2 # bioperl module for Bio::Tools::CodonTable
4 # Please direct questions and support issues to <bioperl-l@bioperl.org>
6 # Cared for by Heikki Lehvaslaiho <heikki-at-bioperl-dot-org>
8 # Copyright Heikki Lehvaslaiho
10 # You may distribute this module under the same terms as perl itself
12 # POD documentation - main docs before the code
16 Bio::Tools::CodonTable - Codon table object
20 # This is a read-only class for all known codon tables. The IDs are
21 # the ones used by nucleotide sequence databases. All common IUPAC
22 # ambiguity codes for DNA, RNA and amino acids are recognized.
24 use Bio::Tools::CodonTable;
26 # defaults to ID 1 "Standard"
27 $myCodonTable = Bio::Tools::CodonTable->new();
28 $myCodonTable2 = Bio::Tools::CodonTable->new( -id => 3 );
34 print join (' ', "The name of the codon table no.", $myCodonTable->id(4),
35 "is:", $myCodonTable->name(), "\n");
37 # print possible codon tables
38 $tables = Bio::Tools::CodonTable->tables;
39 while ( ($id,$name) = each %{$tables} ) {
40 print "$id = $name\n";
44 $aa = $myCodonTable->translate('ACU');
45 $aa = $myCodonTable->translate('act');
46 $aa = $myCodonTable->translate('ytr');
48 # reverse translate an amino acid
49 @codons = $myCodonTable->revtranslate('A');
50 @codons = $myCodonTable->revtranslate('Ser');
51 @codons = $myCodonTable->revtranslate('Glx');
52 @codons = $myCodonTable->revtranslate('cYS', 'rna');
54 # reverse translate an entire amino acid sequence into a IUPAC
57 my $seqobj = Bio::PrimarySeq->new(-seq => 'FHGERHEL');
58 my $iupac_str = $myCodonTable->reverse_translate_all($seqobj);
61 print "Is a start\n" if $myCodonTable->is_start_codon('ATG');
62 print "Is a terminator\n" if $myCodonTable->is_ter_codon('tar');
63 print "Is a unknown\n" if $myCodonTable->is_unknown_codon('JTG');
67 Codon tables are also called translation tables or genetic codes
68 since that is what they represent. A bit more complete picture
69 of the full complexity of codon usage in various taxonomic groups
70 is presented at the NCBI Genetic Codes Home page.
72 CodonTable is a BioPerl class that knows all current translation
73 tables that are used by primary nucleotide sequence databases
74 (GenBank, EMBL and DDBJ). It provides methods to output information
75 about tables and relationships between codons and amino acids.
77 This class and its methods recognized all common IUPAC ambiguity codes
78 for DNA, RNA and animo acids. The translation method follows the
79 conventions in EMBL and TREMBL databases.
81 It is a nuisance to separate RNA and cDNA representations of nucleic
82 acid transcripts. The CodonTable object accepts codons of both type as
83 input and allows the user to set the mode for output when reverse
84 translating. Its default for output is DNA.
88 This class deals primarily with individual codons and amino
89 acids. However in the interest of speed you can L<translate>
90 longer sequence, too. The full complexity of protein translation
91 is tackled by L<Bio::PrimarySeqI::translate>.
94 The amino acid codes are IUPAC recommendations for common amino acids:
111 O Pyl Pyrrolysine (22nd amino acid)
112 U Sec Selenocysteine (21st amino acid)
118 B Asx Aspartic acid or Asparagine
119 Z Glx Glutamine or Glutamic acid
120 J Xle Isoleucine or Valine (mass spec ambiguity)
121 X Xaa Any or unknown amino acid
124 It is worth noting that, "Bacterial" codon table no. 11 produces an
125 polypeptide that is, confusingly, identical to the standard one. The
126 only differences are in available initiator codons.
129 NCBI Genetic Codes home page:
130 (Last update of the Genetic Codes: Apr. 25, 2024)
131 https://www.ncbi.nlm.nih.gov/Taxonomy/Utils/wprintgc.cgi?mode=c
133 The "value notation" / "print form" ASN.1 version is at:
134 ftp://ftp.ncbi.nih.gov/entrez/misc/data/gc.prt
136 Thanks to Matteo diTomasso for the original Perl implementation
143 User feedback is an integral part of the evolution of this and other
144 Bioperl modules. Send your comments and suggestions preferably to the
145 Bioperl mailing lists Your participation is much appreciated.
147 bioperl-l@bioperl.org - General discussion
148 http://bioperl.org/wiki/Mailing_lists - About the mailing lists
152 Please direct usage questions or support issues to the mailing list:
154 I<bioperl-l@bioperl.org>
156 rather than to the module maintainer directly. Many experienced and
157 reponsive experts will be able look at the problem and quickly
158 address it. Please include a thorough description of the problem
159 with code and data examples if at all possible.
161 =head2 Reporting Bugs
163 Report bugs to the Bioperl bug tracking system to help us keep track
164 the bugs and their resolution. Bug reports can be submitted via the
167 https://github.com/bioperl/bioperl-live/issues
169 =head1 AUTHOR - Heikki Lehvaslaiho
171 Email: heikki-at-bioperl-dot-org
175 The rest of the documentation details each of the object
176 methods. Internal methods are usually preceded with a _
180 # Let the code begin...
182 package Bio
::Tools
::CodonTable
;
186 # Object preamble - inherits from Bio::Root::Root
187 use Bio
::Tools
::IUPAC
;
190 use base
qw(Bio::Root::Root);
191 our (@NAMES, @TABLES, @STARTS, $TRCOL, $CODONS, %IUPAC_DNA, $CODONGAP, $GAP,
192 %IUPAC_AA, %THREELETTERSYMBOLS, $VALID_PROTEIN, $TERMINATOR);
195 # set internal values for all translation tables
197 use constant CODONSIZE
=> 3;
199 $CODONGAP = $GAP x CODONSIZE
;
201 # Helper private function to parse the
202 # ftp://ftp.ncbi.nih.gov/entrez/misc/data/gc.prt file which is
203 # below __DATA__ in this module (see the end of the file). This
204 # fills the @NAMES, @TABLES, and @STARTS variables. To update to
205 # a new release of gc.prt, replace the content below __DATA__.
208 # Init tables has with special option (id=0) for ATG-only start
212 ncbieaa
=> "FFLLSSSSYY**CC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
213 sncbieaa
=> "----------**--*--------------------M----------------------------",
217 while (defined(my $line = <DATA
>)) {
218 next if $line =~ /^\s*--/; # skip comment lines
219 if ($line =~ /^\s*\{\s*$/) { # start of a table description
225 if ($line =~ /^\s*(name|id|ncbieaa|sncbieaa)\s+(.+)/) {
232 # The remaining keys --- name, ncbieaa,
233 # and sncbieaa --- are strings which may
234 # be multi-line (e.g., name for table with
235 # id 4). We are assuming that there is no
236 # " character inside the value so we keep
237 # appending lines until we find an end ".
238 while ($rem !~ /^"(.*)"/ && ! eof DATA
) {
244 if ($key eq "name" && ! $name) {
245 # ignore alternative names, e.g. SGC0,
246 # only keep the first name listed.
248 } elsif ($key eq "ncbieaa") {
250 } elsif ($key eq "sncbieaa") {
255 } until (($line = <DATA
>) =~ /^\s*}\s*,?$/); # we reached the end of table description
259 sncbieaa
=> $sncbieaa
265 # print Dumper %tables;
267 my $highest_id = (sort {$a <=> $b} keys %tables)[-1];
268 for (my $i = 0; $i < $highest_id; $i++) {
269 if (defined $tables{$i}) {
270 push @NAMES, $tables{$i}->{name
};
271 push @TABLES, $tables{$i}->{ncbieaa
};
272 push @STARTS, $tables{$i}->{sncbieaa
};
284 my @nucs = qw(t c a g);
286 ($CODONS, $TRCOL) = ({}, {});
290 my $codon = "$i$j$k";
291 $CODONS->{$codon} = $x;
292 $TRCOL->{$x} = $codon;
297 %IUPAC_DNA = Bio
::Tools
::IUPAC
->iupac_iub();
298 %IUPAC_AA = Bio
::Tools
::IUPAC
->iupac_iup();
299 %THREELETTERSYMBOLS = Bio
::SeqUtils
->valid_aa(2);
300 $VALID_PROTEIN = '['.join('',Bio
::SeqUtils
->valid_aa(0)).']';
305 my($class,@args) = @_;
306 my $self = $class->SUPER::new
(@args);
309 $self->_rearrange([qw(ID
313 $id = 1 if ( ! defined ( $id ) );
314 $id && $self->id($id);
315 return $self; # success - we hope!
321 Usage : $obj->id(3); $id_integer = $obj->id();
322 Function: Sets or returns the id of the translation table. IDs are
323 integers from 0 (special ATG-only start) to 25, excluding
324 7-8 and 17-20 which have been removed. If an invalid ID is
325 given the method returns 1, the standard table.
327 Returns : value of id, a scalar, warn and fall back to 1 (standard table)
328 if specified id is not valid
329 Args : newvalue (optional)
334 my ($self,$value) = @_;
335 if( defined $value) {
336 if ( not defined $TABLES[$value] or $TABLES[$value] eq '') {
337 $self->warn("Not a valid codon table ID [$value], using [1] instead ");
340 $self->{'id'} = $value;
342 return $self->{'id'};
349 Function: returns the descriptive name of the translation table
360 my ($id) = $self->{'id'};
367 Usage : $obj->tables() or Bio::Tools::CodonTable->tables()
368 Function: returns a hash reference where each key is a valid codon
369 table id() number, and each value is the corresponding
370 codon table name() string
380 for my $id (0 .. $#NAMES) {
381 my $name = $NAMES[$id];
382 $tables{$id} = $name if $name;
390 Usage : $obj->translate('YTR')
391 Function: Returns a string of one letter amino acid codes from
392 nucleotide sequence input. The imput can be of any length.
394 Returns 'X' for unknown codons and codons that code for
395 more than one amino acid. Returns an empty string if input
396 is not three characters long. Exceptions for these are:
398 - IUPAC amino acid code B for Aspartic Acid and
400 - IUPAC amino acid code Z for Glutamic Acid, Glutamine is
402 - if the codon is two nucleotides long and if by adding
403 an a third character 'N', it codes for a single amino
404 acid (with exceptions above), return that, otherwise
407 Returns empty string for other input strings that are not
408 three characters long.
411 Returns : a string of one letter ambiguous IUPAC amino acid codes
412 Args : ambiguous IUPAC nucleotide string
418 my ($self, $seq, $complete_codon) = @_;
419 $self->throw("Calling translate without a seq argument!") unless defined $seq;
420 return '' unless $seq;
424 $partial = 2 if length($seq) % CODONSIZE
== 2;
429 if ($seq =~ /[^actg]/ ) { #ambiguous chars
430 for (my $i = 0; $i < (length($seq) - (CODONSIZE
-1)); $i+= CODONSIZE
) {
431 my $triplet = substr($seq, $i, CODONSIZE
);
432 if( $triplet eq $CODONGAP ) {
434 } elsif (exists $CODONS->{$triplet}) {
435 $protein .= substr($TABLES[$id],
436 $CODONS->{$triplet},1);
438 $protein .= $self->_translate_ambiguous_codon($triplet);
441 } else { # simple, strict translation
442 for (my $i = 0; $i < (length($seq) - (CODONSIZE
-1)); $i+=CODONSIZE
) {
443 my $triplet = substr($seq, $i, CODONSIZE
);
444 if( $triplet eq $CODONGAP ) {
447 if (exists $CODONS->{$triplet}) {
448 $protein .= substr($TABLES[$id], $CODONS->{$triplet}, 1);
454 if ($partial == 2 && $complete_codon) { # 2 overhanging nucleotides
455 my $triplet = substr($seq, ($partial -4)). "n";
456 if( $triplet eq $CODONGAP ) {
458 } elsif (exists $CODONS->{$triplet}) {
459 my $aa = substr($TABLES[$id], $CODONS->{$triplet},1);
462 $protein .= $self->_translate_ambiguous_codon($triplet, $partial);
468 sub _translate_ambiguous_codon
{
469 my ($self, $triplet, $partial) = @_;
473 my @codons = $self->unambiguous_codons($triplet);
475 foreach my $codon (@codons) {
476 $aas{substr($TABLES[$id],$CODONS->{$codon},1)} = 1;
478 my $count = scalar keys %aas;
480 $aa = (keys %aas)[0];
482 elsif ( $count == 2 ) {
483 if ($aas{'D'} and $aas{'N'}) {
486 elsif ($aas{'E'} and $aas{'Q'}) {
489 $partial ?
($aa = '') : ($aa = 'X');
492 $partial ?
($aa = '') : ($aa = 'X');
497 =head2 translate_strict
499 Title : translate_strict
500 Usage : $obj->translate_strict('ACT')
501 Function: returns one letter amino acid code for a codon input
503 Fast and simple translation. User is responsible to resolve
504 ambiguous nucleotide codes before calling this
505 method. Returns 'X' for unknown codons and an empty string
506 for input strings that are not three characters long.
508 It is not recommended to use this method in a production
509 environment. Use method translate, instead.
513 Args : a codon = a three nucleotide character string
518 sub translate_strict
{
519 my ($self, $value) = @_;
520 my $id = $self->{'id'};
525 return '' unless length $value == 3;
527 return 'X' unless defined $CODONS->{$value};
529 return substr( $TABLES[$id], $CODONS->{$value}, 1 );
535 Usage : $obj->revtranslate('G')
536 Function: returns codons for an amino acid
538 Returns an empty string for unknown amino acid
539 codes. Ambiguous IUPAC codes Asx,B, (Asp,D; Asn,N) and
540 Glx,Z (Glu,E; Gln,Q) are resolved. Both single and three
541 letter amino acid codes are accepted. '*' and 'Ter' are
544 By default, the output codons are shown in DNA. If the
545 output is needed in RNA (tr/t/u/), add a second argument
548 Example : $obj->revtranslate('Gly', 'RNA')
549 Returns : An array of three lower case letter strings i.e. codons
550 Args : amino acid, 'RNA'
555 my ($self, $value, $coding) = @_;
558 if (length($value) == 3 ) {
560 $value = ucfirst $value;
561 $value = $THREELETTERSYMBOLS{$value};
563 if ( defined $value and $value =~ /$VALID_PROTEIN/
564 and length($value) == 1
566 my $id = $self->{'id'};
569 my @aas = @
{$IUPAC_AA{$value}};
570 foreach my $aa (@aas) {
572 $aa = '\*' if $aa eq '*';
573 while ($TABLES[$id] =~ m/$aa/g) {
574 my $p = pos $TABLES[$id];
575 push (@codons, $TRCOL->{--$p});
580 if ($coding and uc ($coding) eq 'RNA') {
581 for my $i (0..$#codons) {
582 $codons[$i] =~ tr/t/u/;
589 =head2 reverse_translate_all
591 Title : reverse_translate_all
592 Usage : my $iup_str = $cttable->reverse_translate_all($seq_object)
593 my $iup_str = $cttable->reverse_translate_all($seq_object,
596 Function: reverse translates a protein sequence into IUPAC nucleotide
597 sequence. An 'X' in the protein sequence is converted to 'NNN'
598 in the nucleotide sequence.
600 Args : a Bio::PrimarySeqI compatible object (mandatory)
601 a Bio::CodonUsage::Table object and a threshold if only
602 codons with a relative frequency above the threshold are
606 sub reverse_translate_all
{
607 my ($self, $obj, $cut, $threshold) = @_;
611 if (!$obj || !$obj->isa('Bio::PrimarySeqI')){
612 $self->throw(" I need a Bio::PrimarySeqI object, not a [".
615 if($obj->alphabet ne 'protein') {
616 $self->throw("Cannot reverse translate, need an amino acid sequence .".
617 "This sequence is of type [" . $obj->alphabet ."]");
620 my @seq = split '', $obj->seq;
622 ## if we're not supplying a codon usage table...
623 if( !$cut && !$threshold) {
624 ## get lists of possible codons for each aa.
627 push @data, (['NNN']);
629 my @cods = $self->revtranslate($aa);
634 #else we are supplying a codon usage table, we just want common codons
636 if(!$cut->isa('Bio::CodonUsage::Table')) {
637 $self->throw("I need a Bio::CodonUsage::Table object, not a [".
640 my $cod_ref = $cut->probable_codons($threshold);
643 push @data, (['NNN']);
646 push @data, $cod_ref->{$aa};
650 return $self->_make_iupac_string(\
@data);
653 =head2 reverse_translate_best
655 Title : reverse_translate_best
656 Usage : my $str = $cttable->reverse_translate_best($seq_object,$cutable);
657 Function: Reverse translates a protein sequence into plain nucleotide
658 sequence (GATC), uses the most common codon for each amino acid
660 Args : A Bio::PrimarySeqI compatible object and a Bio::CodonUsage::Table object
664 sub reverse_translate_best
{
666 my ($self, $obj, $cut) = @_;
668 if (!$obj || !$obj->isa('Bio::PrimarySeqI')){
669 $self->throw(" I need a Bio::PrimarySeqI object, not a [".
672 if ($obj->alphabet ne 'protein') {
673 $self->throw("Cannot reverse translate, need an amino acid sequence .".
674 "This sequence is of type [" . $obj->alphabet ."]");
676 if ( !$cut | !$cut->isa('Bio::CodonUsage::Table')) {
677 $self->throw("I need a Bio::CodonUsage::Table object, not a [".
682 my @seq = split '', $obj->seq;
684 my $cod_ref = $cut->most_common_codons();
686 for my $aa ( @seq ) {
691 if ( defined $cod_ref->{$aa} ) {
692 $str .= $cod_ref->{$aa};
694 $self->throw("Input sequence contains invalid character: $aa");
700 =head2 is_start_codon
702 Title : is_start_codon
703 Usage : $obj->is_start_codon('ATG')
704 Function: returns true (1) for all codons that can be used as a
705 translation start, false (0) for others.
706 Example : $myCodonTable->is_start_codon('ATG')
713 shift->_codon_is( shift, \
@STARTS, 'M' );
719 Usage : $obj->is_ter_codon('GAA')
720 Function: returns true (1) for all codons that can be used as a
721 translation tarminator, false (0) for others.
722 Example : $myCodonTable->is_ter_codon('ATG')
729 my ($self, $value) = @_;
730 my $id = $self->{'id'};
732 # We need to ensure U is mapped to T (ie. UAG)
736 if (length $value != 3 ) {
737 # Incomplete codons are not stop codons
742 # For all the possible codons, if any are not a stop
743 # codon, fail immediately
744 for my $c ( $self->unambiguous_codons($value) ) {
745 my $m = substr( $TABLES[$id], $CODONS->{$c}, 1 );
746 if($m eq $TERMINATOR) {
756 # desc: compares the passed value with a single entry in the given
758 # args: a value (typically a three-char string like 'atg'),
759 # a reference to the appropriate set of codon tables,
760 # a single-character value to check for at the position in the
762 # ret: boolean, true if the given codon table contains the $key at the
763 # position corresponding to $value
765 my ($self, $value, $table, $key ) = @_;
767 return 0 unless length $value == 3;
772 my $id = $self->{'id'};
773 for my $c ( $self->unambiguous_codons($value) ) {
774 my $m = substr( $table->[$id], $CODONS->{$c}, 1 );
775 if ($m eq $key) { return 1; }
780 =head2 is_unknown_codon
782 Title : is_unknown_codon
783 Usage : $obj->is_unknown_codon('GAJ')
784 Function: returns false (0) for all codons that are valid,
786 Example : $myCodonTable->is_unknown_codon('NTG')
793 sub is_unknown_codon
{
794 my ($self, $value) = @_;
797 return 1 unless $self->unambiguous_codons($value);
801 =head2 unambiguous_codons
803 Title : unambiguous_codons
804 Usage : @codons = $self->unambiguous_codons('ACN')
805 Returns : array of strings (one-letter unambiguous amino acid codes)
806 Args : a codon = a three IUPAC nucleotide character string
810 sub unambiguous_codons
{
811 my ($self,$value) = @_;
812 my @nts = map { $IUPAC_DNA{uc $_} } split(//, $value);
815 for my $i ( @
{$nts[0]} ) {
816 for my $j ( @
{$nts[1]} ) {
817 for my $k ( @
{$nts[2]} ) {
818 push @codons, lc "$i$j$k";
823 =head2 _unambiquous_codons
825 deprecated, now an alias for unambiguous_codons
829 sub _unambiquous_codons
{
830 unambiguous_codons
( undef, @_ );
836 Usage : $newid = $ct->add_table($name, $table, $starts)
837 Function: Add a custom Codon Table into the object.
838 Know what you are doing, only the length of
839 the argument strings is checked!
840 Returns : the id of the new codon table
841 Args : name, a string, optional (can be empty)
842 table, a string of 64 characters
843 startcodons, a string of 64 characters, defaults to standard
848 my ($self, $name, $table, $starts) = @_;
850 $name ||= 'Custom' . $#NAMES + 1;
851 $starts ||= $STARTS[1];
852 $self->throw('Suspect input!')
853 unless length($table) == 64 and length($starts) == 64;
856 push @TABLES, $table;
857 push @STARTS, $starts;
862 sub _make_iupac_string
{
863 my ($self, $cod_ref) = @_;
864 if(ref($cod_ref) ne 'ARRAY') {
865 $self->throw(" I need a reference to a list of references to codons, ".
866 " not a [". ref($cod_ref) . "].");
868 my %iupac_hash = Bio
::Tools
::IUPAC
->iupac_rev_iub();
869 my $iupac_string = ''; ## the string to be returned
870 for my $aa (@
$cod_ref) {
872 ## scan through codon positions, record the differing values,
873 # then look up in the iub hash
874 for my $index(0..2) {
876 map { my $k = substr($_,$index,1);
877 $h{$k} = undef;} @
$aa;
878 my $lookup_key = join '', sort{$a cmp $b}keys %h;
881 $iupac_string .= $iupac_hash{uc$lookup_key};
884 return $iupac_string;
890 # Follows the content of
891 # ftp://ftp.ncbi.nih.gov/entrez/misc/data/gc.prt, which is the NCBI
892 # genetic codon table in ASN.1 value notation / print format. We do
893 # not have a ASN.1 decoder for value notation but it's easy enough to
897 --**************************************************************************
898 -- This is the NCBI genetic code table
899 -- Initial base data set from Andrzej Elzanowski
while at PIR International
900 -- Addition of Eubacterial
and Alternative Yeast by J
.Ostell at NCBI
901 -- Base
1-3 of
each codon have been added as comments to facilitate
902 -- readability at the suggestion of Peter Rice
, EMBL
903 -- Later additions by Taxonomy Group staff at NCBI
906 -- Renamed genetic code
24 to Rhabdopleuridae Mitochondrial
909 -- Added Cephalodiscidae mitochondrial genetic code
33
912 -- Added GTG as start codon
for genetic code
3
913 -- Added Balanophoraceae plastid genetic code
32
916 -- Change to CTG
-> Leu
in genetic codes
27, 28, 29, 30
919 -- Added Karyorelict nuclear genetic code
27
920 -- Added Condylostoma nuclear genetic code
28
921 -- Added Mesodinium nuclear genetic code
29
922 -- Added Peritrich nuclear genetic code
30
923 -- Added Blastocrithidia nuclear genetic code
31
926 -- Added Pachysolen tannophilus nuclear genetic code
26
929 -- Updated version to reflect numerous undocumented changes
:
930 -- Corrected start codons
for genetic code
25
931 -- Name of new genetic code is Candidate Division SR1
and Gracilibacteria
932 -- Added candidate division SR1 nuclear genetic code
25
933 -- Added GTG as start codon
for genetic code
24
934 -- Corrected Pterobranchia Mitochondrial genetic code
(24)
935 -- Added genetic code
24, Pterobranchia Mitochondrial
936 -- Genetic code
11 is now Bacterial
, Archaeal
and Plant Plastid
937 -- Fixed capitalization of mitochondrial
in codes
22 and 23
938 -- Added GTG
, ATA
, and TTG as alternative start codons to code
13
941 -- Code
14 differs from code
9 only by translating UAA to Tyr rather than
942 -- STOP
. A recent
study (Telford et al
, 2000) has found
no evidence that
943 -- the codon UAA codes
for Tyr
in the flatworms
, but other opinions exist
.
944 -- There are very few GenBank records that are translated with code
14,
945 -- but a test translation shows that retranslating these records with code
946 -- 9 can cause premature terminations
. Therefore
, GenBank will maintain
947 -- code
14 until further information becomes available
.
950 -- Added GTG start to Echinoderm mitochondrial code
, code
9
953 -- Added code
23 Thraustochytrium mitochondrial code
954 -- formerly OGMP code
93
955 -- submitted by Gertraude Berger
, Ph
.D
.
958 -- Added code
22 TAG
-Leu
, TCA
-stop
959 -- found
in mitochondrial DNA of Scenedesmus obliquus
960 -- submitted by Gertraude Berger
, Ph
.D
.
961 -- Organelle Genome Megasequencing Program
, Univ Montreal
964 -- Added code
21, Trematode Mitochondrial
965 -- (as deduced from
: Garey
& Wolstenholme
,1989; Ohama et al
, 1990)
966 -- Added code
16, Chlorophycean Mitochondrial
967 -- (TAG can translated to Leucine instaed to STOP
in chlorophyceans
971 -- Added CTG
,TTG as allowed alternate start codons
in Standard code
.
972 -- Prats et al
. 1989, Hann et al
. 1992
974 -- Version
3.3 - 10/13/95
975 -- Added alternate intiation codon ATC to code
5
976 -- based on complete mitochondrial genome of honeybee
977 -- Crozier
and Crozier
(1993)
979 -- Version
3.2 - 6/24/95
981 -- 10 Alternative Ciliate Macronuclear renamed to Euplotid Macro
...
982 -- 15 Blepharisma Macro
.. code added
983 -- 5 Invertebrate Mito
.. GTG allowed as alternate initiator
984 -- 11 Eubacterial renamed to Bacterial as most alternate starts
985 -- have been found
in Archea
988 -- Version
3.1 - 1995
989 -- Updated as per Andrzej Elzanowski at NCBI
990 -- Complete documentation
in NCBI toolkit documentation
991 -- Note
: 2 genetic codes have been deleted
993 -- Old id Use id
- Notes
995 -- id
7 id
4 - Kinetoplast code now merged
in code id
4
996 -- id
8 id
1 - all plant chloroplast differences due to RNA edit
999 --*************************************************************************
1001 Genetic
-code
-table
::= {
1006 ncbieaa
"FFLLSSSSYY**CC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1007 sncbieaa
"---M------**--*----M---------------M----------------------------"
1008 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1009 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1010 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1013 name
"Vertebrate Mitochondrial" ,
1016 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIMMTTTTNNKKSS**VVVVAAAADDEEGGGG",
1017 sncbieaa
"----------**--------------------MMMM----------**---M------------"
1018 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1019 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1020 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1023 name
"Yeast Mitochondrial" ,
1026 ncbieaa
"FFLLSSSSYY**CCWWTTTTPPPPHHQQRRRRIIMMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1027 sncbieaa
"----------**----------------------MM---------------M------------"
1028 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1029 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1030 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1033 name
"Mold Mitochondrial; Protozoan Mitochondrial; Coelenterate
1034 Mitochondrial; Mycoplasma; Spiroplasma" ,
1037 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1038 sncbieaa
"--MM------**-------M------------MMMM---------------M------------"
1039 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1040 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1041 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1044 name
"Invertebrate Mitochondrial" ,
1047 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIMMTTTTNNKKSSSSVVVVAAAADDEEGGGG",
1048 sncbieaa
"---M------**--------------------MMMM---------------M------------"
1049 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1050 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1051 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1054 name
"Ciliate Nuclear; Dasycladacean Nuclear; Hexamita Nuclear" ,
1057 ncbieaa
"FFLLSSSSYYQQCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1058 sncbieaa
"--------------*--------------------M----------------------------"
1059 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1060 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1061 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1064 name
"Echinoderm Mitochondrial; Flatworm Mitochondrial" ,
1067 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIIMTTTTNNNKSSSSVVVVAAAADDEEGGGG",
1068 sncbieaa
"----------**-----------------------M---------------M------------"
1069 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1070 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1071 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1074 name
"Euplotid Nuclear" ,
1077 ncbieaa
"FFLLSSSSYY**CCCWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1078 sncbieaa
"----------**-----------------------M----------------------------"
1079 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1080 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1081 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1084 name
"Bacterial, Archaeal and Plant Plastid" ,
1086 ncbieaa
"FFLLSSSSYY**CC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1087 sncbieaa
"---M------**--*----M------------MMMM---------------M------------"
1088 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1089 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1090 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1093 name
"Alternative Yeast Nuclear" ,
1095 ncbieaa
"FFLLSSSSYY**CC*WLLLSPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1096 sncbieaa
"----------**--*----M---------------M----------------------------"
1097 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1098 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1099 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1102 name
"Ascidian Mitochondrial" ,
1104 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIMMTTTTNNKKSSGGVVVVAAAADDEEGGGG",
1105 sncbieaa
"---M------**----------------------MM---------------M------------"
1106 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1107 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1108 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1111 name
"Alternative Flatworm Mitochondrial" ,
1113 ncbieaa
"FFLLSSSSYYY*CCWWLLLLPPPPHHQQRRRRIIIMTTTTNNNKSSSSVVVVAAAADDEEGGGG",
1114 sncbieaa
"-----------*-----------------------M----------------------------"
1115 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1116 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1117 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1120 name
"Blepharisma Macronuclear" ,
1122 ncbieaa
"FFLLSSSSYY*QCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1123 sncbieaa
"----------*---*--------------------M----------------------------"
1124 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1125 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1126 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1129 name
"Chlorophycean Mitochondrial" ,
1131 ncbieaa
"FFLLSSSSYY*LCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1132 sncbieaa
"----------*---*--------------------M----------------------------"
1133 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1134 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1135 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1138 name
"Trematode Mitochondrial" ,
1140 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIMMTTTTNNNKSSSSVVVVAAAADDEEGGGG",
1141 sncbieaa
"----------**-----------------------M---------------M------------"
1142 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1143 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1144 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1147 name
"Scenedesmus obliquus Mitochondrial" ,
1149 ncbieaa
"FFLLSS*SYY*LCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1150 sncbieaa
"------*---*---*--------------------M----------------------------"
1151 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1152 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1153 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1156 name
"Thraustochytrium Mitochondrial" ,
1158 ncbieaa
"FF*LSSSSYY**CC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1159 sncbieaa
"--*-------**--*-----------------M--M---------------M------------"
1160 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1161 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1162 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1165 name
"Rhabdopleuridae Mitochondrial" ,
1167 ncbieaa
"FFLLSSSSYY**CCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSSKVVVVAAAADDEEGGGG",
1168 sncbieaa
"---M------**-------M---------------M---------------M------------"
1169 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1170 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1171 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1174 name
"Candidate Division SR1 and Gracilibacteria" ,
1176 ncbieaa
"FFLLSSSSYY**CCGWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1177 sncbieaa
"---M------**-----------------------M---------------M------------"
1178 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1179 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1180 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1183 name
"Pachysolen tannophilus Nuclear" ,
1185 ncbieaa
"FFLLSSSSYY**CC*WLLLAPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1186 sncbieaa
"----------**--*----M---------------M----------------------------"
1187 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1188 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1189 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1192 name
"Karyorelict Nuclear" ,
1194 ncbieaa
"FFLLSSSSYYQQCCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1195 sncbieaa
"--------------*--------------------M----------------------------"
1196 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1197 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1198 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1201 name
"Condylostoma Nuclear" ,
1203 ncbieaa
"FFLLSSSSYYQQCCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1204 sncbieaa
"----------**--*--------------------M----------------------------"
1205 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1206 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1207 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1210 name
"Mesodinium Nuclear" ,
1212 ncbieaa
"FFLLSSSSYYYYCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1213 sncbieaa
"--------------*--------------------M----------------------------"
1214 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1215 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1216 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1219 name
"Peritrich Nuclear" ,
1221 ncbieaa
"FFLLSSSSYYEECC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1222 sncbieaa
"--------------*--------------------M----------------------------"
1223 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1224 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1225 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1228 name
"Blastocrithidia Nuclear" ,
1230 ncbieaa
"FFLLSSSSYYEECCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1231 sncbieaa
"----------**-----------------------M----------------------------"
1232 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1233 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1234 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1237 name
"Balanophoraceae Plastid" ,
1239 ncbieaa
"FFLLSSSSYY*WCC*WLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSRRVVVVAAAADDEEGGGG",
1240 sncbieaa
"---M------*---*----M------------MMMM---------------M------------"
1241 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1242 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1243 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG
1246 name
"Cephalodiscidae Mitochondrial" ,
1248 ncbieaa
"FFLLSSSSYYY*CCWWLLLLPPPPHHQQRRRRIIIMTTTTNNKKSSSKVVVVAAAADDEEGGGG",
1249 sncbieaa
"---M-------*-------M---------------M---------------M------------"
1250 -- Base1 TTTTTTTTTTTTTTTTCCCCCCCCCCCCCCCCAAAAAAAAAAAAAAAAGGGGGGGGGGGGGGGG
1251 -- Base2 TTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGGTTTTCCCCAAAAGGGG
1252 -- Base3 TCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAGTCAG