In Helianthus × multiflorus il gene CYC2c è espresso ectopicamente in corolle dei fiori del disco modificati in fior

“ray-like”

Nel girasole, HaCYC2c è espresso in fiori del raggio ma non in fiori del disco (Chapman et al., 2008; Tähtiharju et al., 2012). Al contrario, nelle piante mutanti Chry2, il gene è espresso ectopicamente a causa dell'inserimento di un ET, con formazione di fiori del disco zigomorfi (Chapman et al., 2012; Fambrini et al., 2014a).

È stato analizzato il pattern di espressione dei geni CYC2c mediante real- time RT-PCR (qPCR) in H. annuus, H. decapetalus e H. × multiflorus. È stata verificata una corrispondenza con i risultati riportati precedentemente (Chapman et al., 2008; Tähtiharju et al., 2012). Come previsto, CYC2c è espresso nella corolla di fiori del raggio in H. annuus e H. decapetalus. Mentre nei fiori del disco i livelli di espressione sono estremamente ridotti (Fig. 18). Al contrario, in H. × multiflorus il gene CYC2c ha mostrato un evidente ed alto livello di trascrizione sia in corolle dei fiori del raggio esterni (RFE) che in corolle dei fiori del raggio interni (RFI) (Fig. 18); stessa espressione manifestata dalle inflorescenze dbl e Chry2 (Chapman et al., 2012, Fambrini et al., 2014a). È probabile che la sovraespressione del gene H×mCYC2c abbia determinato il passaggio dei fiori del disco, da attinomorfi a zigomorfi, con la produzione di un’infiorescenza di tipo ligulato, atipica per il genere Helianthus.

Fig. 18: Espressione del gene CYC2c. Livello di trascrizione dell’mRNA del

gene CYC2c in corolle di fiori del raggio (RF) e fiori del disco (DF) di Helianthus annuus (Ha), Helianthus decapetalus (Hd) e Helianthus × multiflorus (H×m). L’mRNA è stato estratto da corolle di RF di Ha (HaRF) e H×m (H×mRFE), e da corolle di DF posizionate nel secondo cerchio dell’infiorescenza di Ha (HaDF), Hd (HdDF) e H×m (H×mRFI). I valori relativi di trascrizione sono stati calcolati mediante qRT-PCR. È stato usato mRNA di radice di Helianthus annuus come campione di riferimento e normalizzato a quelli del gene ribosomale Ha-18S. I dettagli sono forniti in Materiali e Metodi. Il grafico mostra la media (± DS) di tre repliche biologiche (n = 3). Lettere uguali indicano assenza di differenze significative in base al test di Tukey dopo analisi della varianza (ANOVA P ≤ 0,05).

4. Conclusioni

In una conferenza tenuta a Chiswick, in Inghilterra e dedicata alle Asteraceae ed in particolare alle specie perenni di Helianthus spp. DeWar (1891) affermava “It … seemed probable … that Helianthus × multiflorus is a garden hybrid of Helianthus annuus and perhaps Helianthus decapetalus, and that it has been produced several times. Helianthus annuus is the only known sunflower besides Helianthus × multiflorus which assumes double forms (ligulate inflorescence), and its hybrids might inherit this tendency …” È probabile che questa affermazione sia scaturita dall’osservazione del mutante Chry di H. annuus, già conosciuto in Europa da circa 300 anni (Heiser, 1976).

Abbiamo dimostrato che H. × multiflorus varietà “Soleil d’Or” presenta un’infiorescenza ligulata, esteticamente molto attrattiva, originata probabilmente dall’espressione ectopica del gene CYC2c, come conseguenza dell’inserzione di un elemento CACTA difettoso nella regione promotrice al 5'. Non si può, tuttavia, escludere che nella formazione di queste nuove ed insolite infiorescenze di aspetto ligulato possano essere coinvolti anche altri geni CYC.

Potrebbe essere interessante approfondire il ruolo dell’inserzione dell’ET CTEHM1 mediante una analisi approfondita delle varietà di H. × multiflorus che presentano infiorescenze di tipo raggiato (es. “Major” e “Meteor”); come sarebbe necessario indagare sulla regolazione, soprattutto su base epigenetica, di questi elementi trasponibili.

Ciononostante i dati raccolti in H. × multiflorus forniscono un ulteriore conferma sul coinvolgimento degli ET nell’evoluzione dell’architettura delle

infiorescenze; in particolare sullo sviluppo di una infiorescenza atipica nel genere Helianthus in cui l’architettura di base è di tipo raggiato.

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6. Supplementare

Fig. S1: Elenco delle sequenze amminoacidiche dei fattori di trascrizione (FT)

codificate dai geni CYCLOIDEA (CYC) e utilizzati per l'analisi filogenetica. Le abbreviazioni delle specie sono le seguenti: Antirrhinum majus (Am), Gerbera hybrida (Gh), Helianthus annuus (Ha), Helianthus decapetalus (Hd), Helianthus × multiflorus (H×m) e Senecio vulgaris (Sv). I numeri di accesso in GenBank sono inclusi tra parentesi. Le sequenze amminoacidiche dei FT sono state allineate con il metodo ClustalW (Thompson et al., 1994) presso il Centro di Bioinformatica dell'Università di Kyoto (http://www.genome.jp/tools/clustalw/) e sottoposte ad analisi filogenetica. L'analisi filogenetica è stata eseguita utilizzando i programmi PHYLIPs, PHYLogeny Inference Package, versione 3.67 (Felsenstein, 1985). È stato utilizzato il programma Proml che implementa il metodo di massima verosimiglianza per le sequenze degli amminoacidi. È stato scelto il modello di probabilità Dayhoff di cambiamento tra amminoacidi. La sequenza amminoacidica di Oryza sativa PCF1 (OsPCF1) è stata utilizzata come “outgroup”. Come supporto per l'albero, un'analisi di “bootstrap” con 100 repliche, è stata eseguita dal programma Seqboot e un albero di consenso è stato ottenuto dal programma Consense.

>OsPCF1 (XP_015633587) MMASSSDLILYNLVPAQPLNPSAIPNPNPDLSIAAAEPPSSDGATPRRVRPRKSPSSSDRHSKVAGRGRRVRIPAMVA ARVFQLTRELGHRTDGETIEWLLRQAEPSIIAATGTGVTPEEAPPAAVAIGSSSVAAAAAAGGHGGAFVHVPYYTALL MQPPNADEPPMASAASASGTTAADENNN >AmCYC (Y16313.1) MFGKNTYLHLPQVSSSLHSRAATSVVDLNGNEIQLHDMLSGHYLTTANAPVLESTALFNNNNNFNHDVVNGLNRDPSP TFPTKQAVKKDRHSKIYTSQGPRDRRVRLSIGIARKFFDLQEMLGFDKPSKTLDWLLTKSKTAIKELVQSKSTKSNSS SPCDDCEEVVSVDSENVTDHSKGKSLKANNKCKEAMDSHQAAAKESRAKARARARERTKEKMCIKQLNEAIVLRNHQF EVSGTREAFVHPVFGFHQQNYGNASHENWDQSNLSSQSNQLCAILNQHKFIN >AmDICH (Q9SNW8.1) MFNKKRYLQYPQVPPSLNPRASTSVVSLNGNEILLHHHDVISGYYLASNPQNLEPDALFNGFHHDVGGTNGDPLVLAN TLAKKHTPKKDRHSKINRPQGPRDRRVRLSIGIARKFFDLQEMLGFDKPSKTLDWLLTKSKEAIKELVQSKSSKSNIS NSPSECDQEVLSADLPYIGSSKGKAAVGLNSNKCKGGRDAVDLAKESRAKARARARERTKEKMCIKQLNQERNKSYEW NPSVLFQSKSSQQFEVSGPSTNYEELNQESIMIKRKLKQNHPSMFGFQPENATENWDYYSNFTSQS NQLCAILDQHKFIN >HdCYC2c MNLYLNFFVNYYCYCFFHIQITMFSSNPFPQIPSSIHVSHPFNSFFDLEKNDVYLNHHEDNNNPFVSGDPFFHSYNNF

LSIDIARKFFVLQDLLGFDKASKTLDWLFTKSKKAIKELIEETKHSSSSTVSTNQCEMAFLETIQGGSGSDEDKGQKM SALKFLDGEKKKMTQKCKSGFHASLARGQSRAEARARARERTKQKLRIKELDNDLKKVPDDYPCHALSPSNTTLQSNS WVQFESQSDYNDILHGSMLEQRFSVSSSTLYTYNHNFVVSDEWISQISSKGMPKFKVIHEEQSHRAMI >HxmCYC2cA MNLYLNFFVNYYCYCFFHIQITMFSSNPFPQIPSSIHVSHPFNSFFDLEKNDVYLNHHEDNNNPFVSGDSFFHSYSNF APQPSALPPVTDYMPSNTQELDSQNQLLDHEGSGLQYCDNYSDLLESVVYPSKKKVVTSKKDGHSKIHTAQGPRDRRV RLSIDIARKFFVLQDLLGFDKASKTLDWLFTKSKKAIKELIEETKHSSSSTVSTNQCEMAFLETIQGGSGSGSDEDKG QKMSALKFLDGEKKKMTQKCKSGFHASLARGQSRAEARARARERTKQKLRIKELDNDLKKVPDDYPCHALSPSNATLQ SNSWGQFESQSDYNDILQGSMLEQRFSVSSSTLYTYNHNFVVSDEWISQISSKGMPKFKVIHEEQSHRAMI >HaCYC2c (EU088370.1) MNLYLNFFVNYYCYCFFHFQITMFSSNPFPQIPSPIHVSHPFNSFFDLERNDVYFNHHEDNNNPFVSGDSFLHSYSSF APQPSTLPPVTDYIPSNAQELDSQNQLLDHEGSGLQYCDNYSDLLESVVYPSKKKVAISKKDGHSKIYTAQGPRDRRV RLSIDIARKFFVLQDLLGFDKASKTLDWLFTKSKKAIKELIEETKHSSSSTVSTNQCEMAFLETIKGGSGSDEDKGQK MSALRFLDGEKKKMTQKCKSGFHASLARGQSRAEARARARERTKQKLRIKELDNDLKKIPDDYPCHALSPSNTTLQSN SWGQFESQSDYNDILHGSMLEQRFSVSSSTLYTYNHNFVVSDECISQISSKGMPKFKVIHEEQSHRAMI >HaCYC2d (EU088371.1) MFPSNLVQPHPSSIHVFPSSSSLFDLEKDGVHFHHHYHHHHNNQFINGDCFFHGYNTAPPPVLTDNIKGSGFHYSDDH NSLLEAVIYPSKKKVANMKKDGHSKIHTAQGPRDRRVRMSIDVARKFFYLQDLLGFDKASKTLDWLFTKSKIAIKELV EEMKHCSSPGATDQCEDVFQETLKRSGEELDKGEKKKSAPKFVDAKKKRKYRSVVDVNQSRAEARARARERTKEKLQS KKLDDESKKVPHESSFWSQIESKKEFNERTNESIMEPKIPMSSSVVYGYQHNFLVPNDSSSQIKYTSFLNLH >HaCYC2b (EU088369.1) MLSSSTNAFSPSKSFFDNQKDNHDFNYPQEIDNPFISTDFVCHTYTGPAHVPQVTEEATTLEQGFVLQHQFSEDHDDL LESVVSSYKKKIVTTKKDGHSKICTAGGLRDRRVRLSIDISRKFFCLQELLGFDKASKTLDWLFTKSLTAIKDLVEQT NHSSSSALTDESNAKFLESIKGDSDDENTQKEKSLLKRVDGKGKKGTQRAKVRSQESFSREQSRAEARERARKRTKEK NRAKKFDDCESGLECSMDQETSNQSKDSSFQVRA >HaCYC2e (EU088372.1) MSSSNPFPQVTSSSHYVFPPSNPFLDHEKDDLYINNFNPFLSGDCFPDNENNTTRKQDLVEEVGLEEECDENDHLFWS VVKNNKGSNKDHHSKIHTAQGPRDRRVRLSIEVAKRFFYLQDLLGFDKGSKTLDWLFNKSKNSIDELIKRKKESSSSS LTDQLEVVFLETDEQAKGRKNKCGEGKRKKIARKCTSGVTVNQSRAEARARARERTKEKMNVKKLDKESKSAPVDCCS SNLTLQSSFWNSFESQNDYNDKIGEALLEDDISLLYSYQHNLAVSNDSSSEFNCMPDVHEHWGGDCAAI >HaCYC2a (EU088368.1) MFSSSNPFTSSIHGLPPSSNIFLGQEKDGVYFNPFFTGEYCSFDPPIKESELQNCENNNNNDNNNNNNNNHLKLSDSV ISPSRKRVAASKKDRHSKIFTAQGPRDRRVRLSIDISKKFFGLQDLLGFDKASKTLDWLFTRSKAAIKDLVEEMKHTS SSTLSDQCEEVFMEKGSDYLHKNGRKKKPAAKCIKRKKKTAQEHKSGIHLNLAARSQLRAEARARARERTLEKLRMKK LNNDVKNVVGDSDYCFLQTTQIQSQSNHEVKIGEALTQGKLPKPYSLLYNSQHNYVYSKGSRSHFKNCAH >GhCYC7 (JN190061.1) MFNSSNPFPSSIHGFPPPSASFFGHENDDVYFTHHPFLVGDCSFHDGNVIAPPPPPAGNIKASGSQYCEDNNNNNNNN NNDNHQLQDSVISTSRKRVAASKKDRHSKIFTAQGPRDRRVRLSIEISRKFFGLQDLLGFDKASKTLDWLFTKSMTAI KDLVQEMKHGSSSSLSDRCEVVFMEDEERKNRGLKKKKPLAKCVNGGKRKKKMTQKHKTGFHVNLATRNQLRAEARAR ARERTIEKLQIKKLHHDSKNVVLDHNNYCYDLQSRTTNLFYTSQQNFVDSKDSSCLIKHSHSPKFIQDQQGDHNPLI >GhCYC4 (EU429305.1) MFSSNPFQDLPPSIHVFPPLDPNIDHDRDDAYYNYHHSNDPFISEDCLLHSYNDISPSSPLDMEKINKIKQDFLRQQR QIYEGSGLQYCEDRDDLLDSVVSSSKKKMATSKKDGHSKIYTAGGPRDRRVRLSIGIAKKFFCLQDLLGFDKASKTLD WLFTKSKTAIKDLVEEKKHCSSRSSTVTEVSFLETINGELDEEDKGQKKKCVDGKPKKMKRKSNSQLQANLARDQSRA EARARARERTREKMRIKKLGNEYLETLVPENFDYHVSPSNPTLQSSCWSNFESQSDYNDITWESIMDPVKYADLPNFG AFPEQQGDHATYDLDLL >GhCYC9 (JN190063.1) MFSSNPFPQPPYVLSPPNTYFHHEEDDGYFNYHRSNTDADDPFISGDCFLPSCNDLAPPSPLVMENLTKNKQDFLEIH DDLLDSVVSSSKKKMPTSKKDGRSKIYTAGGPRDRRVRLSIEIARKFFCLQDLLGFDKPSKTLDWLFAKSKTAIKDLV

EETKNCSSRSSTVTEVSFLEAINGGLDEEGKGKTKKSVDGKSKKMKRKSNSQIQVNPARDQSRAEARARARERTREKM RVKKLDDELMSLVS >GhCYC2 (EU429303.1) MFSTNPFQELNSSSHVFPPPNSFLDHEQDDLFHHRSNHPFVSGDSFFGAMADFKDSGGQHQLFSGSGLEYNDEYNNTL ESGVSKMKNKKKISKKDHHSKIDTAHGPRDRRVRLSIDIARKFFCLQDLLGFDKASKTLDWLFTKSKPAIDELLEGTK NSSSSTVTDQSEVGVLEIINGGSNEEDKAKKKKTTPNCVDEKGKKTTRKHKSGSFPVNQSRAEARARARERTKEKWHI KKLDDGSKKVLNECRCPVSDSNLTLQSSVWSPMI >GhCYC5 (JN190059.1) MFSSNPFPQIPNSIHVSPHPSSSFFNHEENGIYLNHHGHNSNLFVSGDCFFHAYNSIAPPASPPPLLTNYMATNTQDL FRQQQQLSEGSGFHSCDNHNDLLESVIYPSKKKRETSKKDGHSKIYTAQGPRDRRVRLSIEIAQKFFVLQDLLGFDKA SKTLDWLFTKPKTAIKELVEETKQCSYSTVSTDQCEVVFQETFKGRSYEDKCQKKKSPLMCVEGKRKKMAQKYKSGYH VNHARDLSRAEARARARERTKEKLRVKMLDDKSKKVPGDYYCPSNLTIQSSFWSPNESQSEYIDILRGATMEQRFPMP SWIPYSYQHNLLVSDSSGSKVKYTSFPNNAMI >GhCYC3(EU429304.1) MFSSNPFPQLPPSIHVFPPQNSFFELEKDGDYLNNHDHNNQFVSGDCFFNAYNNIAPPPVTDNITEHQNQQLSKGSGL QYCDDNNHFLESVAYSSKKKMGTSKKDGHSKIYTAQGPRDRRVRLSIEIAQKFFVLQDLLGFDKASKTLDWLFTKSKT AIRELVEEMKHCSSSSVTDQCEVVFQETMKEGSDEEDKGQQKKSAPMCLDGKKKKTTRKYKSGVSRAEASARARERTK EKLHIKKLDDDSRKVPDDCYRPGSPSNLTLQSSFWSRTESQSDYNDRTCESVMELKMSMPFSALFGYQHNHLSNDLTT QTKYTSFLNLQ >SvRAY1 (FJ356698.1) MFSSNPFHQIPSSIHVFSPSNSFFDHEKDGGHFNHHNDISQFVSGDCIFDSYNNLAPPPVIADNIMVRQQDQLSKDSG FHFCDDKSNLLESVIYSSKKKMTNLKKDGHSKIHTAQGPRDRRVRLSIEVARKFFSLQDLLGFDKASKTLDWLFTKSK TAIKELVEEMKHCSSSGATDHCEVVFQETIKRGSDEEDKGQKKKSGTKYIEGKKKKMSRKYKSGVDVNQSRAEARARA RERTREKLQNKKLDDDSKKVPNNCYRPASPSNLTLQSSFWSEIESQNELNHRTGESTMVPKVPMPSSMFFAYQHNLLE SNDLSSQIKYTSFLNLH >SvRAY2 (FJ356701.1) MFSTNPFSHLASSIQVSPPSNSFLDFEKEEPYFYHHHQSSNPFDPADCLPAREKFSTSDHNLLEELGLEAYDEYTHFE SDTKKNHSPKKDHHSKIHTAQGTRDRRVRLSIEVAKKFFYLQDLLGFDKASKTLDWLFDKSKIPINELVQTKKQSLSS TVSDQSEVVFGETVEEGPDEQDKGQKKKKCVDRKRKKITRKYQSGSPVNQSRADARARARERTKEKLNSKKLDIESKS VYNGDCCSNIWSVIESDKIEESLIEKEISLPYPMLYSYQPNLTVSNDSSSSSSKFNNNSFPNFTVTYE >SvRAY3 (KT722935) SKTLDWLLTKSLTAIKDLVEETSQCSSSIITTDQYSKVEFLESIKGTSDEDKNCQKKKSLCVDEKKKKMTQRCSKAGL IQDNLTRDQLRAEARARARER >SvCYC4 (KU663021) SDQSEVVFGETVKGEKKKKNVEGKRKKITRKYKSGSPVNQSRVEARARARERTKKKLKLKSSMTNPKVFMVTLVLQIQ QSSQAFGVQLNQKR >SvCYC5 (KU663022) RVRLSIEIAKKFFYLQDLLGFDKASKTLDWLFSQSKIPINELVQSKKQSSSSSVSDQSEVVFGETVKGEKKKKNVEGK RKKITRKYKSGSPVNQSRVEARARARERTKKRLNIKKLDDESKSIHGDSCSSNSTIQSSFWSAIESEKIDDYMSSSML YSYQPNLSVLSVSNNSSSNVTGLPKFTLVHEPQGDRAMI >SvCYC7 (KU663024) DKPSKTLDWLFTKSKIAIKDLVEEMKHTSSSTLSDQCEQVFMEKGDYLEKKGKKKKPVAKCVKRKKKITQKQKSGVQV SLGARSQLRAEARARARVRTIEKLRIKRLSNDEKNVLGDNYYYYLLSTTGNQVQS >GhCYC1 (EU429302.1) MYSSSNTNGNTSSTSSNQPKTSFFCNYNNEYHSKPSQEHYHPPSSSSFHFHYPHYTPLEDEAIFRELFLQQQTFSNDH NYQNTVVVAHEQQPCKTTTNLQCTMEKCSYNKGKYVTNDGEDHFNSHQFLKNSGLTKRSSKKDRHSKIDTARGPRDRR MRLSLDVAKQFFRLQDMLGFDKASNTVEWLLMKSKAAIHYLLPQQLNKSCSLMCVSNSASSASECEVLSGIDDQSMEK

TGDDLVITMDKVKSSYCSGNIEKKRVVRGVRRSAYIDHSLAKETRERARARARKRTNEKRNNKLGWDQYPCLDHGMDQ NLNRLGSWIPFVENQVQTNDQPENRSSHFQWKQGVVGDNSSVMTGNWSPSFLFNNQHNAGPSHEHQFRDFQILGKPW >GhCYC10 (JN190064.1) MYPSLDNNGSKQPRTSFFFNYNNDIINHSRTPQEHYCHPSSFSSSFHLPSPHYIPLEDEAVFCEFFQQQQFFSNDHNY HNTIINLAQEQSTNEMTANVESTMGECSNNNGVQIATNDGDDDHYEFNTHVKPENSSPIRKKSSKRDRHSKINTARGP RDRRMRLSLDVAKKLFGLQDLLRFDKASKTIDWLITKSKTAIQELLPDPSCSFMGVSNSASSTSECEVLSGIGDLPID DQATTKNKAKSSSSSRKKKVKITRVRRTANLHHPLAKATRERARARARERTVEKRNNKLGGGQDSNFKPCMDQATDQD VNRVVSWSPFEERQVQSIDQHKQGFVGDTSSLMMTGNWSPSYFNYQHSAGLTHEHQFNDFQIIGKGWEGNNN >HaCYC1b (EU088367.1) MSSSNLSNYHNDSFHSKTPQEPYHTPSCSSFLLPSPIYTPLEDESAFCELFQQQQFLSNDHNHNLLVHEMSNNVGECS NKKGKITTNDGDDDHCDVNTHVEPDKDSPRKIPSNRDRHSKINTAQGPRDRRMRLSLDVAKKFFGLQDLLGFDKASKT IDWLLTESKTAILDLLPDHSCSFMDVSNSTSSTSECEVVSGIGDQFMVKTGDDQATTKTKAKSSSGNNKKQKEKITRV RKCADLQHPLAKETRVRARQRARERTIQKQNNYYNHGVGQDSTFGPFLDQVMDQDVNQLGSWSTFQESPHKSIDQVNQ MSSNFQYNRGFVGDNSSLSMTRNCTPSYFFDSQHSTGVMDEHQFDDLQKKKKASEGNNT >HaCYC1a (EU088366.1) MPPSTSVFGNCYNDNNPSKTPQEYYNTPSSSSFLLSSPIYIPLEDEAVFCDFFQQQQFFSNDYNHHNSMVVPHEECSN NNGQAVTNEGDDDHCIFNTHVEPESSSPRKRPRKRDRHSKIDTARGPRDRRMRLSLDVAKKLFGLQDLLGFDKASKTV DWLLTKSKASILELLPDRSCSFMDVANCASSTSECEVLSGTADHSLVKTGDDQATTKNKAQSSSGSSKKKKEKVTRVR KCVDFHHPLAKATRERARERARERTIEKRNNKRGGGTQDSNFRSCLDQAMDQDVNQYGSWSTIQENQRQNIGQMSLDF QVHQGFVGDNSSLLMTSNWTPSYLLNYQHSSGVIHEHQFNGLEMNVKAWEGNNN >SvCYC6(KU663023) PSKTLDWLITKSETAILELFPDHSCSFMGVSNSASCSSECEVLSGARAADQPMVVIETGDDQETMKNKENXIFESIKK PQRKSSXKNLGNVADFHHPVAKATRERARERARERTTEKRNNKHKSIGRLEQAMDQNVTQLRSWSPFEENRLEPIDQV DQMSSNLHIKQGFVNDNSSMFMMGNWNPSYMFNYQHTSGLTREH >GhCYC6 (JN190060.1) MDYVFSSSNSETMLVNSNNEANLENLDSRHEEDVPLFLQFPSPFLDDISTTSILPNHHLHHLTSDQRETITTEPPPSP SKTKRVRKKRSAGKKDRHSKIHTAQGLRDRRMRLSLQIARKFFGLQDMLGFDKASKTIEWLFCKSKKAIEEVTESLKS QNTTQSAIRECESPLSDCEVDSGLEFNAPSNKGKQLKLQDEIKDSGNSRKPIESNLLARESRDKARARARERTRERMM ISQLEKSKQMFEANPTDDFGKLQLGFSTNPNSHYMDESSSSPLEYTGSHHFLDQAQVDNMPQNAAASSSTYFSTFDYN KNFANPPAGWLNSSNTFLGFLGGWDSENFRMEAENYGILPNIAPFTGDIHGQNPSSIIISSSNLVHFQTENIRK >GhCYC8 (JN190062.1) MEHVCSSRKDKCLLIDPNKEDGHEILNSSHEEPPLFLHFPSPFFDDTSTTPILANNYTHHHSSPNKPLNMTEEPSPSE SKRVKKRSVGKKDRHSKIHTAQGLRDRRMRLSLHIARKFFDLQDLLGFDKASKTIEWLFCKSNKAIKEVAESFNSQNT NQSLSGEIETMDSLLSESEVGLTSEIETATNKGENSKDDEIKNRESRKRIQTNSLAREQARERARERTREKMMIKDLE KSKQLFKRNPKDEFYKLGLGYSTSPNYQNVDELGYTSSPFDPIQQESSNPSLEPSSTQHLLRRPQSVNNVDSFENYSG GIVGSSRYCSILSNHNPPASWLNSSNGFLGISGGWDADSFITESYNYGIVPSTAPLTGDIYEQTLAYFSCLQPIQSQN EDK >HaCYC3a (EU088373.1) MEHVGSSNIHQEEPPLFLDFPSPFFDSISTTPIQTNDLSTPNKPLNVTEPSPSESKRLKKRCGGKKDRHSKIHTAQGL RDRRMRLSLHIARKFFDLQDLLGFDKASKTIEWLFSKSNKAIKEVSENFSPQHTNTSMNDQDTSEVGLTSENKTATNN LDNSREEDIKNRESRMKRIDPNNSSVRETRDQARARARERTRERLMIKELEKSKQLFGRNPNEEINKLGLGYLPSSNN NNAEELGFITSSPSEPIQQESSTSSLENSHTHHLLPVSQSNKQTNVSSNIPFKVGYENDETRANIFSIPGDREAAFTV PEWQSTTNNLNIFENYSGSVVGTSTTCSTLTNHNPPAGWLNSWNGFLPGEWDADNFVNEYYNYGIVPTTVSLTGDIYD QNPSSFILSSTTNILHVQSQNQGK >HaCYC3c (EU088375.1) MEYAYLSRNNEAILLPSNNEFNHENPESNHEEDAPYFRQFQTPLLDEILHNHQQHMTYSTANKPPPSPSKTKRARKQR SAGKKDRHSKIHTAQGLRDRRMRLSLHIARKFFGLQDMLGFDKASKTIEWLFCKSKKAIDEVTESVKSQNTTRSACNE NIESCESPLSDCEVHSGIEFDAASNKGKQLKVQDEINDSGSSKTATKSDILARELRGKARARARERTRERMLITNPEK SKQMFGANPNDDLDQLQLGFSSNPNNNHYIEDSSSTSPNEYSGTHHFFDHTQLNNNIAQKTEDYLGTAAASSSTYFSG YGYNKSFANPPAGWLNSSNTFLGFLGGWDSENFTENYGFLPNLAPLTGDIHGQNFSFIPPSNLVHFKTQNQRD

>HaCYC3b (EU088374.1) MAYENNESNHENLDSNYEEDAPLYGQFPSPFFDDIPHNHHQHKKSNTTDEPPPSTPKTKRVRKKRSAGKTDRHSKIHT AQGLRDRRMRLSLHIARKFFGLQDMLGFDKASKTIEWLFCKSKEAIDEVTESVKSQNTTQSVAGRENIESCDQCPDSD CEVDYNSGIEFYATSNKGVKQPTESELLARELRDKARARARARTKERMKSKQVYEPNPNDDYDQPQSGFSTNPNDDYD QPQLGYSENPNNPYTEGSSSSHLEYSGTHHFFDPTHLNNVAPSSSAYFSGYGYNKSFENPPAGWLNSSNTFLGFLGGW DSEHFTENYGHYYPNLAPTTGDEYHGQHFSSVVIPPPYNLMHYQTQYQRD

7. Ringraziamenti

Giunto alla fine di questo percorso ringrazio il Dipartimento di Scienze Agrarie, Alimentari ed Agro-ambientali per tutte le conoscenze che mi ha permesso di acquisire.

Desidero ringraziare il Professor Andrea Cavallini, il Professor Claudio Pugliesi e il Professor Piero Picciarelli per i loro preziosi consigli, la disponibilità e la pazienza dimostratami, contribuendo alla stesura di questa tesi.

Un grande ringraziamento va alla mia famiglia per l’aiuto ed il supporto costante di tutti questi anni.

Ringrazio la mia compagna Irene per il fondamentale sostegno morale e l’incoraggiamento datomi di fronte alle difficoltà incontrate.