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Genetic code
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"Codon" redirects here. For the plant genus, see Codon (genus).
A series of codons in part of a messenger RNA (mRNA) molecule. Each codon consists of three nucleotides, usually corresponding to a single amino acid. The nucleotides are abbreviated with the letters A, U, G and C. This is mRNA, which uses U (uracil). DNA uses T (thymine) instead. This mRNA molecule will instruct a ribosome to synthesize a protein according to this code.
The genetic code is the set of rules used by living cells to translate information encoded within genetic material (DNA or mRNA sequences) into proteins. Translation is accomplished by the ribosome, which links amino acids in an order specified by messenger RNA (mRNA), using transfer RNA (tRNA) molecules to carry amino acids and to read the mRNA three nucleotides at a time. The genetic code is highly similar among all organisms and can be expressed in a simple table with 64 entries.[1]
The code defines how sequences of nucleotide triplets, called codons, specify which amino acid will be added next during protein synthesis. With some exceptions,[2] a three-nucleotide codon in a nucleic acid sequence specifies a single amino acid. The vast majority of genes are encoded with a single scheme (see the RNA codon table). That scheme is often referred to as the canonical or standard genetic code, or simply the genetic code, though variant codes (such as in human mitochondria) exist.
While the "genetic code" determines a protein's amino acid sequence, other genomic regions determine when and where these proteins are produced according to various "gene regulatory codes".
Contents
[hide]
1 History
1.1 Codons
1.2 Expanded genetic codes (synthetic biology)
2 Features
2.1 Reading frame
2.2 Start/stop codons
2.3 Effect of mutations
2.4 Degeneracy
2.5 Codon usage bias
3 Standard codon tables
3.1 RNA codon table
3.2 DNA codon table
4 Alternative genetic codes
4.1 Non-standard amino acids
4.2 Variations
4.3 List of alternative codons
5 Origin
6 See also
7 References
8 Further reading
9 External links
History[edit]
The genetic code
Efforts to understand how proteins are encoded began after DNA's structure was discovered in 1953. George Gamow postulated that sets of three bases must be employed to encode the 20 standard amino acids used by living cells to build proteins, which would allow a maximum of 43 = 64 amino acids.[3]
Codons[edit]
The Crick, Brenner experiment first demonstrated that codons consist of three DNA bases. Marshall Nirenberg and Heinrich J. Matthaei were the first to reveal the nature of a codon in 1961.
They used a cell-free system to translate a poly-uracil RNA sequence (i.e., UUUUU...) and discovered that the polypeptide that they had synthesized consisted of only the amino acid phenylalanine.[4] They thereby deduced that the codon UUU specified the amino acid phenylalanine.
This was followed by experiments in Severo Ochoa's laboratory that demonstrated that the poly-adenine RNA sequence (AAAAA...) coded for the polypeptide poly-lysine[5] and that the poly-cytosine RNA sequence (CCCCC...) coded for the polypeptide poly-proline.[6] Therefore, the codon AAA specified the amino acid lysine, and the codon CCC specified the amino acid proline. Using various copolymers most of the remaining codons were then determined.
Subsequent work by Har Gobind Khorana identified the rest of the genetic code. Shortly thereafter, Robert W. Holley determined the structure of transfer RNA (tRNA), the adapter molecule that facilitates the process of translating RNA into protein. This work was based upon Ochoa's earlier studies, yielding the latter the Nobel Prize in Physiology or Medicine in 1959 for work on the enzymology of RNA synthesis.[7]
Extending this work, Nirenberg and Philip Leder revealed the code's triplet nature and deciphered its codons. In these experiments, various combinations of mRNA were passed through a filter that contained ribosomes, the components of cells that translate RNA into protein. Unique triplets promoted the binding of specific tRNAs to the ribosome. Leder and Nirenberg were able to determine the sequences of 54 out of 64 codons in their experiments.[8] Khorana, Holley and Nirenberg received the 1968 Nobel for their work.[9]
The three stop codons were named by discoverers Richard Epstein and Charles Steinberg. "Amber" was named after their friend Harris Bernstein, whose last name means "amber" in German.[10] The other two stop codons were named "ochre" and "opal" in order to keep the "color names" theme.
Expanded genetic codes (synthetic biology)[edit]
Main article: Expanded genetic code
See also: Nucleic acid analogues
In a broad academic audience, the concept of the evolution of the genetic code from the original and ambiguous genetic code to a well-defined ("frozen") code with the repertoire of 20 (+2) canonical amino acids is widely accepted.[11] However, there are different opinions, concepts, approaches and ideas, which is the best way to change it experimentally. Even models are proposed that predict "entry points" for synthetic amino acid invasion of the genetic code.[12]
Since 2001, 40 non-natural amino acids have been added into protein by creating a unique codon (recoding) and a corresponding transfer-RNA:aminoacyl – tRNA-synthetase pair to encode it with diverse physicochemical and biological properties in order to be used as a tool to exploring protein structure and function or to create novel or enhanced proteins.[13] [14]
H. Murakami and M. Sisido extended some codons to have four and five bases. Steven A. Benner constructed a functional 65th (in vivo) codon.[15]
In 2015 N. Budisa, D. Söll and co-workers reported the full substitution of all 20,899 tryptophan residues (UGG codons) with unnatural thienopyrrole-alanine in the genetic code of the bacterium Escherichia coli.[16]
In 2016 the first stable semisynthetic organism was created. It was a (single cell) bacterium with two synthetic bases (called X and Y). The bases survived cell division.[17][18]
In 2017, researchers in South Korea reported that they had engineered a mouse with an extended genetic code that can produce proteins with unnatural amino acids.[19]
Features[edit]
Reading frames in the DNA sequence of a region of the human mitochondrial genome coding for the genes MT-ATP8 and MT-ATP6 (in black: positions 8,525 to 8,580 in the sequence accession NC_012920[20]). There are three possible reading frames in the 5' → 3' forward direction, starting on the first (+1), second (+2) and third position (+3). For each codon (square brackets), the amino acid is given by the vertebrate mitochondrial code, either in the +1 frame for MT-ATP8 (in red) or in the +3 frame for MT-ATP6 (in blue). The MT-ATP8 genes terminates with the TAG stop codon (red dot) in the +1 frame. The MT-ATP6 gene starts with the ATG codon (blue circle for the M amino acid) in the +3 frame.
Reading frame[edit]
A reading frame is defined by the initial triplet of nucleotides from which translation starts. It sets the frame for a run of successive, non-overlapping codons, which is known as an "open reading frame" (ORF). For example, the string 5'-AAATGAACG-3' (see figure), if read from the first position, contains the codons AAA, TGA, and ACG ; if read from the second position, it contains the codons AAT and GAA ; and if read from the third position, it contains the codons ATG and AAC. Every sequence can, thus, be read in its 5' → 3' direction in three reading frames, each producing a possibly distinct amino acid sequence: in the given example, Lys (K)-Trp (W)-Thr (T), Asn (N)-Glu (E), or Met (M)-Asn (N), respectively (when translating with the vertebrate mitochondrial code). When DNA is double-stranded, six possible reading frames are defined, three in the forward orientation on one strand and three reverse on the opposite strand.[21]:330 Protein-coding frames are defined by a start codon, usually the first AUG (ATG) codon in the RNA (DNA) sequence.
In eukaryotes, ORFs in exons are often interrupted by introns.
Start/stop codons[edit]
Translation starts with a chain-initiation codon or start codon. The start codon alone is not sufficient to begin the process. Nearby sequences such as the Shine-Dalgarno sequence in E. coli and initiation factors are also required to start translation. The most common start codon is AUG, which is read as methionine or, in bacteria, as formylmethionine. Alternative start codons depending on the organism include "GUG" or "UUG"; these codons normally represent valine and leucine, respectively, but as start codons they are translated as methionine or formylmethionine.[22]
The three stop codons have names: UAG is amber, UGA is opal (sometimes also called umber), and UAA is ochre. Stop codons are also called "termination" or "nonsense" codons. They signal release of the nascent polypeptide from the ribosome because no cognate tRNA has anticodons complementary to these stop signals, allowing a release factor to bind to the ribosome instead.[23]
Effect of mutations[edit]
Examples of notable mutations that can occur in humans.[24]
During the process of DNA replication, errors occasionally occur in the polymerization of the second strand. These errors, mutations, can affect an organism's phenotype, especially if they occur within the protein coding sequence of a gene. Error rates are typically 1 error in every 10–100 million bases—due to the "proofreading" ability of DNA polymerases.[25][26]
Missense mutations and nonsense mutations are examples of point mutations that can cause genetic diseases such as sickle-cell disease and thalassemia respectively.[27][28][29] Clinically important missense mutations generally change the properties of the coded amino acid residue among basic, acidic, polar or non-polar states, whereas nonsense mutations result in a stop codon.[21]:266
Mutations that disrupt the reading frame sequence by indels (insertions or deletions) of a non-multiple of 3 nucleotide bases are known as frameshift mutations. These mutations usually result in a completely different translation from the original, and likely cause a stop codon to be read, which truncates the protein.[30] These mutations may impair the protein's function and are thus rare in in vivo protein-coding sequences. One reason inheritance of frameshift mutations is rare is that, if the protein being translated is essential for growth under the selective pressures the organism faces, absence of a functional protein may cause death before the organism becomes viable.[31] Frameshift mutations may result in severe genetic diseases such as Tay-Sachs disease.[32]
Although most mutations that change protein sequences are harmful or neutral, some mutations have benefits.[33] These mutations may enable the mutant organism to withstand particular environmental stresses better than wild type organisms, or reproduce more quickly. In these cases a mutation will tend to become more common in a population through natural selection.[34] Viruses that use RNA as their genetic material have rapid mutation rates,[35] which can be an advantage, since these viruses thereby evolve rapidly, and thus evade the immune system defensive responses.[36] In large populations of asexually reproducing organisms, for example, E. coli, multiple beneficial mutations may co-occur. This phenomenon is called clonal interference and causes competition among the mutations.[37]
Degeneracy[edit]
Main article: Codon degeneracy
Degeneracy is the redundancy of the genetic code. This term was given by Bernfield and Nirenberg. The genetic code has redundancy but no ambiguity (see the codon tables below for the full correlation). For example, although codons GAA and GAG both specify glutamic acid (redundancy), neither specifies another amino acid (no ambiguity). The codons encoding one amino acid may differ in any of their three positions. For example, the amino acid leucine is specified by YUR or CUN (UUA, UUG, CUU, CUC, CUA, or CUG) codons (difference in the first or third position indicated using IUPAC notation), while the amino acid serine is specified by UCN or AGY (UCA, UCG, UCC, UCU, AGU, or AGC) codons (difference in the first, second, or third position).[38]:102–117 :521–522 A practical consequence of redundancy is that errors in the third position of the triplet codon cause only a silent mutation or an error that would not affect the protein because the hydrophilicity or hydrophobicity is maintained by equivalent substitution of amino acids; for example, a codon of NUN (where N = any nucleotide) tends to code for hydrophobic amino acids. NCN yields amino acid residues that are small in size and moderate in hydropathy; NAN encodes average size hydrophilic residues. The genetic code is so well-structured for hydropathy that a mathematical analysis (Singular Value Decomposition) of 12 variables (4 nucleotides x 3 positions) yields a remarkable correlation (C = 0.95) for predicting the hydropathy of the encoded amino acid directly from the triplet nucleotide sequence, without translation.[39][40] Note in the table, below, eight amino acids are not affected at all by mutations at the third position of the codon, whereas in the figure above, a mutation at the second position is likely to cause a radical change in the physicochemical properties of the encoded amino acid.
Grouping of codons by amino acid residue molar volume and hydropathy. A more detailed version is available.
Codon usage bias[edit]
The frequency of codons, also known as codon usage bias, can vary from species to species with functional implications for the control of translation. The following codon usage table is for the human genome.[41]
[show]Human genome codon frequency table
Human genome codon frequency
Codon
AA[A]
Fraction[B]
Freq ‰[C]
Number[D]
Codon
AA
Fraction
Freq ‰
Number
Codon
AA
Fraction
Freq ‰
Number
Codon
AA
Fraction
Freq ‰
Number
UUU F 0.46 17.6 714,298 UCU S 0.19 15.2 618,711 UAU Y 0.44 12.2 495,699 UGU C 0.46 10.6 430,311
UUC F 0.54 20.3 824,692 UCC S 0.22 17.7 718,892 UAC Y 0.56 15.3 622,407 UGC C 0.54 12.6 513,028
UUA L 0.08 7.7 311,881 UCA S 0.15 12.2 496,448 UAA * 0.30 1.0 40,285 UGA * 0.47 1.6 63,237
UUG L 0.13 12.9 525,688 UCG S 0.05 4.4 179,419 UAG * 0.24 0.8 32,109 UGG W 1.00 13.2 535,595
CUU L 0.13 13.2 536,515 CCU P 0.29 17.5 713,233 CAU H 0.42 10.9 441,711 CGU R 0.08 4.5 184,609
CUC L 0.20 19.6 796,638 CCC P 0.32 19.8 804,620 CAC H 0.58 15.1 613,713 CGC R 0.18 10.4 423,516
CUA L 0.07 7.2 290,751 CCA P 0.28 16.9 688,038 CAA Q 0.27 12.3 501,911 CGA R 0.11 6.2 250,760
CUG L 0.40 39.6 1,611,801 CCG P 0.11 6.9 281,570 CAG Q 0.73 34.2 1,391,973 CGG R 0.20 11.4 464,485
AUU I 0.36 16.0 650,473 ACU T 0.25 13.1 533,609 AAU N 0.47 17.0 689,701 AGU S 0.15 12.1 493,429
AUC I 0.47 20.8 846,466 ACC T 0.36 18.9 768,147 AAC N 0.53 19.1 776,603 AGC S 0.24 19.5 791,383
AUA I 0.17 7.5 304,565 ACA T 0.28 15.1 614,523 AAA K 0.43 24.4 993,621 AGA R 0.21 12.2 494,682
AUG M 1.00 22.0 896,005 ACG T 0.11 6.1 246,105 AAG K 0.57 31.9 1,295,568 AGG R 0.21 12.0 486,463
GUU V 0.18 11.0 448,607 GCU A 0.27 18.4 750,096 GAU D 0.46 21.8 885,429 GGU G 0.16 10.8 437,126
GUC V 0.24 14.5 588,138 GCC A 0.40 27.7 1,127,679 GAC D 0.54 25.1 1,020,595 GGC G 0.34 22.2 903,565
GUA V 0.12 7.1 287,712 GCA A 0.23 15.8 643,471 GAA E 0.42 29.0 1,177,632 GGA G 0.25 16.5 669,873
GUG V 0.46 28.1 1,143,534 GCG A 0.11 7.4 299,495 GAG E 0.58 39.6 1,609,975 GGG G 0.25 16.5 669,768
A Amino acid. B Fraction of each codon among all those specifying a given amino acid. C Frequency among 40,662,582 codons of 93,487 coding sequences. D Number.
Standard codon tables[edit]
RNA codon table[edit]
Amino acids biochemical properties nonpolar polar basic acidic Termination: stop codon
Standard genetic code
1st
base
2nd base
3rd
base
U
C
A
G
U
UUU (Phe/F) Phenylalanine UCU (Ser/S) Serine UAU (Tyr/Y) Tyrosine UGU (Cys/C) Cysteine
U
UUC UCC UAC UGC
C
UUA (Leu/L) Leucine UCA UAA[B] Stop (Ochre) UGA[B] Stop (Opal)
A
UUG UCG UAG[B] Stop (Amber) UGG (Trp/W) Tryptophan
G
C
CUU CCU (Pro/P) Proline CAU (His/H) Histidine CGU (Arg/R) Arginine
U
CUC CCC CAC CGC
C
CUA CCA CAA (Gln/Q) Glutamine CGA
A
CUG CCG CAG CGG
G
A
AUU (Ile/I) Isoleucine ACU (Thr/T) Threonine AAU (Asn/N) Asparagine AGU (Ser/S) Serine
U
AUC ACC AAC AGC
C
AUA ACA AAA (Lys/K) Lysine AGA (Arg/R) Arginine
A
AUG[A] (Met/M) Methionine ACG AAG AGG
G
G
GUU (Val/V) Valine GCU (Ala/A) Alanine GAU (Asp/D) Aspartic acid GGU (Gly/G) Glycine
U
GUC GCC GAC GGC
C
GUA GCA GAA (Glu/E) Glutamic acid GGA
A
GUG GCG GAG GGG
G
A The codon AUG both codes for methionine and serves as an initiation site: the first AUG in an mRNA's coding region is where translation into protein begins.[42]
B ^ ^ ^ The historical basis for designating the stop codons as amber, ochre and opal is described in an autobiography by Sydney Brenner[43] and in a historical article by Bob Edgar.[44]
Inverse table for the standard genetic code (compressed using IUPAC notation)
Amino acid
Codons
Compressed
Amino acid
Codons
Compressed
Ala / A
GCU, GCC, GCA, GCG GCN
Leu / L
UUA, UUG, CUU, CUC, CUA, CUG YUR, CUN
Arg / R
CGU, CGC, CGA, CGG, AGA, AGG CGN, MGR
Lys / K
AAA, AAG AAR
Asn / N
AAU, AAC AAY
Met / M
AUG
Asp / D
GAU, GAC GAY
Phe / F
UUU, UUC UUY
Cys / C
UGU, UGC UGY
Pro / P
CCU, CCC, CCA, CCG CCN
Gln / Q
CAA, CAG CAR
Ser / S
UCU, UCC, UCA, UCG, AGU, AGC UCN, AGY
Glu / E
GAA, GAG GAR
Thr / T
ACU, ACC, ACA, ACG ACN
Gly / G
GGU, GGC, GGA, GGG GGN
Trp / W
UGG
His / H
CAU, CAC CAY
Tyr / Y
UAU, UAC UAY
Ile / I
AUU, AUC, AUA AUH
Val / V
GUU, GUC, GUA, GUG GUN
START
AUG
STOP
UAA, UGA, UAG UAR, URA
DNA codon table[edit]
Main article: DNA codon table
The DNA codon table is essentially identical to that for RNA, but with U replaced by T.
Alternative genetic codes[edit]
Non-standard amino acids[edit]
In some proteins, non-standard amino acids are substituted for standard stop codons, depending on associated signal sequences in the messenger RNA. For example, UGA can code for selenocysteine and UAG can code for pyrrolysine. Selenocysteine became to be seen as the 21st amino acid, and pyrrolysine as the 22nd.[45] Unlike selenocysteine, pyrrolysine-encoded UAG is translated with the participation of a dedicated aminoacyl-tRNA synthetase.[46] Both selenocysteine and pyrrolysine may be present in the same organism.[47] Although the genetic code is normally fixed in an organism, the achaeal prokaryote Acetohalobium arabaticum can expand its genetic code from 20 to 21 amino acids (by including pyrrolysine) under different conditions of growth.[48]
Variations[edit]
See also: List of genetic codes
Genetic code logo of the Globobulimina pseudospinescens mitochondrial genome. The logo shows the 64 codons from left to right, predicted alternatives in red (relative to the standard genetic code). Red line: stop codons. The height of each amino acid in the stack shows how often it is aligned to the codon in homologous protein domains. The stack height indicates the support for the prediction.
Variations on the standard code were predicted in the 1970s.[49] The first was discovered in 1979, by researchers studying human mitochondrial genes.[50] Many slight variants were discovered thereafter,[45] including various alternative mitochondrial codes.[51] Small variants such as translation of the codon UGA as tryptophan in Mycoplasma species, and translation of CUG as a serine rather than leucine in yeasts of the "CTG clade" (such as Candida albicans).[52][53][54] Because viruses must use the same genetic code as their hosts, modifications to the standard genetic code could interfere with viral protein synthesis or functioning.[55] However, viruses such as totiviruses adapted to the host's genetic code modification.[56] In bacteria and archaea, GUG and UUG are common start codons. In rare cases, certain proteins may use alternative start codons.[45] Surprisingly, variations in the interpretation of the genetic code exist also in human nuclear-encoded genes: In 2016, researchers studying the translation of malate dehydrogenase found that in about 4% of the mRNAs encoding this enzyme the stop codon is naturally used to encode the amino acids tryptophan and arginine.[57] This type of recoding is induced by a high-readthrough stop codon context[58] and it is referred to as functional translational readthrough.[59]
Variant genetic codes used by an organism can be inferred by identifying highly conserved genes encoded in that genome, and comparing its codon usage to the amino acids in homologous proteins of other organisms. For example, the program FACIL[60] infers a genetic code by searching which amino acids in homologous protein domains are most often aligned to every codon. The resulting amino acid probabilities for each codon are displayed in a genetic code logo, that also shows the support for a stop codon.
Despite these differences, all known naturally occurring codes are very similar. The coding mechanism is the same for all organisms: three-base codons, tRNA, ribosomes, single direction reading and translating single codons into single amino acids.[61]
List of alternative codons[edit]
[show]List of alternative codons
Amino acids biochemical properties nonpolar polar basic acidic Termination: stop codon
Comparison between codon translations with alternative and standard genetic codes
Code
Translation
table
DNA codon involved
RNA codon involved
Translation
with this code
Standard translation
Notes
Standard 1 Includes translation table 8 (plant chloroplasts).
Vertebrate mitochondrial 2 AGA AGA Ter (*) Arg (R)
AGG AGG Ter (*) Arg (R)
ATA AUA Met (M) Ile (I)
TGA UGA Trp (W) Ter (*)
Yeast mitochondrial 3 ATA AUA Met (M) Ile (I)
CTT CUU Thr (T) Leu (L)
CTC CUC Thr (T) Leu (L)
CTA CUA Thr (T) Leu (L)
CTG CUG Thr (T) Leu (L)
TGA UGA Trp (W) Ter (*)
CGA CGA absent Arg (R)
CGC CGC absent Arg (R)
Mold, protozoan, and coelenterate mitochondrial + Mycoplasma / Spiroplasma 4 TGA UGA Trp (W) Ter (*) Includes the translation table 7 (kinetoplasts).
Invertebrate mitochondrial 5 AGA AGA Ser (S) Arg (R)
AGG AGG Ser (S) Arg (R)
ATA AUA Met (M) Ile (I)
TGA UGA Trp (W) Ter (*)
Ciliate, dasycladacean and Hexamita nuclear 6 TAA UAA Gln (Q) Ter (*)
TAG UAG Gln (Q) Ter (*)
Echinoderm and flatworm mitochondrial 9 AAA AAA Asn (N) Lys (K)
AGA AGA Ser (S) Arg (R)
AGG AGG Ser (S) Arg (R)
TGA UGA Trp (W) Ter (*)
Euplotid nuclear 10 TGA UGA Cys (C) Ter (*)
Bacterial, archaeal and plant plastid 11 See translation table 1.
Alternative yeast nuclear 12 CTG CUG Ser (S) Leu (L)
Ascidian mitochondrial 13 AGA AGA Gly (G) Arg (R)
AGG AGG Gly (G) Arg (R)
ATA AUA Met (M) Ile (I)
TGA UGA Trp (W) Ter (*)
Alternative flatworm mitochondrial 14 AAA AAA Asn (N) Lys (K)
AGA AGA Ser (S) Arg (R)
AGG AGG Ser (S) Arg (R)
TAA UAA Tyr (Y) Ter (*)
TGA UGA Trp (W) Ter (*)
Blepharisma nuclear 15 TAG UAG Gln (Q) Ter (*) As of Nov. 18, 2016: absent from the NCBI update.
Chlorophycean mitochondrial 16 TAG UAG Leu (L) Ter (*)
Trematode mitochondrial 21 TGA UGA Trp (W) Ter (*)
ATA AUA Met (M) Ile (I)
AGA AGA Ser (S) Arg (R)
AGG AGG Ser (S) Arg (R)
AAA AAA Asn (N) Lys (K)
Scenedesmus obliquus mitochondrial 22 TCA UCA Ter (*) Ser (S)
TAG UAG Leu (L) Ter (*)
Thraustochytrium mitochondrial 23 TTA UUA Ter (*) Leu (L) Similar to translation table 11.
Pterobranchia mitochondrial 24 AGA AGA Ser (S) Arg (R)
AGG AGG Lys (K) Arg (R)
TGA UGA Trp (W) Ter (*)
Candidate division SR1 and Gracilibacteria 25 TGA UGA Gly (G) Ter (*)
Pachysolen tannophilus nuclear 26 CTG CUG Ala (A) Leu (L)
Karyorelict nuclear 27 TAA UAA Gln (Q) Ter (*)
TAG UAG Gln (Q) Ter (*)
TGA UGA Ter (*) or Trp (W) Ter (*)
Condylostoma nuclear 28 TAA UAA Ter (*) or Gln (Q) Ter (*)
TAG UAG Ter (*) or Gln (Q) Ter (*)
TGA UGA Ter (*) or Trp (W) Ter (*)
Mesodinium nuclear 29 TAA UAA Tyr (Y) Ter (*)
TAG UAG Tyr (Y) Ter (*)
Peritrich nuclear 30 TAA UAA Glu (E) Ter (*)
TAG UAG Glu (E) Ter (*)
Blastocrithidia nuclear 31 TAA UAA Ter (*) or Gln (Q) Ter (*)
TAG UAG Ter (*) or Gln (Q) Ter (*)
TGA UGA Trp (W) Ter (*)
Origin[edit]
The genetic code is a key part of the story of life, according to which self-replicating RNA molecules preceded life as we know it. The main hypothesis for life's origin is the RNA world hypothesis. Any model for the emergence of genetic code is intimately related to a model of the transfer from ribozymes (RNA enzymes) to proteins as the principal enzymes in cells. In line with the RNA world hypothesis, transfer RNA molecules appear to have evolved before modern aminoacyl-tRNA synthetases, so the latter cannot be part of the explanation of its patterns.[62]
A hypothetical randomly evolved genetic code further motivates a biochemical or evolutionary model for its origin. If amino acids were randomly assigned to triplet codons, there would be 1.5 × 1084 possible genetic codes.[63]:163 This number is found by calculating the number of ways that 21 items (20 amino acids plus one stop) can be placed in 64 bins, wherein each item is used at least once.[64] However, the distribution of codon assignments in the genetic code is nonrandom.[65] In particular, the genetic code clusters certain amino acid assignments.
Amino acids that share the same biosynthetic pathway tend to have the same first base in their codons. This could be an evolutionary relic of an early, simpler genetic code with fewer amino acids that later evolved to code a larger set of amino acids.[66] It could also reflect steric and chemical properties that had another effect on the codon during its evolution. Amino acids with similar physical properties also tend to have similar codons,[67][68] reducing the problems caused by point mutations and mistranslations.[65]
Given the non-random genetic triplet coding scheme, a tenable hypothesis for the origin of genetic code could address multiple aspects of the codon table, such as absence of codons for D-amino acids, secondary codon patterns for some amino acids, confinement of synonymous positions to third position, the small set of only 20 amino acids (instead of a number approaching 64), and the relation of stop codon patterns to amino acid coding patterns.[69]
Three main hypotheses address the origin of the genetic code. Many models belong to one of them or to a hybrid:[70]
Random freeze: the genetic code was randomly created. For example, early tRNA-like ribozymes may have had different affinities for amino acids, with codons emerging from another part of the ribozyme that exhibited random variability. Once enough peptides were coded for, any major random change in the genetic code would have been lethal; hence it became "frozen".[71]
Stereochemical affinity: the genetic code is a result of a high affinity between each amino acid and its codon or anti-codon; the latter option implies that pre-tRNA molecules matched their corresponding amino acids by this affinity. Later during evolution, this matching was gradually replaced with matching by aminoacyl-tRNA synthetases.[69][72][73]
Optimality: the genetic code continued to evolve after its initial creation, so that the current code maximizes some fitness function, usually some kind of error minimization.[69][70]
Hypotheses have addressed a variety of scenarios:[74]
Chemical principles govern specific RNA interaction with amino acids. Experiments with aptamers showed that some amino acids have a selective chemical affinity for their codons.[75] Experiments showed that of 8 amino acids tested, 6 show some RNA triplet-amino acid association.[63][73]
Biosynthetic expansion. The genetic code grew from a simpler earlier code through a process of "biosynthetic expansion". Primordial life "discovered" new amino acids (for example, as by-products of metabolism) and later incorporated some of these into the machinery of genetic coding.[76] Although much circumstantial evidence has been found to suggest that fewer amino acid types were used in the past,[77] precise and detailed hypotheses about which amino acids entered the code in what order are controversial.[78][79]
Natural selection has led to codon assignments of the genetic code that minimize the effects of mutations.[80] A recent hypothesis[81] suggests that the triplet code was derived from codes that used longer than triplet codons (such as quadruplet codons). Longer than triplet decoding would increase codon redundancy and would be more error resistant. This feature could allow accurate decoding absent complex translational machinery such as the ribosome, such as before cells began making ribosomes.
Information channels: Information-theoretic approaches model the process of translating the genetic code into corresponding amino acids as an error-prone information channel.[82] The inherent noise (that is, the error) in the channel poses the organism with a fundamental question: how can a genetic code be constructed to withstand noise[83] while accurately and efficiently translating information? These "rate-distortion" models[84] suggest that the genetic code originated as a result of the interplay of the three conflicting evolutionary forces: the needs for diverse amino acids,[85] for error-tolerance[80] and for minimal resource cost. The code emerges at a transition when the mapping of codons to amino acids becomes nonrandom. The code's emergence is governed by the topology defined by the probable errors and is related to the map coloring problem.[86]
Game theory: Models based on signaling games combine elements of game theory, natural selection and information channels. Such models have been used to suggest that the first polypeptides were likely short and had non-enzymatic function. Game theoretic models suggested that the organization of RNA strings into cells may have been necessary to prevent "deceptive" use of the genetic code, i.e. preventing the ancient equivalent of viruses from overwhelming the RNA world.[87]
Stop codons: Codons for translational stops are also an interesting aspect to the problem of the origin of the genetic code. As an example for addressing stop codon evolution, it has been suggested that the stop codons are such that they are most likely to terminate translation early in the case of a frame shift error.[88] In contrast, some stereochemical molecular models explain the origin of stop codons as "unassignable".[69]
See also[edit]
List of genetic engineering software
References[edit]
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Further reading[edit]
Griffiths AJ, Miller JH, Suzuki DT, Lewontin RC, Gilbert WM (1999). An Introduction to genetic analysis (7th ed.). San Francisco: W.H. Freeman. ISBN 0-7167-3771-X.
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular biology of the cell (4th ed.). New York: Garland Science. ISBN 0-8153-3218-1.
Lodish HF, Berk A, Zipursky SL, Matsudaira P, Baltimore D, Darnell JE (2000). Molecular cell biology (4th ed.). San Francisco: W.H. Freeman. ISBN 0-7167-3706-X.
Caskey CT, Leder P (Apr 2014). "The RNA code: nature's Rosetta Stone". Proceedings of the National Academy of Sciences of the United States of America. 111 (16): 5758–9. Bibcode:2014PNAS..111.5758C. doi:10.1073/pnas.1404819111. PMC 4000803  . PMID 24756939.
External links[edit]
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The Genetic Codes → Genetic Code Tables
The Codon Usage Database → Codon frequency tables for many organisms
History of deciphering the genetic code
American Scientist: Ode to the code (Origin)
Alphabet of Life (Origin)
Symmetries in the genetic code
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Mere Sapno Ki Rani
From Wikipedia, the free encyclopedia
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"Mere Sapno Ki Rani"
Film song by Kishore Kumar
from the album Aradhana
Language
Hindi
Released
1969
Length
5:00
Label
Saregama
Composer(s)
Sachin Dev Burman
Lyricist(s)
Anand Bakshi
"Mere Sapno Ki Rani" is a popular Hindi song from the 1969 Bollywood movie Aradhana. The song was composed by Sachin Dev Burman and performed by Kishore Kumar. In the film, Rajesh Khanna, the male protagonist, sings the song atop an open jeep virtually chasing the heroine, Sharmila Tagore, who was aboard the toy train in Darjeeling (Darjeeling Himalayan Railway).[1][2] R. D. Burman composed the song after his father, S.D. Burman, became too ill to finish the film's music. The song became a superhit in India.[2] Many other versions of this song were made such as the song "Atha Dilisena Hiru" by famous Sri Lankan singer H. R. Jothipala[3] The song was re-used in the Tamil film, Eli (2015).[4]
Picturization[edit]
In the film, Rajesh Khanna stars as Arun Verma, a young pilot from the Indian Air Force. On the Darjeeling toy train, Sharmila Tagore is Vandana, a recent college graduate who is returning to her hill station home. Arun and his friend, Madan (Sujit Kumar) have taken leave without permission, and as Madan drives the open top jeep, Arun sings the song in Kishore Kumar's voice, flirtatiously eying Vandana, and later in the film, when they meet again, they fall in love.
Singer[edit]
Originally, Mohammed Rafi was supposed to have sung all the songs in the film, but the composer, Sachin Dev Burman fell ill. His son, R.D. Burman reportedly ghost-composed this song, and took a chance and gave the song to Kishore Kumar.[5] It, along with "Roop Tera Mastana", turned Kishore into an overnight sensation along with Rajesh Khanna, and the two became best friends in the industry.
References[edit]
Jump up ^ Mahalingam, Sudha (March 2001). "Darjeeling: Where the journey is the destination". Travelogues. Outlook Traveller. Outlook Publishing (India) Private Limited. Archived from the original on 2006-12-10. Retrieved 2007-03-09.
^ Jump up to: a b "Darjeeling Toy Train". Theme India: Train Tourism in India. IndiaLine. Retrieved 2007-03-09. External link in |work= (help)
Jump up ^ SinhalaJukebox - A Jukebox of Songs from Sri Lanka (siinhalajukebox.org)
Jump up ^ Naig, Udhav (13 May 2015). "Won't rule out another stint in politics, says Vadivelu". The Hindu. Retrieved 26 May 2015.
Jump up ^ Bharatan, Raju (31 October 2000). "The Aradhana syndrome and SD Burman". Retrieved 2012-10-18.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Mere_Sapno_Ki_Rani&oldid=784492917"
Categories:
1969 songs
Hindi film songs
Indian songs
Kishore Kumar songs
Songs with music by S. D. Burman
Songs with lyrics by Anand Bakshi
Transport culture of India
Railway culture
Hidden categories:
CS1 errors: external links
Music infoboxes with unknown value for type
Articles with hAudio microformats
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This page was last edited on 8 June 2017, at 16:47.
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text:
Love It or List It
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Love It or List It
Genre
Reality renovations, upgrading
Created by
Maria Armstrong
Catherine Fogarty
Directed by
Various
Starring
Hilary Farr
David Visentin
Narrated by
Jacqueline Hennessy
Theme music composer
Lou Pomanti
Country of origin
Canada
United States
Original language(s)
English
No. of seasons
10
No. of episodes
130 (List of episodes)
Production
Executive producer(s)
Maria Armstrong
Catherine Fogarty
Producer(s)
Maria Armstrong
Location(s)
Toronto, Ontario, Canada, Raleigh–Durham, North Carolina, United States
Running time
44 minutes
Production company(s)
Big Coat Productions
Release
Original network
W Network
HGTV
Original release
September 8, 2008 – present
External links
Website
Production website
Love It or List It is a Canadian home design TV show currently airing on HGTV, W Network, and on OWN Canada, and is the original show in the Love it or List It franchise. The show is produced by Big Coat Productions and was based in Toronto and other surrounding areas in Ontario, Canada. The show premiered as a primetime program on W Network on September 8, 2008, and has since aired on OWN Canada as well as HGTV in the United States. In September 2014, the show began filming in the United States in North Carolina.[1]
In September 2017, HGTV ordered 20 additional episodes of the program. The next season is to start airing in July 2018.[2]
Contents
[hide]
1 Format
2 Host and crew
3 Episodes
4 Franchise
5 International syndication
6 Reception
7 Controversy
8 References
9 External links
Format[edit]
This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. (November 2016) (Learn how and when to remove this template message)
Every episode of Love It or List It follows the same formula. A couple (usually, but not in all cases) presents their living situation in the current house that they own. In most episodes, the couple is split on whether or not they want to stay in the house.
Interior designer Hilary Farr and real estate agent David Visentin tour the couple's home before meeting with them. The tour usually consists of David finding mostly negative things to say about the residence while Hilary is convinced that she can work magic with whatever plans she is given.
At the meetup between the couple and the hosts of the show, both Hilary and David are briefed as to what the couple's desires are. Hilary is given a list of renovations the couple wants for the current house and her budget for the entire project. David, meanwhile, is tasked with searching for a new home for the couple that both meets their needs and stays within their desired budget.
Common challenges faced by Hilary are an inadequate budget to complete the entire request list from the homeowners, often due to discovery of unforeseen issues with the house that are uncovered during the renovation such as lack of compliance with modern building codes. Common issues for David, meanwhile, depend on the homeowners' desires; for instance, the couple has children enrolled in the neighborhood school and they do not desire to change, or the potential house is too distant from family members or a workplace.
After Hilary's renovation is complete, the homeowners are given a tour of their current home to see what she was able to accomplish. After the tour, David meets with them and hands them an evaluation of the home's current market value following the renovations. He will then remind the couple what they could have in one of the new homes they looked at and that they would not get that in their current home.
After a moment to deliberate, Hilary and David pose a question to the homeowners. They must choose to either Love It, meaning that they will continue to live in their current home with the renovations, or to List It, meaning that they will buy one of the homes David showed them and sell their current home. After they reveal their decision, the homeowners explain their reasoning to Hilary and David (who, if they decided to Love It, reacts with incredulity) before bidding them farewell.
Host and crew[edit]
Hosts
Hilary Farr – Hilary Farr is a home designer from Toronto, Ontario. She has lived in Australia, England, California, and New York City. Farr honed her skills on properties in Los Angeles, Santa Barbara, New York and Toronto. When she first moved back to Toronto, she became the first designer to "stage" properties for sale. She continues to build and design homes in the downtown core where she herself owns properties.[3]
David Visentin – David Visentin is a real estate agent in Southern Ontario with Country Living Realty Limited. He has been practicing since 1987.[4]
Assistant Designer
Desta Ostapyk (Canadian episodes) – Desta is a Toronto-based designer who graduated in 2004 from the Toronto International Academy of Design and Technology in Interior Design, and has since focused on a career in the Television Industry. She first started working with Big Coat Productions during her last semester of school as an intern for HGTV's hit series My Parents House. She soon became the Design Stylist for the show.[5]
Contractors
Eric Eremita (North Carolina episodes) – General contractor and designer Eric Eremita was selected to be the one and only general contractor on HGTV's "Love It or List It" U.S. version, after HGTV network took notice of him when he competed in its "Brother vs. Brother" reality show.
Eddie Richardson (Canadian episodes) – Eddie Richardson was a contractor on Love It or List It who started his own family business. Richardson has also been a pro-beach volleyball player and professional bass fisherman.
Fergus McLaren (Canadian episodes) – Fergus McLaren started up his own construction company, R-Mac Solutions, 10 years ago.
Behind the Scenes
Architect: Simon West
Senior Production Coordinator: Linda Johnstone
Construction Coordinator: David Violante
Construction Assistants: Chris Blinn, Adam Dalgarno, Ahren Mrowietz, Dale George
Design Coordinator: Kaaveh Shoman
Episodes[edit]
Main article: List of Love It or List It episodes
Season
Start date
End date
Episodes
Hilary wins
David wins
1
September 8, 2008 January 5, 2009 12 9 3
2
April 6, 2009 December 7, 2009 18 10 8
3
May 3, 2010 November 8, 2010 21 9 12
4
March 14, 2011 November 11, 2011 19 12 7
5
February 20, 2012 November 19, 2012 18 10 8
6
January 19, 2013 July 1, 2013 14 8 6
7
January 8, 2014 April 9, 2014 14 11 3
8
September 8, 2014 January 19, 2015 16 8 8
9
March 9, 2015 December 7, 2015 22 13 9
Total
154
90
64
Victories for Hilary are families or clients who decided to love their home and stay. Victories for David are families and clients who decided to list and move into a new or better home.
Franchise[edit]
Main article: Love It or List It (franchise)
Love It or List It has spawned five spinoffs. The first,[6] known as Love It or List It Vancouver (or Love it or List it Too in the US), was launched in winter 2012 and is hosted by Jillian Harris and Todd Talbot. The second spin-off, a British version known as Love It or List It UK, debuted in 2015,[7] is hosted by Kirstie Allsopp and Phil Spencer. The third spin off, Love It or List It Vacation Homes[8] debuted in spring 2016 and is hosted by Dan Vickery and Elisa Goldhawke. A fourth spin off, Vendre ou renover au Quebec debuted in January 2017 and is hosted by Maika Desnoyers and Daniel Corbin.[9] The fifth spin off, Love It Or List It Australia debuted in September 2017 and is hosted by Andrew Winter and Neale Whittaker.[10]
International syndication[edit]
Country / Region
Name
Television Network
Dubbing / Subtitles
Australia Love It or List It Lifestyle Home N/A
Canada Love It or List It W Network N/A
USA Love It or List It HGTV[11] N/A
Spain Tu casa a juicio Divinity Spanish
Brazil Ame-a ou Deixe-a Discovery Home & Health Portuguese
Norway Bolighjelpen TV 2 (Norway) Norwegian
Bulgaria Podnovi ili proday Fox Life (Bulgaria) Bulgarian
Italy Prendere o lasciare Cielo dubbed in Italian
Reception[edit]
On August 31, 2010, Love It or List It was nominated for two Gemini Awards: Best Reality Program or Series and Best Direction in a Reality Program or Series. When HGTV premiered the show on the network, the company stated that Love It or List It has been the highest rating reality series since Candice Olson's Candice Tells All.[12]
In 2012, New York Times' columnist Gail Collins noted that it was US Secretary of State Hillary Clinton's favorite TV show.[13][14] According to Collins, Clinton finds the show "very calming"[15] after being interviewed about her departure from politics.[16][17][18]
In a 2013 interview with Las Vegas Magazine, Vanna White from Wheel of Fortune said it was one of her favourite HGTV programs.[19] Actress Julianne Moore also gave similar praise for the show in an interview with Katie Couric and the Daily Mail.[20][21]
According to the Wall Street Journal, the show and its Vancouver spinoff is one of Canada's beloved reality series.[22]
Controversy[edit]
In April 2016, homeowners Deanna Murphy and Tim Sullivan who had participated in a 2015 Love It or List It episode filed suit against production company Big Coat TV, as well as the North Carolina contractor (Aaron Fitz Construction)[23] who had been hired by the show to do the renovations on their home. The couple alleges that the renovation funds that they provided were not properly disbursed, and that the work on their home was done to a substandard quality.[24] Moreover, the lawsuit states that the television personalities on the show do not play an active role in the renovation process, and that they were not shown homes on the market by any licensed North Carolina real estate agent. Big Coat TV has commented that they "do intend to vigorously defend what [they] consider to be false allegations."[25]
The suit was settled in April 2017.[26] The plaintiffs had signed a confidentiality agreement; their lawyer would not comment on the settlement.[27] Big Coat had previously filed a countersuit for libel, slander and product disparagement; parts of that suit had been dismissed by the time of the settlement but that was under appeal by Big Coat. After the agreements had been concluded, both suits were dismissed.[28]
References[edit]
Jump up ^ http://www.bigcoatproductions.com/blog/post/love-it-or-list-it-is-on-the-move-to-the-us. Big Coat Productions. Retrieved 12 December 2014
Jump up ^ "HGTV Orders 20 New Episodes of Hit Series 'Love It or List It' | Scripps Networks Interactive Newsroom". newsroom.scrippsnetworksinteractive.com. Retrieved 2018-01-02.
Jump up ^ Host Bios; Hilary Farr "Archived copy". Archived from the original on July 18, 2011. Retrieved June 10, 2011.
Jump up ^ Host Bios; David Visentin "Archived copy". Archived from the original on July 18, 2011. Retrieved June 10, 2011.
Jump up ^ "Big Coat Productions". Big Coat Productions. Archived from the original on July 16, 2013. Retrieved March 22, 2013.
Jump up ^ Love it or List it launches new spin off
Jump up ^ "Kirstie and Phil's Love It or List It will return for a second series:Channel 4 orders six more episodes of the show to air in 2016". Digital Spy. 2 June 2015. Retrieved 14 March 2016.
Jump up ^ http://www.bigcoatmedia.com/
Jump up ^ http://www.hgtv.ca/tags/love-it-or-list-it-vacation-homes/
Jump up ^ http://www.news.com.au/entertainment/tv/reality-tv/i-fear-pineapples-are-very-much-the-new-owls-love-it-or-list-it-hosts-kiss-and-tell-on-renos/news-story/935d73b586c68943a6a455f02354bef9
Jump up ^ Vlessing, Etan (November 28, 2011). "HGTV Acquires Canadian Lifestyle Series 'Love It Or List It'". The Hollywood Reporter.
Jump up ^ W Network's Top Rated Show; http://corusent.com/home/Corporate/PressReleases/tabid/1697/Default.aspx?Id=2323
Jump up ^ Maga, Carly (March 29, 2012). "Hillary Clinton's favorite show is Canada's 'Love It Or List It'". Ca.omg.yahoo.com. Retrieved March 22, 2013.
Jump up ^ Miller, Julie. "Hillary Clinton's Unlikely Favorite TV Show Revealed". Vanity Fair. Retrieved March 22, 2013.
Jump up ^ "Vanity Fair: Hillary Clinton's Unlikely Favorite TV Show Revealed". Huffingtonpost.com. November 13, 2012. Retrieved March 22, 2013.
Jump up ^ "Hillary Clinton likes 'Love It or List It'". Politico.Com. Retrieved March 22, 2013.
Jump up ^ "Love it or list it News, Video and Gossip". Jezebel. November 11, 2012. Retrieved March 22, 2013.
Jump up ^ Collins, Gail (November 10, 2012). "Hillary's Next Move". The New York Times.
Jump up ^ "Q&A WITH VANNA WHITE". Las Vegas Magazine. September 13, 2013.
Jump up ^ Daily Mail. London http://www.dailymail.co.uk/home/you/article-2376232/Julianne-Moore-My-50s-important-years.html. Missing or empty |title= (help)
Jump up ^ "Julianne Moore Opens Up About Her Family". KatieCouric.com. Retrieved August 17, 2014.
Jump up ^ "W Network Announces Return of Home-grown Favourites in Fall 2013 Lineup". The Wall Street Journal. July 17, 2013.
Jump up ^ https://www.bizjournals.com/charlotte/bizwomen/news/latest-news/2017/05/lawsuit-against-producer-of-love-it-or-list-it.html
Jump up ^ CNN, Lisa Respers France. "'Love It or List It' homeowners file suit". CNN. Retrieved 2016-05-03.
Jump up ^ "'Love It Or List It' homeowners sue over Raleigh renovation". miamiherald. Retrieved 2016-05-03.
Jump up ^ ""Raleigh property owners settle lawsuit with 'Love It or List It'"". News & Observer. Retrieved 2017-11-26.
Jump up ^ http://www.charlotteobserver.com/news/local/article150134792.html
Jump up ^ https://www.bizjournals.com/charlotte/bizwomen/news/latest-news/2017/05/lawsuit-against-producer-of-love-it-or-list-it.html
External links[edit]
Love It or List It on HGTV
Love It or List It on W Network
Love It or List It at the Internet Movie Database
[hide]
v
t
e
Love It or List It franchise
Love It or List It
Hosts
Hilary Farr
David Visentin
Other
Episodes
Love It or List It Vancouver
Hosts
Jillian Harris
Todd Talbot
Love It or List It UK
Hosts
Kirstie Allsopp
Phil Spencer
Retrieved from "https://en.wikipedia.org/w/index.php?title=Love_It_or_List_It&oldid=834260200"
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text:
AV input
From Wikipedia, the free encyclopedia
Jump to navigation Jump to search
This article does not cite any sources. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (December 2009) (Learn how and when to remove this template message)
AV input is a common label on a connector to receive (AV) audio/visual signals from electronic equipment that generates AV signals (AV output).
These terminals are commonly found on such equipment as a television, DVD recorder or VHS recorder, and typically take input from a DVD player, a TV tuner, VHS recorder or camcorder.
Types of plugs used for video input[edit]
Composite video
RCA connector
BNC connector
UHF connector
1/8 inch minijack phone connector
S-video
DIN plug (also used for Apple Desktop Bus)
Component video
RCA connector
RGBHV
Digital video
HDMI (High Definition Multimedia Interface)
DVI (Digital Video Interface)
IEEE 1394 (FireWire)
SPDIF (Sony Philips Digital Interface)
Retrieved from "https://en.wikipedia.org/w/index.php?title=AV_input&oldid=760660360"
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