ARTICLE 87

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citreoroseum,

P. chrysogenum P. P. P. griseoroseum P. P. citreoroseum P. et al.   6LQFH However, , which were described , which were described nomen conservandum DQGWRUHMHFWWKHROGHU Key words: Key words: Fleming indoor mycology phylogeny taxonomy P. griseoroseum is predated by P. et al   VXJJHVWHG WR ZDVGHVFULEHGE\7KRP  

P. chrysogenum sensu lato P.  7KLV SURSRVDO ZDV DFFHSWHG and with its synonyms DQGV\QRQ\PL]HG P. chrysogenum P. IMA FUNGUS · VOLUME 2 · NO 1: 87–95 · NO 2 · VOLUME FUNGUS IMA P. griseoroseum. with P. is a widely used name, Frisvad is currently listed as P. brunneorubrum and P. P. chrysogenum P. does not produce but chrysogenum Penicillium P. chrysogenum P. et al.  P. griseoroseum P. P. brunneorubrum P. P. chrysogenum P. P. rubens P. Penicillium chrysogenum 7KH DLP RI WKLV VWXG\ ZDV WR ¿QG RXW ZKHWKHU WKH WZR ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get Any of the above conditions can be waived ivecommons.org/licenses/by-nc-nd/3.0/legalcode. P. chrysogenum but the name P. citreoroseum, P. E\ 'LHUFN[   3LWW   DFFHSWHG and P. brunneorubrum P. chrysogenum P. name and E\ WKH &RPPLWWHH IRU )XQJL DQG /LFKHQV DQG WKH QDPH   DQG .R]DNLHZLF] preserve the name morphological, physiological and extrolite data showed data showed morphological, physiological and extrolite WKDW WKHVH VSHFLHV ZHUH FRQVSHFL¿F ZLWK )ULVYDG  )LOWHQERUJ  %DQNH 0F1HLOO most common clades in et al UHSUHVHQWRQHRUWZRVSHFLHV7RZDUGVWKLVDLP we have studied the most important strains in the history of penicillin production by combining extrolite and molecular GDWD)XUWKHUPRUHWKHKLVWRU\RIWKHWD[RQRP\RI)OHPLQJ¶V P. chrysogenum P. SHQLFLOOLQSURGXFHUVWUDLQLVGLVFXVVHG

1 is OXPSLGLQZKLOH and , such isolates revealed the presence of two highly isolates revealed the  )ULVYDG et al P. rubrumLVGLVFXVVHG P. P. chrysogenum P. WRWKRVHFODGHV )XUWKHUPRUHWKHZHOONQRZQFODLPWKDW$OH[DQGHU)OHPLQJ P. chrysogenum P. P. meleagrinum, P. P. chrysogenum P. et al.   UHJDUGHG P. dipodomyis P. et al.  )lUEHU  Penicillium chrysogenum series Chrysogena is a commonly occurring mould in indoor environments and foods, mould in indoor environments and foods, is a commonly occurring DQG5REHUW$6DPVRQ  and P. rubens P. The taxonomy of this species

et al.  *UDYHVHQ  P. chrysogenum P. is a commonly occurring mould P. notatum, P. , VHULHV´0RUHUHFHQWO\RWKHUVSHFLHV Penicillium et al   VWXGLHG WKH WD[RQRP\ RI et al  )ULVYDG  6DPVRQ  

et al.  P. nalgiovense, P.  +RZHYHU 6DPVRQ  XVLQJ PXOWLORFXV VHTXHQFH W\SLQJ 0/67  , -HQV&)ULVYDG 1 DUHUHLGHQWL¿HGDV Penicillium chrysogenum Penicillium chrysogenum

 7KHVH VSHFLHV DUH SKHQRW\SLFDOO\ VLPLODU EXW H[WUROLWH DQDO\VLV VKRZV WKDW P. chrysogenum P. P. chrysogenum P.   antibiotic much attention for its use in the production SHQLFLOOLQ 6DPVRQ Abstract: DQGKDVJDLQHGPXFKDWWHQWLRQIRULWVXVHLQWKHSURGXFWLRQRIWKHDQWLELRWLFSHQLFLOOLQ3K\ORJHQHWLFDQDO\VLV DOWKRXJKDOOVSHFLHVSURGXFHSHQLFLOOLQ )ULVYDG chrysogenum P. GDWDDQGVKRZHGWKDWLWFRXOGEHVXEGLYLGHGLQWRIRXUFODGHV VOLUME 2 · NO. 1 this series as one broad species and did not accept Raper and 7KRP¶V³ from and these species are clearly different INTRODUCTION Penicillium chrysogenum in indoor environments such as dust, indoor air, and damp indoor air, in indoor environments such as dust, EXLOGLQJ PDWHULDOV &KDQJ IUHTXHQWO\LGHQWL¿HGDVDIRRGVSRLODJHDJHQWDQGKDVJDLQHG  1 Jos Houbraken Fleming’s penicillin producing strain is not strain producing penicillin Fleming’s GRLLPDIXQJXV P. rubens P. et al.  %DQNH have been included in as +XQWHU  /HD   )XUWKHUPRUH Houbraken et al.

Table 1. Overview of Penicillium chrysogenum and P. rubensVWUDLQVVWXGLHG No. Other collection no. Name Substrate, locality Remarks 155/ '72* ,%7 P. chrysogenum Unrecorded source Ex-lectotype of P. griseoroseum ,%7 ,0,

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ARTICLE &%6 155/ ,%7 P. chrysogenum Cheese, Storrs, Connecticut, USA Ex-lectotype of P. chrysogenum ,0, '72, P. chrysogenum Fungal growth on ceiling of archive, 8WUHFKWWKH1HWKHUODQGV '72% ,%7 & P. chrysogenum +RXVHGXVW:DOODFHEXUJ21 5HSUHVHQWDWLYHRIJURXSLQWKH Canada study of Scott et al  &%6 155/ ,%7 P. chrysogenum Branches of Hyssopus1RUZD\ Ex-type of P. notatum '72' ,0, &%6 155/ P. rubens Must contaminant, Belgium The strain first used for producing penicillin in VXEPHUJHGFXOWXUH 5DSHU  7KRP± &%6 155/ ,%7 P. rubens Culture contaminant in bacterial )OHPLQJ¶VRULJLQDOSHQLFLOOLQ ,0, FXOWXUH8. producing strain '72( ,%7 P. rubens Cap of beer bottle, Belgium 155/T '72( ,%7 P. rubens Unrecorded source Ex-lectotype of P. rubens &%6 155/ ,%7 P. rubens MouldyFDQWDORXSH3HRULD,OOLQRLV ³:LVFRQVLQVWUDLQ´SDUHQWRI ,0, USA most high yielding penicillin producing strains :LVFRQVLQ P. rubens MouldyFDQWDORXSH3HRULD,OOLQRLV )XOOJHQRPHVHTXHQFHGVWUDLQ USA

MATERIAL AND METHODS EUDQFK OHQJWK RSWLPL]DWLRQ ZDV FDUULHG RXW 6WDPDWDNLV et al   0D[LPXP SDUVLPRQ\ DQDO\VLV ZDV SUHIRUPHG DV Strains D VHFRQG PHDVXUH RI EUDQFK VXSSRUW 7KLV DQDO\VLV ZDV An overview of Penicillium chrysogenum strains used UXQLQ0(*$DQGERRWVWUDSYDOXHVZHUHGHWHUPLQHGXVLQJ LQ WKLV VWXG\ LV JLYHQ LQ 7DEOH  7KUHH SRLQW SRVLWLRQ  UHSOLFDWHV Penicillium griseofulvum &%6  ZDV inoculations of these strains were made in order to study VHOHFWHGDVDQRXWJURXS1HZO\JHQHUDWHGVHTXHQFHVZHUH WKHLU PDFURPRUSKRORJ\ 7KH VWUDLQV ZHUH LQRFXODWHG RQ GHSRVLWHGLQ*HQEDQNXQGHU-)±-) &]DSHN\HDVWH[WUDFWDJDU &<$ \HDVWH[WUDFWVXFURVHDJDU <(6  FUHDWLQH DJDU &5($  DQG PDOW H[WUDFW DJDU 0($  Extrolite analysis 2[RLG $OOPHGLDZHUHSUHSDUHGDVGHVFULEHGE\6DPVRQ 6WUDLQVZHUHJURZQIRU±GDWƒ&RQWKHDJDUPHGLD et al  &RORQ\FKDUDFWHULVWLFVZHUHUHFRUGHGDIWHUG <(6 &<$ DQG 0($ SULRU WR H[WUROLWH H[WUDFWLRQ ([WUROLWHV RILQFXEDWLRQDWƒ&0LFURVFRSLFPRXQWVZHUHPDGHIURP ZHUH H[WUDFWHG DQG DQDO\]HG E\ +3/&'$' DV GHVFULEHG MEA or OA and microscopical features were studied by light E\ )ULVYDG  7KUDQH   DQG PRGL¿HG E\ 6PHGVJDDUG PLFURVFRS\ 2O\PSXV%+DQG=HLVV$[LRVNRS3OXV    ,GHQWL¿FDWLRQ ZDV SHUIRUPHG E\ FRPSDULVRQ RI WKH UV spectra and retention times of the extrolites with those of Molecular analysis SXUL¿HGDQGFKHPLFDOO\FKDUDFWHUL]HGFRPSRXQGVSUHVHQWLQ '1$H[WUDFWLRQZDVSHUIRUPHGXVLQJWKH8OWUDFOHDQ0LFURELDO WKHFROOHFWLRQDWWKH%LRFHQWUXP'78 /\QJE\  '1$ LVRODWLRQ .LW 0R%LR 6RODQD %HDFK 86$  DFFRUGLQJ WKH PDQXIDFWXUHU¶V LQVWUXFWLRQV 3DUWV RI WKH ȕWXEXOLQ DQG FDOPRGXOLQJHQHVZHUHDPSOL¿HGDQGVHTXHQFHGDFFRUGLQJ RESULTS to the method described by Houbraken et al.  $SDUW RIWKH53%ORFXVZDVDPSOL¿HGXVLQJWKHSULPHUSDLU53% Phylogenetic analysis )B3F ¶ *$7*$&&*7*$&&$&77&**¶ DQG 53% The structure of the partitions of the alignment was as follows: &5B3F &5 ¶ &&&$7**&77*777*&&&$7¶ 7KH WKH FDOPRGXOLQ SDUWLWLRQ ZDV  EDVHSDLUV ORQJ DQG KDG alignments were combined and maximum likelihood analysis  GLVWLQFW DOLJQPHQW SDWWHUQV WKH 53% GDWDVHW LQFOXGHG ZDV SHUIRUPHG XVLQJ 5$[0/ Y  (DFK GDWDVHW ZDV  FKDUDFWHUV DQG  RI WKRVH KDG D GLVWLQFW DOLJQPHQW WUHDWHGDVDVHSDUDWHSDUWLWLRQ7KH*75VXEVWLWXWLRQPDWUL[ SDWWHUQ DQG WKH WXEXOLQ JHQH SDUWLWLRQ ZDV  EDVHSDLUV was used for all three partitions and individual per partition ORQJ  ZHUH IRXQG WR KDYH D GLVWLQFW DOLJQPHQW SDWWHUQ ,Q

88 IMA FUNGUS Penicillium rubens, the original penicillin-producer

P. rubens CBS 205.57, Fleming’s penicillin producer ARTICLE

P. rubens CBS 307.48, Wisconsin strain wild-type

P. rubens Wisconsin 54-1255, full genome sequenced strain

P. rubens CBS 197.46, first strain used for submerged penicillin production 100/99 P. rubens, NRRL 792T, ex-lectotype

P. rubens DTO 95E9

100/99 P. chrysogenum CBS 355.48, type of P. notatum

P. chrysogenum DTO 87I2

P. chrysogenum DTO 102B4 100/93 P. chrysogenum NRRL 820, ex-lectotype of P. griseoroseum

73/- T 100/99 P. chrysogenum CBS 306.48 , ex-lectotype

P. flavigenum CBS 419. 89T

100 P. confertum CBS 171.87T 99

P. mononematosum CBS 172.87T

T 100/99 P. dipodomyis CBS 110412

P. nalgiovense CBS 352.48T

P. persicinum CBS 111235T

P. aethiopicum CBS 484.84T

P. griseofulvum CBS 185.27T 0.01

Fig. 1.%HVWVFRULQJ0/WUHHRIFRPELQHGSDUWLDOȕWXEXOLQFDOPRGXOLQDQG53%GDWDVHWV7KHERRWVWUDSYDOXHVRIWKHPD[LPXPOLNHOLKRRG DQDO\VLV 0/¿UVWQXPEHU DQGPD[LPXPSDUVLPRQ\DQDO\VLV 03VHFRQGQXPEHU DUHVKRZQDWWKHQRGHV7KHERRWVWUDSYDOXHVDERYH DUHVKRZQDQGEUDQFKHVVXSSRUWHGOHVVWKDQLQHLWKHUWKH03RU0/DQDO\VLV DUHLQGLFDWHGZLWKDK\SKHQ

WKHPD[LPXPSDUVLPRQ\DQDO\VLVVLWHVZHUHSDUVLPRQ\ centered around the ex-type strain of P. chrysogenum &%6 LQIRUPDWLYH 03WUHHVOHQJWK FRQVLVWHQF\LQGH[  $OVRWKHH[W\SHVWUDLQVRIP. notatum &%6  UHWHQWLRQLQGH[ 7KHSHUFHQWDJHRIUHSOLFDWHWUHHVLQ P. griseoroseum 155/   DQG VHYHUDO RWKHU VWUDLQV DUH which the associated taxa clustered together in the bootstrap DFFRPRGDWHG LQ WKLV FODGH 7KH RWKHU FODGH LV FHQWHUHG WHVW LV VKRZQ QH[W WR WKH EUDQFKHV )HOVHQVWHLQ   7KH around the ex-type strain of P. rubens 155/ )OHPLQJ¶V EHVWVFRULQJPD[LPXPOLNHOLKRRGWUHHLVVKRZQLQ)LJ RULJLQDOSHQLFLOOLQSURGXFLQJVWUDLQ &%6 WKHZLOGϢtype 3K\ORJHQHWLF DQDO\VLV RIWKHFRPELQHG SDUWLDO ȕWXEXOLQ :LVFRQVLQ VWUDLQ LVRODWHG IURP D PRXOG\ FDQWDORXSH &%6 FDOPRGXOLQDQG53%GDWDVHWVUHYHDOHGWZRKLJKO\VXSSRUWHG   155/   WKH IXOO JHQRPH VHTXHQFHG VWUDLQ clades in Penicillium chrysogenum sensu lato2QHFODGHLV GHULYHGIURP155/ :LVFRQVLQϢ  YDQGHQ%HUJ

VOLUME 2 · NO. 1 89 Houbraken et al.

Table 2([WUROLWHVSURGXFHGE\Penicillium chrysogenum and P. rubens strains studied, with a focus on the important strains for the history of SHQLFLOOLQ No. Other collection no. Name Extrolites

&%6T 155/ ,%7 P. chrysogenum $QGUDVWLQ$DQGUDVWLQ%FKU\VRJLQHFLWUHRLVRFRXPDULQFI OXPSLGLQPHOHDJULQSHQLFLOOLQURTXHIRUWLQH&VHFDORQLFDFLG' ARTICLE secalonic acid F, sorbicillins &%6 155/ ,%7 '72' P. chrysogenum $QGUDVWLQ%FKU\VRJLQHFIOXPSLGLQPHOHDJULQSHQLFLOOLQURTXH- fortine C, secalonic acid D, sorbicillins 155/ '72* ,%7 ,%7 P. chrysogenum $QGUDVWLQ$DQGUDVWLQ%FKU\VRJLQHFDQPHOHDJULQFI OXPSLGLQURTXHIRUWLQH&VRUELFLOOLQV '72% ,%7 & P. chrysogenum $QGUDVWLQ$DQGUDVWLQ%FKU\VRJLQHPHOHDJULQFIOXPSLGLQ secalonic acid D, secalonic acid F &%6 155/ ,%7 P. rubens &KU\VRJLQHPHOHDJULQSHQLFLOOLQURTXHIRUWLQH&DQGUDVWLQ$ sorbicillins 155/T '72( ,%7 P. rubens &KU\VRJLQHPHOHDJULQSHQLFLOOLQFIURTXHIRUWLQHVRUELFLOOLQV &%6 155/ ,%7 P. rubens &KU\VRJLQHPHOHDJULQSHQLFLOOLQURTXHIRUWLQH&URTXHIRUWLQH' sorbicillins '72( ,%7 P. rubens 0HOHDJULQURTXHIRUWLQH&

et al. WKHVWUDLQ¿UVWIRXQGWRSURGXFHVDWLVIDFWRU\\LHOGV Synonyms: Penicillium griseoroseum Dierckx, Ann. Soc. Sci. RI SHQLFLOOLQ ZKHQ JURZQ VXEPHUJHG &%6   155/ Brux. 25  QRPUHM   DUH DFFRPPRGDWHG LQ WKLV FODGH 7KHVH WZR FODGHV Penicillium citreoroseum Dierckx, Ann. Soc. Sci. Brux. 25 evidently represent two different species, P. chrysogenum  QRPUHM and P. rubens Penicillium brunneorubrum Dierckx, Ann. Soc. Sci. Brux. 25:   QRPUHM Phenotypic and extrolite analysis Penicillium notatum :HVWOLQJArk. Bot11     3KHQRW\SLF H[DPLQDWLRQ XVLQJ PDFUR DQG PLFURVFRSLFDO characters of strains of the Penicillium rubens and Type: USA: Connecticut 6WRUUV LVRO H[ FKHHVH  C. P. chrysogenum FODGHV GLG QRW UHYHDO FOHDU GLIIHUHQFHV Thom ,0,±W\SFRQV  7KHPDMRULW\RIWKHH[DPLQHGVWUDLQVVKRZHGVLPLODUJURZWK characteristics, such as good growth and sporulation on Penicillium rubens Biourge, La Cellule 33   YES agar, weak or moderate growth on CREA with poor acid production and large clear or yellow coloured exudate Type: Unrecorded source, P. Biourge &%6 + ± GURSOHWV RQ 0($ DQGRU &<$ )LJ   7KUHH VWUDLQV KDG lectotypus hic designatus IURP %LRXUJH   WR 7KRP GHYLDWLQJJURZWKFKDUDFWHULVWLFV&%6 H[W\SHFXOWXUH  WR155/  WR&%6FXOWXUHV of P. notatum KDGVPDOOHUFRORQLHVRQDOOWHVWHGDJDUPHGLD H[OHFWRW\SH±155/ ,%7 $7&& &%6 DQG155/ H[W\SHFXOWXUHRIP. rubens)LJ DQG&%6     155/  ZLOGW\SH IXOO JHQRPHVHTXHQFHG VWUDLQ GHYLDWHGLQKDYLQJUHGXFHGVSRUXODWLRQRQPRVWDJDU Penicillium chrysogenum has, in addition to the species PHGLD names mentioned above, also various other synonyms Extrolite analysis differentiated the P. chrysogenum and 6DPVRQ et al  6DPVRQ  )ULVYDG   7KHVH P. rubens clades. An overview of extrolites produced by include P. meleagrinum %LRXUJH  P. cyaneofulvum these species, with emphasis on the important strains in the %LRXUJH  P. chlorophaeum %LRXUJH  P. roseo- KLVWRU\RISHQLFLOOLQLVVKRZQLQ7DEOH7KHVHUHVXOWVVKRZ citreum  P. camerunense +HLP  QRP LQYDO that P. chrysogenumSURGXFHVVHFDORQLFDFLG'DQG)DQG P. chrysogenumYDUbrevisterigma )RUVWHU QRP LQYDO RUDPHWDEROLWHUHODWHGWROXPSLGLQ /DUVHQet al ZKLOH  P. aromaticum I microsporum 5RPDQNRYD  P. rubensGRHVQRWSURGXFHWKHVHPHWDEROLWHV P. harmonense%DJKGDGLP. verrucosumYDUcyclopium VWUDLQ ³DQDQDVROHQV´ 5DPtUH]  DQG P. chrysogenum ³PXW fulvescens” Takashima et al. H[ 5DPtUH]  $ TAXONOMY more detailed molecular or extrolite analysis is needed to properly place these species in either P. chrysogenum, Penicillium chrysogenum Thom, Bull. Bur. Anim. Ind. P. rubens, or other currently unrecognised species belonging to USDA 118  QRPFRQV P. chrysogenum sensu lato

90 IMA FUNGUS Penicillium rubens, the original penicillin-producer ARTICLE

Fig. 2. Penicillium rubens&%6 )OHPLQJ¶VRULJLQDOSHQLFLOOLQSURGXFHU A–C.&RORQLHVGROGƒ&A.&<$B.0($C.<(6D–H. &RQGLRSKRUHVI.&RQLGLD%DUV —P

VOLUME 2 · NO. 1 91 Houbraken et al. ARTICLE

Fig. 3Penicillium rubens155/A–C.&RORQLHVGROGƒ&A.&<$B.0($C.<(6D–H.&RQGLRSKRUHVI.&RQLGLD Bars —P

92 IMA FUNGUS Penicillium rubens, the original penicillin-producer

DISCUSSION 6WROO DVGH¿QHGE\%LRXUJH DQGWKLVLGHQWL¿FDWLRQZDV ARTICLE accurate at that time and not incorrect as claimed in Identity of Fleming’s original penicillin- OLWHUDWXUH 7KRP5DSHU 7KRP/D[  producing strain )XUWKHUPRUH %LRXUJH   ZURWH LQ KLV GHVFULSWLRQ RI ,W KDV ORQJ EHHQ FODLPHG WKDW )OHPLQJ   PLVLGHQWL¿HG P. rubrum that the species was a common laboratory the original penicillin-producing strain as “Penicillium rubrum FRQWDPLQDQW ³«XQDQLPDOGHODERUDWRLUH´ ZKLFKFRXOG %LRXUJH ´ 5DSHU 7KRP  /D[   +RZHYHU KDYH VWUHQJWKHQHG /D7RXFKHLQ KLV LGHD RI KDYLQJ WKH LW ZDV WKH P\FRORJLVW &KDUOHV - /D 7RXFKH ±  FRUUHFWLGHQWL¿FDWLRQVLQFH)OHPLQJ¶VLVRODWHZDVDOVRD ZRUNLQJLQDURRPRQWKHÀRRUEHORZWKDWRFFXSLHGE\)OHPLQJ ODERUDWRU\FRQWDPLQDQW ZKRDFWXDOO\LGHQWL¿HGWKHVWUDLQDVPenicillium rubrum2XU current study shows that the correct name of this strain is P. Name changes of Fleming’s penicillin- rubens:HGLVFXVVWZRSRVVLEOHVFHQDULRVKHUHZKLFKFRXOG producing strain KDYHOHG/D7RXFKHWRKLVLGHQWL¿FDWLRQRI)OHPLQJ¶VVWUDLQDV :KHQ&KDUOHV7KRPZDVDERXWWR¿QLVKKLVPRQRJUDSKLQ P. rubrum ERWK VKRZLQJ WKDW KLV LGHQWL¿FDWLRQ ZDV UDWKHU  KHUHFHLYHGWKH)OHPLQJVWUDLQ &%6 155/ accurate:   ,0,   WKDW KDG EHHQ LGHQWL¿HG DV Penicillium 1. /D 7RXFKHLGHQWL¿HG )OHPLQJ¶V VWUDLQ DV P. rubrum, but rubrumDQGKHUHLGHQWL¿HGLWDVP. notatum 7KRP  this was a lapsus for P. rubens 7KH HSLWKHW rubrum +H GLG QRW UHDOL]H WKDW WKH )OHPLQJ LVRODWH ZDV LGHQWL¿HG means “red´ DQG rubens “to be red´ DQG WKXV LW LV QRW E\ WKH H[SHULHQFHG P\FRORJLVW &KDUOHV /D 7RXFKH VLQFH XQOLNHO\ WKDW WKHVH HSLWKHWV KDYH EHHQ FRQIXVHG %RWK KH ZURWH ³1RW EHLQJ D P\FRORJLVW KH >$OH[DQGHU )OHPLQJ@ yellow and red-brown reverse colours have been undertook to identify the mold from the literature and observed in P. chrysogenum sensu lato and in the true selected the name Penicillium rubrum%LRXUJH´ 7KRP  P. rubrum LHP. purpurogenum  3LWW 7KHVHWZR However, the authority name Biourge is not mentioned in species clearly differ in micro-morphology from each )OHPLQJ¶VZRUNDQG7KRPZRXOGKDYHEHHQPRUHDFFXUDWH RWKHUDQGWKHP\FRORJLVW/D7RXFKHZRXOGKDYHQRWLFHG if he had named it “Penicillium rubrum *UDVVEHUJHU WKHVH GLIIHUHQFHV 3LQN DQG YLQDFHRXV GUDE FRORXUV Stoll sensu %LRXUJH´ 6XEHTXHQWO\ 6DPVRQ et al   DUH PHQWLRQHG E\ 5DSHU  7KRP   IRU VWUDLQV RI broadened the concept of P. chrysogenum and placed P. meleagrinum and reddish colours are part of the names P. notatumLQV\QRQ\P\ZLWKWKHIRUPHUVSHFLHV7KLVWUDQVIHU of synonyms of P. chrysogenum sensu lato, such as ZDVODWHUVXSSRUWHGLQVHYHUDORWKHUZRUNV 3LWW)ULVYDG P. citreoroseum, P. roseocitreum, P. brunneorubrum, and  )LOWHQERUJ  %DQNH et al  6DPVRQ  )ULVYDG P. griseoroseum  )URPWKLVSRLQWRQ)OHPLQJ¶VSHQLFLOOLQSURGXFHUZDV 2. ,QWKHRWKHUVFHQDULR/D7RXFKHFRUUHFWO\LGHQWL¿HGWKH LGHQWL¿HGDVP. chrysogenum2XUFXUUHQWVWXG\VKRZVWKDW strain as P. rubrum based on the taxonomic schemes the correct name of this strain is P. rubens SUHVHQWDWWKDWWLPH,QWKHPRVWUHFHQWPRQRJUDSK on PenicilliumZDVWKDWRI%LRXUJH  DQG/D7RXFKH Origin of the Fleming strain ZRXOG KDYH XVHG WKLV PRQRJUDSK IRU LGHQWL¿FDWLRQ &KDUOHV /D 7RXFKH ZDV ZRUNLQJ ZLWK PRXOGV IURP LQGRRU RI )OHPLQJ¶V SHQLFLOOLQSURGXFLQJ VWUDLQ 7KLV LV DOVR DLU DQG HVSHFLDOO\ WKRVH LVRODWHG IURP FREZHEV +H LVRODWHG VXSSRUWHG E\ WKDW PRQRJUDSK EHLQJ FLWHG LQ )OHPLQJ¶V KXQGUHGVRIVWUDLQVIURPSRRUHUSDUWVRI/RQGRQDQG6KHI¿HOG UHSRUWRIWKHRULJLQDOGLVFRYHU\RISHQLFLOOLQ )OHPLQJ  +DUH.LQJVWRQ DQGDODUJHSHUFHQWDJHRIWKHP Penicillium rubrum LV OLVWHG LQ %LRXUJH¶V PRQRJUDSK DV were Penicillium chrysogenum and P. rubens. Furthermore, “P. rubrum " *UDVVEHUJHU6WROO´DQGWKLVVSHFLHV VXEVHTXHQWUHVHDUFKKDVVKRZQWKDWWKHVHPLJKWEHWKHPRVW ZDV FODVVL¿HG LQ %LRXUJH¶V VXEVHFWLRQ ³Les Radiés, common PenicilliumLQLQGRRUDLU 6FRWWet al6DPVRQet Radiata´7KLVVXEVHFWLRQDOVRFRPSULVHVP. chrysogenum al /D7RXFKH¶VK\SRWKHVLVZDVWKDWWKHVHPRXOGVFRXOG and the species P. griseoroseum, P. cyaneofulvum, FDXVHDVWKPDDQGWKLVZDVODWHUFRQ¿UPHGE\IRUH[DPSOH P. brunneorubrum, P. notatum, P. citreoroseum, and :DUGet al  ,WFRXOGYHU\ZHOOEHWKDWRQHRI/D7RXFKH¶V P. baculatum, which are currently all listed as synonyms FXOWXUHV IURP LQGRRU DLU FRQWDPLQDWHG )OHPLQJ¶V SODWH VLQFH of P. chrysogenum([DPLQDWLRQRI%LRXUJH¶VGUDZLQJVRI they were working in the same building and Fleming always “P. rubrum " *UDVVEHUJHU6WROO´VKRZVWKDWKLVGH¿QLWLRQ KDG KLV ZLQGRZV VKXW +DUH   %RWK P. rubens and P. of P. rubrum is very similar to that of P. chrysogenum chrysogenum produce penicillin, and in that respect it is not DQG GRHV QRW ¿W LQ WKH RULJLQDO FRQFHSW RI P. rubrum surprising that P. rubens ZDV SUHVHQW ,W LV QRWLFHDEOH WKDW 6WROO3LWW 7KLVZDVDOVRQRWHGE\%LRXUJH WKHKLVWRULFDOLPSRUWDQWSHQLFLOOLQSURGXFHUV :LVFRQVLQVWUDLQ   DQG WKHUHIRUH KH DGGHG WKH TXHVWLRQ PDUN WR 155/)OHPLQJ¶VVWUDLQ&%6 155/ ,0, his description of P. rubrum )XUWKHUPRUH WKH FORVH  DQG WKH VWUDLQ ¿UVW XVHG IRU SURGXFLQJ SHQLFLOOLQ LQ connection between P. rubrum *UDVVEHUJHU6WROO sensu VXEPHUJHGFRQGLWLRQV&%6 DOOSURYHGWREHP. rubens Biourge and P. rubensZDVDOVRQRWHGE\7KRP    ZKR DVVLJQHG WKH IRUPHU VSHFLHV WR P. rubens ,Q Current taxonomy and implications FRQFOXVLRQ/D7RXFKHXVHG%LRXUJH¶VWD[RQRPLFVFKHPH +HUHZHUHFRJQL]HWKHSUHVHQFHRIWZRVSHFLHVLQPenicillium DQGLGHQWL¿HG)OHPLQJ¶VVWUDLQDVP. rubrum*UDVVEHUJHU chrysogenum sensu lato 7KH SUHVHQFH RI IRXU OLQHDJHV

VOLUME 2 · NO. 1 93 Houbraken et al.

in P. chrysogenum was described by Scott et al   )ULVYDG-&+DZNVZRUWK'/.R]DNLHZLF]=3LWW-,6DPVRQ5$6WRON 6FRWW¶V OLQHDJHV  DQG  DUH P. chrysogenum, and lineage $&  3URSRVDOWRFRQVHUYHLPSRUWDQWQDPHVLQAspergillus  FRUUHVSRQGV WR P. rubens7KH VWDWXV RI OLQHDJH  LV VWLOO and Penicillium,Q6DPVRQ5$3LWW-, HGV Modern concepts XQGHULQYHVWLJDWLRQ,WZDVVXJJHVWHGE\6FRWWet al   in Penicillium and Aspergillus FODVVL¿FDWLRQ ± 1HZ 3URFHHGLQJV RI WKH 7KLUG ,QWHUQDWLRQDO &RQIHUHQFH LQ RWKHU JHRJUDSKLF UHJLRQV 7KH IXOO JHQRPH VHTXHQFHG RQ)XQJL0\FRWR[LQVDQG%LRDHURVROV6DUDWRJD6SULQJV1HZ VWUDLQGHULYHGIURP155/ YDQGHQ%HUJ LVDOVRD 5HJQXP9HJHWDELOHYRO@ 5XJJHOO$5**DQWQHU9HUODJ.* RISHQLFLOOLQApplied and Environmental Microbiology 60± 3LWW -,   >³´@ The genus Penicillium and its teleomorphic  states Eupenicillium and Talaromyces/RQGRQ$FDGHPLF3UHVV )HOVHQVWHLQ-  &RQ¿GHQFHOLPLWVRQSK\ORJHQLHV$QDSSURDFK 5DSHU .% 7KRP &   Manual of the Penicillia %DOWLPRUH XVLQJWKHERRWVWUDSEvolution 39± :LOOLDPV :LONLQV )OHPLQJ $   2Q WKH DQWLEDFWHULDO DFWLRQ RI FXOWXUHV RI D 6DPVRQ 5$ +DGORN 5 6WRON$&  $ WD[RQRPLF VWXG\ RI WKH Penicillium, with special reference to their use in the isolation Penicillium chrysogenumVHULHVAntonie van Leeuwenhoek 43: of %LQÀXHQ]DHBritish Journal of Experimental Pathology 10: ± ± 6DPVRQ 5$ )ULVYDG -&   Penicillium subgenus Penicillium: )ULVYDG-&)LOWHQERUJ2:LFNORZ'7  Terverticillate penicillia QHZ WD[RQRPLF VFKHPHV DQG P\FRWR[LQV DQG RWKHU H[WUROLWHV isolated from underground seed caches and cheek pouches of Studies in Mycology 49± EDQQHUWDLOHGNDQJDURRUDWV Dipodomys spectabilis Canadian Samson RA, Houbraken J, Thrane U, Frisvad JC, Andersen B Journal of Botany 65±  , Food and Indoor Fungi.>&%6 /DERUDWRU\ Manual series )ULVYDG -& )LOWHQERUJ 2   Terverticillate penicillia: chemo- QR@8WUHFKW&%6.1$:)XQJDO%LRGLYHUVLW\&HQWUH WD[RQRP\DQGP\FRWR[LQSURGXFWLRQMycologia 81± 6FRWW - 8QWHUHLQHU :$ :RQJ % 6WUDXVV 1$ 0DOORFK '   )ULVYDG-&6DPVRQ5$  3RO\SKDVLFWD[RQRP\RIPenicillium *HQRW\SLF YDULDWLRQ LQ Penicillium chrysogenum from indoor subgenus Penicillium D JXLGH WR LGHQWL¿FDWLRQ RI IRRG DQG HQYLURQPHQWVMycologia 96± DLUERUQHWHUYHUWLFLOODWHSHQLFLOOLDDQGWKHLUP\FRWR[LQVStudies in 6PHGVJDDUG -   0LFURVFDOH H[WUDFWLRQ SURFHGXUH IRU Mycology 49± VWDQGDUGL]HG VFUHHQLQJ RI IRU IXQJDO PHWDEROLWH SURGXFWLRQ LQ )ULVYDG-&7KUDQH8  6WDQGDUGL]HG+LJK3HUIRUPDQFH/LTXLG FXOWXUHVJournal of Chromatography A 760± &KURPDWRJUDSK\RIP\FRWR[LQVDQGRWKHUIXQJDOPHWDEROLWHV 6WDPDWDNLV $ +RRYHU 3 5RXJHPRQW -   $ UDSLG ERRWVWUDS EDVHGRQDON\OSKHQRQHLQGLFHVDQG899,6VSHFWUD GLRGHDUUD\ DOJRULWKPIRUWKH5$[0/:HE6HUYHUVSystematic Biology 75: GHWHFWLRQ Journal of Chromatography 404± ±

94 IMA FUNGUS Penicillium rubens, the original penicillin-producer

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