamole
Estradiol
Ethanol
Hexobendine
NBMPR
Pentoxifylline
Progesterone
Propentofylline
PMAT
Decynium-22
Enzyme
(inhibitors)
XO
Allopurinol
Amflutizole
Benzbromarone
Caffeic acid
Cinnamaldehyde
Cinnamomum osmophloeum
Febuxostat
Myo-inositol
Kaempferol
Myricetin
Niraxostat
Oxipurinol
Phytic acid
Pistacia integerrima
Propolis
Quercetin
Tisopurine
Topiroxostat
Others
Aminopterin
Azathioprine
Methotrexate
Mycophenolic acid
Pemetrexed
Pralatrexate
Many others
Others
Precursors: Adenine
Adenosine
AMP
ADP
ATP
Cytosine
Cytidine
CMP
CDP
CTP
Guanine
Guanosine
GMP
GDP
GTP
Hypoxanthine
Inosine
IMP
IDP
ITP
Ribose
Uracil
Uridine
UMP
UDP
UTP
Others: Chrysophanol (rhubarb)
See also: Receptor/signaling modulators
Authority control
NDL: 00568741
NKC: ph123363
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text:
University of Michigan School of Public Health
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University of Michigan School of Public Health
Type
Public
Established
1941
Dean
Martin Philbert
Location
Ann Arbor, Michigan, US
Campus
Suburban
Website
sph.umich.edu
Henry F. Vaughan School of Public Health
The University of Michigan School of Public Health is one of the professional graduate schools of the University of Michigan. Located in Ann Arbor, Michigan, UM SPH is one of the oldest schools of public health in the country and is also considered one of the top schools focusing on health in the United States. Founded in 1941, the School of Public Health grew out of the University of Michigan's degree programs in public health, some of which date back to the 19th century.
According to the US News & World Report's report on graduate programs, the University of Michigan School of Public Health was ranked as the #4 School of Public Health in the country and also had the #1 Healthcare Management program in the country in 2011.[1][2][3]
Contents
[hide]
1 Notable faculty
2 Notable alumni
3 References
4 External links
Notable faculty[edit]
Thomas Francis, Jr., mentor of Jonas Salk; ran field trials of oral polio vaccine that deemed it safe and effective
John R. Griffith, Andrew Pattullo Collegiate Professor in the Department of Health Management and Policy; co-author of The Well-Managed Health Care Organization
George Kaplan, social epidemiologist
Neal Krause, health behavior and health education expert
Paula Lantz, past Chair of the Department of Health Management and Policy
Jonas Salk, developer of the polio vaccine
Ken Warner, past Dean of the School of Public Health; worked successfully to implement a non-smoking University of Michigan campus on July 1, 2011
Marc A. Zimmerman, former chair Health Behavior & Health Education; Director of the Prevention Research Center and Michigan Youth Violence Prevention Center
Notable alumni[edit]
Shervin Assari, ranked 4 among 31,000 peer reviewers by Publons; NYAM Fellow, President of SAPHIR, Chair of Associate Members Committee, American College of Epidemiology, Chair of Membership Council, American Academy of Health Behavior (AAHB)
Larry Brilliant, physician, epidemiologist, technologist, author and the former director of Google's philanthropic arm Google.org
Julio Frenk, former Minister of Health in Mexico; former dean of Harvard School of Public Health
David Michael Green, MacArthur Foundation Fellow
Albert H. Wheeler, first African-American Mayor of Ann Arbor, Michigan
Mona Hanna-Attisha, pediatrician who was a key whistleblower revealing the Flint Water Crisis which exposed children to dangerous levels of lead in Flint, Michigan.
References[edit]
Jump up ^ US News & World Report - Education: Grad Schools: Public Health Ranking - USNews.com
Jump up ^ [1]
Jump up ^ US News & World Report - Breaking News, World News, Business News, and America's Best Colleges - USNews.com
External links[edit]
University of Michigan School of Public Health
[show]
v
t
e
University of Michigan
Located in: Ann Arbor, Michigan
Academics
Ross School of Business
School of Dentistry
School of Education
College of Engineering
Rackham School of Graduate Studies
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School of Public Health
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Hopwood Award
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Michigan Wolverines
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Founded: 1817
Students: 44,718
Endowment: 7.05 billion
Retrieved from "https://en.wikipedia.org/w/index.php?title=University_of_Michigan_School_of_Public_Health&oldid=801073687"
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This page was last edited on 17 September 2017, at 14:02.
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Walter Egan
From Wikipedia, the free encyclopedia
Jump to: navigation, search
For the golfer, see Walter Egan (golfer). For the politician, see Walter D. Egan.
This biography of a living person 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. (April 2017) (Learn how and when to remove this template message)
Walter Egan
Born
(1948-07-12) July 12, 1948 (age 69)
Queens, New York, United States
Genres
Soft rock, folk rock, rock
Occupation(s)
Singer, guitarist, songwriter
Instruments
Guitar, vocals
Years active
1969–present
Labels
Culture Factory, Razor & Tie, Columbia, Renaissance, Red Steel, Spectra
Walter Egan (born July 12, 1948) is an American rock musician, best known for his 1978 gold status hit single "Magnet and Steel" from his second album release, Not Shy, produced by Lindsey Buckingham and Richard Dashut.[1] The song reached #8 on the Billboard Hot 100 and #18 on the Easy Listening chart.[2] Overseas, it peaked at #32 on the Australian Singles Chart, Kent Music Report.[3]
Contents
[hide]
1 Early years and education
2 Career
2.1 Music
2.2 Game show appearances
3 Discography
4 References
5 External links
Early years and education[edit]
Egan was born in Queens, New York, United States. In 1970, as a sculpture major, he earned a Bachelor of Fine Arts degree at Georgetown University in Washington, D.C. as one of Georgetown's first art majors. Egan started his career in the arts, concentrating on printmaking and painting, and the music business. As early as 1969, Egan was playing guitar and composing songs. While an undergraduate, he was a lead guitarist and backup vocalist for the college-formed group Sageworth and Drums, also known as Sageworth.[4] The group evolved from a group of Georgetown University undergraduate students playing weekend mixers and parties to a tight, professional headlining act with a reputation as one of the best and most original bands in the greater Washington, D.C. area.[5] Other members included Peter Barry Chowka, lead vocal and guitar; John Zambetti, guitar and backing vocals; Tom Guidera, bass and backing vocals; and Matthew Sheppard, drums, with Annie McLoone added later as a lead singer.[6][5]
In late 1971, Sageworth relocated to Boston and spent the next two years performing around the Northeast before finally breaking up, at which point Walter moved to Los Angeles and began his career as a solo artist.
Career[edit]
Music[edit]
In 1977, Lindsey Buckingham of Fleetwood Mac co-produced Walter Egan's first album, Fundamental Roll, with Stevie Nicks, also of Fleetwood Mac.[1] The hit song, "Magnet and Steel", inspired by Nicks and from his second album "Not Shy",[7] was featured in the 1997 film Boogie Nights, the 1998 film Overnight Delivery, and the 1999 film Deuce Bigalow: Male Gigolo. Egan also wrote "Hearts on Fire", which was covered by Gram Parsons on his album Grievous Angel, and "Hot Summer Nights", which was the first hit for the band Night, which included such session musicians as Nicky Hopkins and Robbie McIntosh. Egan scored minor hits with his own version of "Hot Summer Nights", as well as "Only the Lucky" and "Fool Moon Fire".[8]
Egan later toured as a member of a latter-day version of Spirit.[9]
Egan is listed as co-writer on the Eminem hit "We Made You".[10] The song's producer, Dr. Dre, believed he was inspired by the bass line from (and used samples of)[11] Egan's "Hot Summer Nights".
In 2008, he participated in a music industry seminar (with Bill Danoff) at Georgetown University.[5]
In 2011, from his home in Franklin, he produced a new album entitled Raw Elegant that was released on Spectra Records.
Game show appearances[edit]
In 1985, Egan was a four-time champion on the game show Catchphrase. In 1986, Egan appeared as a contestant on the television game show Scrabble. During his introduction segment, Egan identified himself as a singer and songwriter. Host Chuck Woolery asked Egan to sing a bit from one of his songs, and he sang the main hook from "Magnet and Steel".
Discography[edit]
Albums
Fundamental Roll (Culture Factory, 1977)
Not Shy (Columbia, 1978)
HiFi (Columbia, 1979)
The Last Stroll (Columbia, 1980)
Wild Exhibitions (Renaissance, 1983)
Walternative (We, 1999)
The Lost Album (Renaissance, 2000)
Maddog (Red Steel, 2001)
Apocalypso Now (Gaff, 2002)
The Meaning of Live (Red Steel, 2004)
Raw Elegant (Spectra, 2011)
Myth America (Classic Music Vault, 2014)
Singles
"Only The Lucky"/"I'd Rather Have Fun" (Columbia 10531, 1977)
"When I Get My Wheels"/"Waitin'" (Columbia 10591, 1977)
"Magnet And Steel"/"Tunnel O' Love" (Columbia 10719, 1978)
"Hot Summer Nights"/"She's So Tough" (Columbia 10824, 1978)
"Magnet And Steel"/"Only The Lucky" (Columbia Hall of Fame 33353, 1979)
"Unloved"/"Make It Alone" (Columbia 10916, 1979)
"You're The One"/"Like You Do" (Columbia 11046, 1979)
"That's That"/"Hi Fi Love" (Columbia 11116, 1979)
"Baby Let's Runaway"/"Johnny Z (Is A Real Cool Guy)" (Columbia 11297, 1980)
"Fool Moon Fire"/"Tammy Ann" (Backstreet 52200, 1983)
"Star Of My Heart"/"Joyce" (Backstreet 52249, 1983)
"Magnet And Steel"/"Only The Lucky" (Collectables 33353)
References[edit]
^ Jump up to: a b "Not Shy - Walter Egan - Songs, Reviews, Credits". AllMusic. Retrieved 11 April 2017.
Jump up ^ Whitburn, Joel (2002). Top Adult Contemporary: 1961-2001. Record Research. p. 85.
Jump up ^ "One Hit Wonders". Australian-charts.com. October 22, 2008. Retrieved 2011-08-06.
Jump up ^ "Walter Egan - Biography & History". AllMusic. Retrieved 11 April 2017.
^ Jump up to: a b c Reger, Jeff (March 13, 2008). "Take Me Home". The Georgetown Voice. Retrieved May 6, 2017.
Jump up ^ "WalterEgan.com - Official Web Site of Walter Egan". Walteregan.com. Retrieved 11 April 2017.
Jump up ^ "The Penguin Q&A Sessions: Walter Egan". FleetwoodMac.net. October 13–26, 1999. Retrieved 2013-05-06.
Jump up ^ Whitburn, Joel (2000). Top Pop Singles 1955-1999. Menomonee Falls, WI: Record Research. pp. 201, 466. ISBN 978-0-89820-139-0.
Jump up ^ Adelson, Martin E. "Walter Egan Question and Answer Session". Fleetwoodmac.net. Retrieved 11 April 2017.
Jump up ^ "Eminem - Relapse". Discogs.com. Retrieved 11 April 2017.
Jump up ^ "Eminem - Relapse (CD, Album)". Discogs.com. Retrieved 2012-03-14.
External links[edit]
Walter Egan's MySpace page
Walternative - Walter Egan Fansite
- Gram Parsons Guitar Pull site
Authority control
WorldCat Identities
VIAF: 33668513
MusicBrainz: c0c71b1c-d524-44c1-bd20-04531270cc77
IATH: w6s58zdp
Retrieved from "https://en.wikipedia.org/w/index.php?title=Walter_Egan&oldid=807105771"
Categories:
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