AR TICLE Homothallism: an Umbrella Term for Describing

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[ 9 [ O % V % $ < ] W / + N Z % ; ; % + 569 [; @ < U ; ; W / + N Z % ; ; % + 56 Article info:@ [57U 56!4K/ [!4`56!4K; [!"`56!4 V % %% . K $ V % % Abstract: / 665EK . [ [^ O 9 [; 665E@ / © 2015 International Mycological Association © 2015 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: ! 5 / U Homothallism: an umbrella term for describing diverse sexual behaviours diverse for describing umbrella term an Homothallism: [ Non-commercial:No derivative works:Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. G !""7; 56!4W $ $ . % 9 566# V * I + !"67 $ V [ . [ + !"67 [ $ 9% O P . % < % . [ % O % $ % $ V % @ OO 56!7 % ? % $ . N [ % @$ % 9% $ % % V% .9 O INTRODUCTION [!644"E\ 56!46F6!!_ VOLUME 6 · NO. 1 Wilson et al.

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ARTICLE !"5#N [ CONVENTIONAL CATEGORIES OF . HOMOTHALLISM W Neurospora N. Primary homothallism tetrasperma <!"5#V ; "6 z . % 9 !""5 [ ? $ MAT1-1 $ U/V.! $ MAT1-2 > et al.56!!KW ! U/V.5 <!"5#V V $ MAT genes % U/V.! U/V.5 . V * I 9 566# ] % O P $ $ % U et al.!""FV ; et al.!""# et al. æ!6 z N. tetrasperma are !"""] et al.5664 V % $ !""5V % % V Aspergillus nidulans$ $ [ ; % et al. !"4_ N % !""5/ % $ % [ % et al. 56!6] [ A. nidulans MAT1-1-1 % MAT1-2-1 % 9U] MATING TYPE SWITCHING $ [ $ ; et al.566#V / % Aspergillus A. parasiticus A. fumigatus % MAT1-1- $ * I9 1 MAT1-2-1 gene 9 et al.566"QR] et al. 566# N % 566" % % Pseudohomothallism V $ ; [ ? $ * I 9 566#KW ! $ . $ !"7" Bidirectional Mating Type Switching 9% V[ % $ in Saccharomyces cerevisiae, ? $ !"7" I* !"_# * I <!"4#* I9 566# * !"7_!"77N N $ % . . . % Q W !V % % . <!"5#N % ? MAT1-1 MAT ? MAT1-2 MATa W V < !"4#V % % ? $

208 IMA FUNGUS Homothallism in fungi ARTICLE

Fig. 1./ % % Primary homothallism: V MAT in Aspergillus nidulans Pseudohomothallism: @. N $ seen in Neurospora tetrasperma Bidirectional mating type switching: G V $ $ % Saccharomyces cerevisiaeUnidirectional mating type switching:G $ $ V ? % $ V mating system is seen in Chromocrea spinulosa Unisexual reproduction:G $ type partner as in Cryptococcus neoformans

9% $ %MAT1 et al.566_ . % V [ U/V. ? $ ! U/V.5 ? Q MAT1 V ? UV* . \ MAT1 > In S. cerevisiae % I V !"E6 % MAT1 MAT1 ? MAT [ UV* UV*V . 9Q MAT1-1? HML % V 9Q MAT1-2? HMRa % O VUV*% % MAT1 <>/ G? $ > !"E5V\ UV* MAT $

VOLUME 6 · NO. 1 209 Wilson et al.

which a new MAT1 via gene % V % V $ [ O UV* MAT MATa N " I + !"F# U

ARTICLE $ ? Ceratocystidaceae9 IU>!""# et $ N al.5666 et al. 56!7 et al.56!7 *et % 9Q9Q. al56!4 % . " . C. albifundus % MAT1 $ V MAT1-1 MAT1-1-1 MAT1-1-2 MAT1-2 MAT1-2-1/ % % % 9 $ !""E56!5 . . ] . . Unidirectional Mating Type Switching that MAT1-2-1 V % S. cerevisiae .[ MAT1-1 et al. 5666 et V al. 56!7 9 % %% % . \ MAT1-2-1 et Chromocrea, Ceratocystis, al.56!7N [ Glomerella, Sclerotinia !"46U !"45 ZIW !"E_; !"E#9 IU>!""# Ceratocystidaceae V % et al.5666 et al.56!7V % . MAT1-2-1 [ [ . * et $ % al.56!4N % C. Z % Chromocrea spinulosa O seen in Chromocrea spinulosa % . [ $ U !"45V V $ % . % l N % l+ mating U !"45 % C. spinulosa N % . . % V$ % % / V $ %* I9 $ . 566#>% % % U !"45 [ N . l+ $ C. spinulosa % V % l % ? % > [ U !"45 V $ % % Q RN I* !"_#* !"7_ l+ via l V $ $ UNISEXUALITY– AN UNUSUAL FORM OF U HOMOTHALLISM !"45V % % ] I@ !""7 % % ? $ in Neurospora africana V %. ? Neurospora species, Z . N. galapagosensis, N. dodgei N. lineolata + et Ceratocystidaceae + et al. 56!7 al. !""7 Neurospora species

210 IMA FUNGUS Homothallism in fungi

% mat A MAT1-1 / % ARTICLE mat a MAT1-2 ] et V al. !""6 @ I !""6 . V % mat A mat a MAT1 N $ [ ? + et al.!""79% mat A U N. africana * et al. 5664 V V? % $ $ $ $ V $ MAT ] I @ !""7 N % $ . $ $ with the N. africana mat A? . G? ] I@ !""7V $ [ $ V $ % . ? O mat A? ? [ The mating strategy in N. africana $ % $ ; et al.56!_ N % / $ % N % [ Candida albicans V $ $ MAT1-1 MAT1- $ > I` 2 566#N $ C. albicans can $ V $ ? V [ . 9I` $ !""" % $ . $ O $ MAT $ + I`566_V %% et al.56!7KW ! a\a \ / % ? $ . + I`566_ Cryptococcus neoformans * et al. 5664 Z $ C. albicans % W O I 9 56!_ > et al. 56!_ V via . O $ $ a / . $ et al. 566" V [ %% % * et al. 5664 Z $ Bar1N Bar1 a . > C. neoformans @ et al. 566# % a % {"" z . 2.G I + !"#E + $ % @ 5 V % $ % $ . / et al. .$ 566" V .$ a % % 2.G % %% 9 a.a !"#FN ? %% N $ a O $. C. neoformans + ! % et al.!""FV $ aV . O P $ a % N O $ . . % $ mating / et al.566" ? * V $ $ et al.5664 [ Huntiella moniliformis V $ C. neoformans via et al.56!4 Ceratocystidaceae Q %% % Ceratocystis+ et al.56!7 Z $ % $ that in N. africana U/V.5

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[ $ $ $ V . / . H. omanensis, $ % % U/V.! $ U/V.5 $ $ % W $ $ @ $ Cryphonectria parasitica

ARTICLE MAT1-2 % et al.56!4 % Cryphonectriaceae Z $ DothideomycetesU et al!""_G / % $ V % via which $ . N. africana % % [ ? % @. % 9% % [ $ $ Neurospora species O O % $ V % [ % $ via ] et al.!""6 ] I@ !""7 et al.56!7 % % W Cryptococcus neoformans N $ % ? et al.56!7 % de novo W % % V % [ . % $ N $ > et al.56!_ % . 9% % CONCLUSIONS ? % % V @. % . N $ % $ % % @$ ACKNOWLEDGEMENTS % V [ Z % ; < % % / I 9 566# @ V <@V\> W @$ > W G S$ V 9 + % ] N Z % ; N V V > 566F > W 9 % @ / ] [ ? $ ZN<E_"57V] [ V % $ [ > W $ > W % . N % ] + S N% Z % ; G % $ % % % V % REFERENCES ? . O / <2 9 W 566# $ [ the MAT1 ? N[ Sex in Fungi: molecular determination and $ evolutionary implication 9 ` 2 ` V ` Z $ G */[45# 4_7

212 IMA FUNGUS Homothallism in fungi

460[E"6 E"_ Cryptococcus neoformans ARTICLE /*U!"_79 %% . . American Journal of Epidemiology 108[__# _76 Pleurage anserina Mycologia 26[ *< $` [+< [U`56!4X _"5 7!7 % . + >;@ U* ] >* !""7 U O . Ceratocystis albifundus . Neurospora, Plant Pathology[ [!6!!!!\ !5_7" Gelasinospora AnixiellaMycological Research 98[!_6" !_!F * 9 `566#U + ``/<566_G $ N[ Sex in Candida albicans in Fungi: molecular determination and evolutionary implication EMBO Journal 22[5464 54!4 9 `2 `V `G */[_4 4# + /W !"67 @$ Mucorineae Saccharomyces Annals of the Missouri Botanical Neurospora Journal of Garden 31[56_ 5!F Agricultural Research 35[5E" _64 U U !"45 / <+Q!"4# Neurospora Chromocrea spinulosa W; Annals of Botany ? Bulletin of the Torrey Botanical Club 16[77" 7FE 84[!E5 !EE U @V > U/ ` <` > % Genetics 143[#E" #"" W O U9 `56!_] % $ U U] * @S S / * G !""_ S $ Cryptococcus neoformansPLoS Cryphonectria Genetics 9[!66_FEE parasiticaHeredity 70[_E4 _"5 ] `SG %@SGGU *` ` et al.5664 > 2/!"E5U Annual @? Aspergillus nidulans % Review of Genetics 16[7_" 466 with A. fumigatus A. oryzaeNature 438[!!64 !!!4 > 2/V 2!"E6V ] >* ] ` U O * !""6 Neurospora Cell 19[#4_ #F7 crassa / . Proceedings of the National > 566F$Science 311["F6 "F! Academy of Sciences, USA 87[7"!5 7"!F > U W O U * W./% / U O ; et ] >*U O * >+!""6 9 Sordariaceae al. 56!_ Z $ $ [ a mating % de novo in the ?Experimental Mycology 14[5#7 5E" yeast Cryptococcus neoformansPLoS Biology 11[!66!F4_ ] >* @ U* !""7 @ . > UW O U@@ 9 `56!!@$ Neurospora africana Annual Review of Genetics 45[764 7_6 Molecular and General Genetics 244[76! 76" > @U ` /< 566# ] Candida albicans, a 9 `S !""E U . Saccharomyces Annual Review of Genetics 41[ cerevisiaeAnnual Review of Genetics 32[4F! 4"" !"_ 5!! 9 `S 56!5 U . ] MAT switching in QR] GU W 9V < ;@ 566" < % $ Saccharomyces cerevisiaeGenetics 191[__ F7 Aspergillus fumigatus 9 VG U> <* !""# @. Z . Nature 457[7#! 7#7 . Ceratocystis coerulescens, a ; ` 56!4 S% O [ Current Genetics 32[45 4" Molecular Ecology 24[56!E 56_# 9 + O.; `9G N566"@$ ; U@ W// U`2 >G %/Uet al.566# \ $ . Aspergillus U . parasiticusFungal Genetics and Biology 46[!F" !#4 Aspergillus nidulansCurrent Biology 17[!_E7 !_E" 9GU`/<!"""N [ . ; <<!"E#U . [ $ Candida albicansScience 285[ Genetics 115[5!4 5!F !5#! !5#4 ; @@W O U9 `56!_Z $ 2 ` @ G !""# U [ % Annual Review of Genetics 31[574 5#F $ via Eukaryotic Cell 2.G2`!"#FU Filobasidiella neoformans, 12[!!44 !!4" $ Cryptococcus neoformans Mycologia 68[ ; @ @ 2 Z Q O 9< !""# U . E5! E__ Sordaria macrospora are 2.G2`+ `S!"#E< a mating $ Genetics

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147[4F# 4E6 9!"46] GlomerellaXNNN/ ; % ] `/ 9 */ U < <2 + American Journal of Botany ` !"4_ V Aspergillus nidulans Advances in 37[_67 _!5 Genetics 5[!7! 5_E 9!"7"9 $ Biological >+!""5W Reviews 24[7!! 77#

ARTICLE Neurospora tetraspermaMycological +*U USS ZS `G!""F< Research 96[!6_ !!F Cryptococcus neoformans[ 2 W O U @ @ 9 ` 56!7 Z $ Proceedings of the National Academy Advances in Genetics 85[544 _64 of Sciences, USA 93[#_5# #__! *> 2 /* ` 566_ V ;U @ SV [ U` < + [ Saccharomyces +<56!7<>/ " cerevisiaeAnnual Review of Biochemistry 72[7E! 4!F . PloS One 9["5!E6 @ OO G 56!7 W . /] V% > U/ ;U [U` Fungal Biology and Biotechnology 1[# et al. 56!4 Z $ Huntiella moniliformis @ <G ; U+ `566#+ % Fungal Genetics and Biology 80[! " in Candida albicans Q * Q !"_# Q Eukaryotic Cell 6["6# "!E Saccharomyces @ G G !""6 Neurospora crassa . . Comptes Rendus des Travaux du Proceedings of the National Academy of Sciences, USA Laboratoire Carlsberg 22["" !!# 87[7"!# 7"5! G 9 VG [ +< @ `; [ V% .9 O U/ U O > @ .+ ` 566" U` 5666 < MAT-2 . Branchiomma bairdi[ . . Ceratocystis Current . ]G Polychaeta[SabellidaeMarine Genetics 38[7E 45 Biodiversity Records 2[7_ @.9 + U* Q V +] !""" S% V +] Q5666; . % . [ Fungal Genetics and Biology Proceedings of the National Academy of Sciences, 31[! 4 USA 96[44"5 44"# Z ` W 9 !"E_ 9 Sclerotinia trifoliorumPhytopathology 73[4F"M4#5 G.* @ .+ @ +< 56!6 Aspergillus nidulans @ @ / $ % O Fungal Genetics and Biology 47[ #E" #"" + S !"F# V . . "Mycologia 59[5!5 55!

214 IMA FUNGUS