.4iieri;v 1995: .^(1. Primed in Beigiu,

ALLERGY

Special report

Allergen nomenclature Te Piao King, Hoffman D, Lowenstem H, Marsh DG. Platts-Mills TAE. Thomas W. AUereen nomenclature. .Allergy 1995: 50; "765-774. © Munksgaard 1995.

T. P. King, D. Hoffman, H. L0wenstein, D. G. Marsh, T. A. E. Platts-Mills, W. Thomas Accepted for publication 2 September 1994

A. Introduction Rapid advances have been made in the past few years in allergen characterization and sequence determination by chemical and molecular biological approaches. This is indicated by the hst of allergens with known partial or complete amino acid sequences in Table 1. A number of other important allergens are known in addition to those in Table 1 but their sequences are as yet not known. A useful source for known allergens is the Allergen Database (ALBE) m which are compiled their known biochemical and immunological properties together with their sequence data if known (1). To take into account these advances, a revision of the current allergen nomenclature system (2) is given below. As in the current nomenclature system, the proposed revisions are for allergens which induce IgE-mediated {atopic) allergy in humans. In addition to the expected thorough immunochemieal characterization of any newly discovered allergen, investigators are urged to obtain partial, or preferably complete, sequence data before using the official nomenclature system. Also it is expected that investigators would screen a reasonable population size so as to establish the frequency of response in patients to the newly discovered allergens.** Investigators frequently refer to allergens as major or minor ones. The generally accepted meaning of this terminology is that an allergen is designated as either major or minor depending on whether greater or less than 50'\, of patients tested have the corresponding allergen-specific igEs (cf. 3-5). The revised nomenclature for allergens is given * This document has appeal in WHO Bulletin Vol. 72. August 1994. ** Investigators are invited to consult our committee memhers for assigning an allergen numher hefore publication so as to avoid duplication.

below together with the proposed nomenclatures for (a) allergen genes. niRNAs and cDNAs and (b) recombinant and synthetic peptides of allergenic interest,

B. Revised allergen nomenclature /. Allergens Allergens are designated according to the accepted taxonomic name of their source as follows: the first three letters of the genus, space, the first letter of the species, space, and an Arabic number. The numbers are assigned to the allergens in the order of their identification, and the same number is generally used to designate homologous allergens of related species. As two examples, Eol p 1 refers to the first pollen allergen identified from Lolium perenne ryegrass, and Cyn d 1 refers to the homologous pollen allergen from Cynodon dactylon, Bermuda grass. In some instances, the above system of the first 3 letters of a genus and the first letter of a species has to be modified to include an additional letter for designation of the exact genus or species. For example. 4 of the many vespids which can cause insect allerg} are Vespuki vulgaris, Vespuki vidua, Vespula consohrina and Vespa craho. The homologous major \enom allergens, antigen 5s. from Vespida vulgaris and Vespula vidua are both designated as Ves v 5 in the existing nomenclature, and those from Vespula consohrina and Vespa crabo are designated as Ves c 5. To avoid these ambiguities, antigen 5s from Vespula vulgaris and Vespula vidua will be designated as Ves V 5 and Ves vi 5 respectively, and those from Vespula consobrina and Vespa craho as Ves c 5 and Vesp c 5. In the examples given, the modified nomenclature is used for the allergens from Vespula vidua and Vespa craho, as the allergens from Vespula vulgaris were characterized prior to those for Vespula vidua and Vespa craho. 765

Te Piao King et a!. Another example is for allergens from the domestic dog iCanis domesticus) and the mold Candida alhicans. TO avoid ambiguity, the modified system is used to designate Can d and Cand a allergens from these two sources respectively. In the current nomenclature system (2) the letters arc italicized and the numerals are Roman numerals. In the revised system, only letters of normal type and Arabic numbers are used. The proposed changes conform to the accepted nomenclature used in baclerial genetics (6) and the HLA system (7) in that ltalici/ed and normal characters are used to represent genotypes and phenotypes respectively. 2. Allergens and isoallerirens

same allergen group are designated by suffixes of a period followed by four Arabic numerals. The first two numerals 01 to 99 refer to a particular isoallergen, and the two subsequent numerals 01 to 99 refer to a particular variant of a particular isoallergen designated by the preceding two numerals. In cases where there is only one known isoallergen but there are several variants, the system of a suffix of 4 numerals will still apply. These numerals will be chosen in the order of the identification of allergens and/or their cDN As irrespective of the physicochemical properties of the allergens. In cases of silent mutations, there can be more than one suffix of 4 numerals designating the same allergen and in that case the suffix with the lowest number will be used

An allergen from a smde spec.es mav consist of to desigriate the allergen of interest, several closelv similar molecules. These'similar mol. ^\^ ^ ^ ^ ' ^ ' ° " of suffixes of 4 numerals to designate ecules are des"iunated as isoallergens when thev share '^^allergens and their variants will permit their unthe following ^common biochemical properties: a. ^"^^iguous designation. In many cases it is unnesimilar molecular size: b. identical biological func"'''^''^"•>^ ^^ ^P^^^^> ^^^ isoallergen or variant and the t.on, if known, e.g. enzymatic action; and c. 67" corresponding suffixes may be deleted; e.g. Bet v 1 ident.tv of am.nolic.d sequences. ' represents any Bet v 1 allergen and Bet v 1.0101 It is'recognized thai the recommended 67",. seLepresents variant number 1 of isoallergen Bet v 1. quencc idcnlitv for 2 allergens to be assigned to the ^''•'' '^^^^' examples of this nomenclature system are same group is cMily A guide. There are likelv to be 2'^^" f^Z\ . • . u , i borderline cases. As an example, the ragweed alterP " ^ ^ ^ ' ^ ^'«"'"g:. ^"^^^^^ ^. ^^^^^^^'^^^ "^"l^^Pl^ uens Amb a 1 and 2 share 65"., am.no acid sequence P^lvniorph.c forms which differ from each other by Identity (8). These allergens were assigned to differ^""e ";' ' " \ ' sequences (8). Four such forms of ent groups because of tlie.r ditlerent immunochem'"^"'^f ^ '"'^ ^"^'!''" ^^"^ ^^^> ^^^ designated as Amb .carpropert.es befo.-e the.r sequences were known. '' ^'l^' ]-^^- 1.03 and 1.04 Lach isoallergen of the Alle.-ge.is fro.r. different spec.es of the same or '^"^^ '' ^ ^'"^^"P '^ ^^^^"^ ^« ^^^^'^ ^ ^"^ ^ ''^^•^"^^ ^"^ diilerenl uenus which share the above-ment.oned 97",, of sequence identity. These variants of the Amb common biochemical p.opert.es a.-e considered to '^ lf\ -[^'^^P ' ' ' ' " ^^ designated as Amb a 1.0101, bclonu to tlie same u.'oup and the sequence identitv . *^^^' requ.i-e.iient ca.i he < 67",, as .s the case for alle.'" ''^'"^'"f ^ ^° ^^"^^ ^ ^' ^^"'-^ ^"^ ^^''"^^ ^^ ^"^'^

uensofthesamcspec.es. For example, Amb a 5 and

^ " ^^'''''' ^^'^"^ '''' ""^^"^ ^^'^^"^"g- '^^^^^ ^o™^

A.iib t 5 l.-om shoi-t and mant ragweed pollens have about 45 "„ sequence .dentitv (9) \ i n d also have s.ni.lar te.-t.a.-y struclu.-es (10, 11). Another example is the .uiniM- pt">ilen allc^gens A m b a 10, Poa p 10 and Foi p 10 fro.11 ragweed (12), Kentucky bluegrass (13) and .-yegrass (14). Although their sequences are not know n, the\ cU'c assiuned to the sa.ne allertzen croup as the> clearK hax eUic same b.olouical functfon of cviochriMiie c "

^^*^"^'"'" ^'^'^^ polymorphic sites and they have > 99"^ ' " " ' " ^ '^^"'^ sequence identity. They are designated ''' '^"^^ '' -^^^^^^ ^ " ^ ~-^^^--

^. hoallcriicns and varianis cDNA cloning ol" alle.-gens often show s nucleotide mutations wh.ch are either silent or which can lead to single or multiple a.nino acid substitutions. In the re\ised system, members of an allergen group which ha\e >67",, amino acid sequence identit\ are designated as isoallergens. Fach isoallergen may have multiple forms of closely similar sequences, which a.-e designated as va.-iants. isoailergens and their variants belonging to the 766

C. Nomenclature for allergen genes, mRNAS and cDNAs '^^ P'"".^"^ ^"^^ genomic structures of allergens are known in at least two cases; cat allergen Fel d 1 and mouse urinaiy allergen Mus m 1. Knowledge of the genomic structure can provide an understanding of how the different polymorphic forms are generated by differential splicing and or exon usage. By adopting the revised nomenclature for allergens, we can reserve the italicized characters to designate an allergen gene. Normal characters are used for designation of mRNAs and cDNAs. For example, Fel d 1 is a protein consisting of two polypeptide chains (15) which are encoded by two separate genes (16). The two allergen genes for chains 1 and 2 of Fe! d 1 will be represented by Fel

Allergen nomenclature d lA and Fel d IB respectively. Allelic forms of mRNAs and cDNAs of the Fel d lA gene are designated as mRNA or cDNA Fel d lA.OlOl where the numerals are to correspond to those of the polymorphic allergens.

D. Nomenclature for recombinant and synthetic peptides of allergenic interest There is interest in the possible use of fragments of allergens as reagents to modulate allergen-specific immune responses. Such fragments may be prepared by recombinant technology or by chemical synthesis. Therefore it is useful to establish a generally accepted nomenclature for such peptides of allergenic interest. The nomenclature for recombinant and synthetic peptides of allergenic interest is to be based on the nomenclature for naturally occurring allergens since it is well accepted by the scientific and the clinical communities. An allergen which is prepared by recombinant (r) or chemical synthetic (s) means is to be differentiated from a natural (n) allergen by the addition of the prefix of r or s followed by a suffix of the amino acid residue positions which are given in parenthesis. For example, a recombinant hornet venom allergen Dol m 5.0201 which contains the entire sequence of 204 residues will be designated as rDol m 5.0201, and a recombinant or synthetic peptide of residue 151-165 of Dol m 5.0201 will be designated as r or sDol m 5.0201 (151-165) respectively. Natural allergens may contain post-translational modifications as many proteins do. These modifications include glycosylation, acylation, methylation, etc. Recombinant or synthetic allergen, designated by the prefix r or s, is taken to indicate that it does not contain the post-translational modification of the natural allergen. If the recombinant or synthetic allergen does contain the exact same modification as that of the natural allergen, it will be designated by a prefix of rn or sn. For example, the honeybee venom allergen phospholipase A., Api m 1, is a glycoprotein with an oligosaccharide attached at the asparagine residue number 13 (17, 18). A synthetic peptide of residues 1-20 of Api m 1 without the oligosaccharide at residue 13 will be designated as sApi m 1

(1-20), and a synthetic peptide of residues 1-20 with the exact same oligosaccharide of the natural allergen will be designated as snApi m 1 (1-20). For recombinant or synthetic fragments which are derivatives of sequences contained w ithin the native allergen structure, an additional suffix enclosed in square brackets will be used to indicate that the peptide referred to is an analog. Substitutions or modifications of amino acid residues are given w ith the standard one-letter code and superscript numbers indicate the residue positions at which modifications occur. The one-letter codes for L-amino acids are given in upper-case letters and those for D-amino acids are in lower-case letters. The modifications, which can be substitution, insertion or deletion, are specified in parenthesis within the brackets. Obviously when there are many changes, it will not be practical to follow this nomenclature but to give the fully modified sequences. Examples of these analogs of sDol m 5.0201 (151-165) are given belowUnmodified: sDol m 5.0201 (151-165) Substitution: sDol m 5.0201 (151-165) [K " ' ] - L-lysine residue at position 153 of sDol m 5.0201 (151-165) is substituted with D-lysine Insertion: sDol m 5.0201 (151-165) [ + K " ' ] - one residue of L-lysine is inserted between positions 153-154 Deletion: sDol m 5.0201 (151-165) [ - K ' ' ' ] - L-lysine residue at position 153 is deleted N-terminal modification: sDol m 5.0201 (151165) [N-Ac] - N-terminal amino group is acetylated C-terminal modification: sDol m 5.0201 (151165) [C-NH,] - C-terminal carboxyl group is in the form of carboxamide The nomenclature proposed above is very similar to that used for synthetic peptides representative of immunoglobulin sequences (19).

Acknowledgments We Ihank Drs, Martir Chapman, irwin Gntfith. Shayam Mohaptra. Rosalia Rodri Licz and Alec SCIKMT for rcxiewing this document.

767

1 e Fiao King et a!. Tdble '

Sjme dl'ergens with known sequences Allergens: systematic

Sequenc

A. Weed pollei As'.erales imb a 1; antigen E

8, 20 8, 21

^mb a 2: antigen K ib a 3: Ra3

11,23 24,25 26 27

ib a 5: Ra5 ib a 6: Ra6 *mb a 7: Ra7 ib a^ Ambrosia tritida fgiant ragweed!

9, 10.

Artemisia vulgaris (mugwort)

29

B. Grass pollens Poales

Dactylis glomer (orchard (irass)

Dace! 1:AgDg

32

Dace1 2 Dace

33

Lolp LOl

1: gro up gro up

p

Loip Lol p

34

3' 5

gro:upl

Lol p g- Lol p li

31 31 35

35 36 37 38 34 39 40. 41 42 43 44 45

C. Tree pollei

Cor a 1

Que a 1

Crv

55,56 57,58

768

Allergen nomenclature Table 1. Continued, Sequence

Allergens; systematic Allergen source

and original names

(kDa!

data

Juniper sabinoides (mountain cedar)

Juns 1

50

58

Junv 1

45-50

58

Olee 1

16

Juniper virginiana (eastern red cedar)

Olea europaea (olive) D. Mites Dermatophagoides pteronyssinus Derp 1: antigen P, Derp 2

62

Der p 3: trypsin

63

Der p 4: amylase

64 65

Derp 5 Der p 6; chymotrypsin Der p 7

25 22-28

Dermatophagoides microceras (mite) Dermatophagoides farinae (mite)

Derf 1

69

Derf 2

70, 71

Derf 3

72

Lepidoglyphus destructor (storage mite) E. Mammals Cams domesticus'^ Cand2

27

74, 75 74 75

Fel d 1; cat-1

38

15

Mus m 1; MUP

19

76 77

Can d 1

Felis domesticus (cat saliva) Mus musculus (mouse urine) Rattus norvegicus

78, 79

Ratn 1 F. Fungi Aspergillus fumigatus Aspf 1 AspP Aspf?

90 55

Candida albicans 83 Alternana alternata Alt a 1

84-86

Tnchophyton tonsurans Tnt 1 G, Insects Apis mellifera (honeybee)

Api m 1; phosphoiipase A Api m 2: hyaluronidase Api m 4; melittin

769

Te F*iao King et al. Table 1 Continued, Allergens: systematic Allerger: source

and original names

MW (kDa)

Bombui> pennsvlvanicu^ ibumbiebfei

91 91

Bom p 1; phospholipase Bom p 4: protease

Blattel'.t germanica Blag 2

20

92

Chi t 1: hemoglobin

16

93

Chirono imidge) Dolichovespula maculata iwhitp face hornet

94 95 96,97

Dol m 1; phospholipase A, Dol m 2; hyaluronidase Dol m 5: antigen 5

Dolicho\/espula arenana (yellow hornet)

Dol a 5: antigen 5

Polistes annulans

99 99

Pol a 1: phospholipase A. Pol a 2: hyaiuronidase Pol a 5: antigen 5 Polistes exclamans (wa.sp!

Pol e 1: phospholipase A, Pol e 5: antigen 5

Polistes fuscatus (wa.spi

100

Pol f 5: antigen 5

Polistes metncus

100

Poi m 5: antigen 5

23

Vesf 5: antigen 5

23

100

Ves g 5; antigen 5

23

100

Ves m 1; phosphoiipase A,

33.5 44 23

102 103

Vespula' flavopilosa (yellow jacke Vespula germanica (yellow lacke Vespulamaculifrons ivellow lacke

Ves m 2: hvaluronidase Ves m 5: antigen 5

Ves p 5: antigen 5

100

Ves s 5: antigen 5

100

Ves vi 5

100

Vespula squamosa (yellow lacketi Vespula vidua iwasp! Vespula ['ulgans (yellow jacket!

Ves V 1: phopholipase A. Ves V 2: hvaluronidase Ves V 5' antigen 5

Vespa crabo (European horn

Vesp c 1: phospholipase

101

Vespc 5.0101: antigen 5

100

Vesp c 5.0102' antigen 5

100

Sol I 2

104, 105

Sol, 3

104

Son 4

104

bolenopsis invcta (fire ant!

Gadus callanas (cod!

770

Allergen nomenclature

Allergens; systematic and original names

Allergen source Gallus domesticus (chicken)

Penaeus aztecus (brown shrimp)

Brassica juncea (oriental mustard)

Gal Gal Gal Gal

d 1; ovomucoid d 2; ovalbumin d 3; conalbumin (Ag22) d 4; lysozyme

Pen a 1

MW (kDa!

107, 107, 107, 107,

lOS lOi 10S lOi

109 110

Pen a 2; tropomyosin

Bra i 1; 25 albumin

112 Sinapis alba (yellow mustard)

Horv 1;BMAI-1

113 Sin a 1; 25 albumin

I. Others Ascaris suum (worm) tievea brasiliensis (rubber)

Hev b 1; elongation factor

' References refer to those where partial (P) or complete (C) sequence data are available. The original references describing the initial characterization studies are not given because of limited space. Also we apologize to our colleagues whose allergen sequence data we may have overlooked for inclusion in this table, ^ Sequence data for groups 5 and 9 allergens from several grass pollens indicate that they are highly homologous proteins. Comparison of complete sequence data of groups 5 and 9 allergens from a single grass species will clarify whether these two groups are the same protein, ^ Canis domesticus is also designated as Cams familiaris.

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allergen database. IUIS. Baltimore, MD. Edition 1,0, 2. MARSH. D . G . . L. GOODFRIEND. T . P. KLNG. H . LOWENSTEIN. AND T. A. E. PEATTS-MILLS. 1986, Allergen no-

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AND T.P. KING. 1979. Cytcx-hromes C: New ragweed pollen allergens. Eed. Proc. 38: 1-^15. 13. EKRAMODDOUEIAH. A. K. M.. E. T. KISIE. AND A. LL

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teristics of the asparagine-linked oligosaccharide from honeybee venom phospholipase A2. Comp. Biochem. Physiol, 83B:321-324. 771

Te Piao King et al. IS WujFR \ H S c H R O D i R k [ H \LBERc, -WD I \\\RZ I'A" Sptcitie inlLi ittion of Iiil antibod'ts with acaibohx driiL Lpilopt (>( lionLvbct vtnoni ph ispholipast \ 2 MkrgN 42:464-470, 19 S i w w o R i H D R k I DoRRiNCiON T h H I G H K RiilJ \ \ [ > M W liRNLR 1990 Nomenclature foi sMithctn. pcpiidts iLpits^ni ltn.. of lmmunoglohulin chain scqucnccs Bull V\H()6S 109^111 ^0

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I Rr)Gt RS W D I) G K i x p p i R 1991 Cloning of \ m b 1 1 ( mlifcii 1 ) the niiioi illcmcn lannl\ of short rat.n\ccd pollc-iK E Biol Chem 266: 1229-1236. RoGi RS B l I P MoRf.irNsrtRN I I C i R i m i H \ B ^1 ( \1 ( f ) i N s u i \Vv B R X I I R 7 P K I N G R D C A R M A N XNO M C K I O 1991 C omplctc sequence of the ilicigcn

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uith I cells h o m I iL'uccd illciiu patients I ImmiuK 1 147:2547-2552. k i \Pi'iR [) G 1 G o o D i R i i N D AND I I) C \ P R \ 19S0 \niino aeid scquuice of i mweed lllcr^cn K i3 Biochcmis iry 19: 5:'29-5734. GHOSH

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19S9 C ompletc p n m a n stiuctuic of a t olinm peiennc (peienmal i\e grass) pollen allei gen l o l p I I I Compaiison with known I ol p I and II sequences Biochemistrv 2S S6658670 39. SiNGH. M. B.. T. H o u G H . P. T H E E R A K U L P I S U T . .A. \\IIOGII

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SiMPSON I D \ \ XRD I M C C L L S K E \ R P L \ \ N D R B K N O X 1991 KolatuMi of eDN \ encoding a newh identihed m qor allerL'emc piotein of i\c grass pollen lntia .ellulai taigeting to the am\loplost P U K Nail Acad Sci SS n s 4 - l ^ S S ^ \1XTIHIH-SEN T \ K N I E L S E N T I S O G A ^ R D S K L \ SN E R \ N D H EOWENSTEIN 1992 NH2-teiminal sc qucnces of foi'i immunoafihnit\ puiihcd mass pollen allei auis Phi p I Poa p 1 Sec c 1 and C\n d I (abst ) \lleig\ (SuppM"^ ^1 E \ P F t R S S \ R 1 \ L \ D K R X T T AND O S C H E I N E R 1992 a ) N \ JomnL^ of a maioi alleigcn of i \ c (Secale ec I eaie t uid tiiiiotln ill iss (Phleuiii pr ilcmse ) W th I Ul Con jicssof \llemoloL'\ iiidChn Immunol Pans MxTTHItsEN E \ND H LOWFNSTEIN 1991 Gioup \ allcigcns m eias^ pollen^ I Puiihe ilion and chaiacteii/a lion of tile eioup \ alleiuen fioiii PhleL 11. piatcns^ pollen Phi p \ Chn l x p \ l l e i c \ 2 ! 2^>''-M)'' \ R I X I A S S L A F H E R \ 1 DtCHENE D K R X H O S( HFI\ER \ND R \ X L ? N T \ l992 ^ DN \ eiomniz ol a nuiKM mass pollen allemen fiom tmunhMzi ass (Phleum pi at cnse) ldentific ifon as Phi p \ a piotein v ith possible tai oelin>j to the am\ loplast W t h l u i C onsziess of \lleigok>g\ md C hn Immunc^l Pans PiriRsc^N \ W \ l B L C K L R \ N D M ScHLXAk 1992 C 11 u ae tei lzatnn of isoloi Ills t>f the maioi allcigeii Phi p \ b\ imii'unoblotting md niK losequcncing Int \ i c h \llcig\ Immunol 9s i o s l l i ) 9 Bt f E

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XND M S C H L X X K 199^ Paitial mRN \ sequence of Phi p \ I MBl acce>-sion numbei \ ^ 0 9 4 2 46 F sc H R t XND D G Kl XPPER 19S9 Isolation and JiaiaUei 1/ation of i ma|oi u o s s leaetne siass moup 1 al lemenie determinant Mo! Immunol 26 ^ V - ^ 6 1 4~ OisFN E L / H A N G R D H I L L E T KISIL \ H SF HON XND S M O H X P A T R A 1991 Identihcafon and chai i. tenzation of the Poa p I \ moup ot basic alleiaens ol Kentuck\ blucgiass pollen I Immunol 1 4 ' ' 2 O ^ - 2 ' i ! 4S \ \ IlOGl ' \ ^ M SiNGH AND R B K N O X 1993 Se quencc ana!\sis ot Soi h 1 the gioup 1 allergen of Johnson mass pt>!!en and its eompanson to r\e-giass Eol p I (abst) I Mleigx Chn Immunol 91 340 49, B R E I T E N E D E R . H.. E. E E R R E I R A . A. R E I K E R S T O R F E R . M.

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^1 S\iiiH P M M B SiNc li x N o R B k \ ( ) \ I^H)^ ( I^ „ leteii/tiion md J o m i u ol IIK m uo! illcmcn of Beimud i " i i s s ( M , d l In Molcailii Bioio.x uid Immunolo N )f Mlei^ens (D K\ Ut md \ Selion eds I C RC Piess Boe I Ralon, pp, 1.-^" 160, '>^ M u i i i R i S c. P i i i K i W D B [ ) X \ I D 19s^ Puiili. 1 tion i i u K h i i i e t u i itioii ol i m uoi lilcijeii fiom I) i^txhs ^loniei 111 pi^ilen Ihe \ ' Hii 1 Im Vich \llci.\ \ p p l lmnumol, "S: 2S3-2S9, r , ROBIRIS \ M I 1 BlXXN P S FlORX 1 IH'SON VND M R W XI KI R 19«)^, NuJcotide scq lcncc ot . t ) N \ en codiiu the Ciioup I! iHeiiieii of eoci-sfoot oiehard m iss ( D i a x h s aiomei tta) Dae e 11 I ui I \!leig\ C !m lnunu U

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