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Rahim, Md Abdur (2014) ASSOCIATION OF THE EXPRESSION LEVELS OF TRANSCRIPTION FACTORS WITH THE PHENOTYPES AND GENOTYPES OF PEACH FRUITS THAT DIFFER IN THEIR QUALITATIVE CHARACTERISTICS. [Tesi di dottorato]

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Abstract (inglese)

Anthocyanins are plant bioactive compounds that control color of many vegetables, flowers, fruit, roots and other plant parts. In peach [Prunus persica (L.) Batsch], color is a key determinant for fruit quality and depends by flovonoids including anthocyanins. These bioactive molecules have potential benefits to human health including protection against cancer, cardiovascular diseases, inflammation and other chronic diseases. The R2R3-MYB transcription factors (TFs) control the expression of genes for the synthesis of anthocyanin pigments with the help of co-activators belonging to the basic-helix-loop-helix (bHLH) and WD40 repeat family. In the peach genome, through this study three MYB10-like (MYB10.1, MYB10.2 and MYB10.3) and three bHLH-like (bHLH3, bHLH33 and GL3) TFs were identified as best candidates to be the regulators of the anthocyanin accumulation in peach. Among these MYB10s, MYB10.1 is the highly expressed gene in ripe peach fruits. The accumulation of anthocyanins in yellow flesh peach fruits is highest in the peel, abundant in the part of the mesocarp surrounding the stone and lowest in the mesocarp. The expression of MYB10.1 and MYB10.3 genes were correlated with anthocyanin levels of different peach parts. They have also positive correlation with the key structural genes of the anthocyanin pathway, like chalcone synthase (CHS), flavanone-3?-hydroxylase (F3H), dihydroflavonol 4-reductase (DFR) and UDP-glucose:flavonoid-3-O-glucosyltransferase (UFGT). Functions of peach MYB10s were tested in tobacco and shown to activate key genes in the anthocyanin pathway when bHLHs were co-expressed as partners. Over-expression of MYB10.1/bHLH3 and MYB10.3/bHLH3 activated anthocyanin production by up-regulating NtCHS, NtDFR and NtUFGT while other combinations were not effective.
To better understand the role of MYB10.1 in anthocyanin synthesis during peach ripening, its coding sequence has been constitutively expressed in tobacco transgenic plants. Its over-expression induce anthocyanin pigmentation in the reproductive parts of transgenic tobacco lines through the up-regulation of transcript levels of many important genes involved in the anthocyanin biosynthetic pathway like phenylalanineammonia-lyase (NtPAL), NtCHS, chalcone isomerase (NtCHI), NtF3H, NtDFR, anthocyanin synthase (NtANS) and NtUFGT. The pigment accumulation was always limited to reproductive parts and never present in the vegetative part like stem and leaves. Besides the regulation of anthocyanin biosynthesis, the ectopic expression of MYB10.1 leads to two different types of phenotype in transgenic as compared to WT tobacco. The strong phenotype, called type-I, is characterized by irregular leaf shape and size, reduced plant height, stamens with reduced filament length and non-dehiscing anthers, reduced pistil length, no nectary gland formation and reduced capsule development but pigmented reproductive parts including androecium, gynoecium and petals. On the contrary, so-called type-II plants, have a mild phenotype with regular plant growth and development, but purple seed coats. Surprisingly, the over-expression of MYB10.1 altered the floral development in transgenic tobacco lines. In type-I transgenic plants, the over-expression of peach MYB10.1 leads to down-regulation of NtMYB305, a gene similar, but not orthologous, to MYB10.1, which is required for floral development. Moreover, MYB10.1 up-regulates the expression of jasmonic acid (JA) biosynthesis (allene oxide synthase, NtAOS) and signaling (NtJAZd) pathway genes as well as 1-aminocyclopropane-1-carboxylate oxidase (NtACO), a key gene in ethylene synthesis. Furthermore, NECTARIN1 (NtNCE1) that is known to be involved in nectary gland formation, was repressed since its transcription is regulated by NtMYB305. On the contrary, type-II transgenic plants showed opposite results for NtMYB305, NtAOS, NtJAZd and NtNEC1. Therefore, it can be concluded that the over-expression of peach MYB10.1 in tobacco not only regulates flavonoid biosynthesis in the reproductive parts but also affects (directly and/or indirectly) other process like vegetative and reproductive development. These new findings will promote further investigation to elucidate more diverse role of R2R3-MYB TFs. Furthermore, they will promote more and better insights on the regulation of anthocyanin biosynthesis and will aid fruit breeders to introduce new peach cultivars with higher antioxidant level as well as colored fruit.

Abstract (italiano)

Le antocianine sono composti vegetali bioattivi che controllano il colore di molte verdure, fiori, frutti, radici e altre parti di piante. Nel pesco [Prunus persica (L.) Batsch], il colore è un fattore determinante per la qualità del frutto ed è regolato dai flovonoidi tra cui gli antociani. Queste molecole bioattive hanno benefici effetti per la salute umana, compresa la protezione contro il cancro, le malattie cardiovascolari, infiammatorie e altre malattie croniche. I fattori di trascrizione (TF) R2R3-MYB controllano l'espressione dei geni della sintesi delle antocianine con l'aiuto di co-attivatori appartenenti alle famiglie basic-helix-loop-helix (bHLH) e WD40. Nel genoma di pesco, attraverso questo studio sono stati identificati tre geni di tipo MYB10-like (MYB10.1, MYB10.2 e MYB10.3) e tre bHLH-like (bHLH3, bHLH33 e GL3) quali migliori candidati ad essere i regolatori dell’accumulo degli antociani in pesca. Tra questi MYB10s, MYB10.1 è il gene più espresso nei frutti maturi. L'accumulo di antociani in frutti di pesco è più alto nella buccia, seguito dal mesocarpo attorno all’endocarpo e minima nel resto del mesocarpo. L'espressione dei geni MYB10.1 e MYB10.3 è correlata con i livelli di antocianine delle diverse parti della pesca. Essi sono correlati positivamente con i principali geni implicati nella via di sintesi delle antocianine, come la calcone sintasi (CHS), flavanone-3?-idrossilasi (F3H), diidroflavonol 4-reduttasi (DFR) e UDP-glucose:flavonoid-3-O-glucosiltrasferase (UFGT). Il ruolo dei MYB10 di pesco è stato testato in tabacco e si è visto che nel caso vengano co-espressi con bHLHs si attivano alcuni geni chiave nella via di sintesi delle antocianine. Solo le sovraespressioni di MYB10.1/bHLH3 e MYB10.3/bHLH3 attivano la produzione di antociani regolando i geni NtCHS, NtDFR e NtUFGT mentre altre combinazioni non sono state efficaci, confermando anche dal punto di vista funzionale la maggior importanza di MYB10.1 rispetto a MYB10.3 e MYB10.2.
Per meglio comprendere il ruolo di MYB10.1 nella regolazione della biosintesi delle antocianine nella pesca in maturazione, la porzione codificante del gene è stata fatta sovra-esprimere in modo costitutivo in piante transgeniche di tabacco. La sua sovra-espressione induce la tipica pigmentazione dovuta ad antocianine nelle parti riproduttive di linee di tabacco transgeniche attraverso l’induzione dei livelli di trascrizione di molti importanti geni della via biosintetica degli antociani, come fenilalanin-ammonio-liase (NtPAL), NtCHS, calcone isomerasi (NtCHI), NtF3H, NtDFR, antocianina sintasi (NtANS) e NtUFGT. L'accumulo dei pigmenti è sempre stato limitato alle parti riproduttive e mai a quelle vegetative come fusto e foglie. Accanto alla regolazione della biosintesi di antocianine, l'espressione ectopica di MYB10.1 conduce a due diversi fenotipi nel tabacco rispetto al WT. Nel fenotipo forte, detto Tipo I, si ha forma irregolare della foglia e dimensione della pianta ridotta in altezza, ridotta lunghezza del filamento dello stame, antere non-deiscenti, ridotta lunghezza del pistillo, nessuna formazione di nettari e ridotto sviluppo della capsula; inoltre, parti del fiore, tra cui androceo, gineceo e petali, sono più pigmentate. Nel fenotipo blando, detto di tipo II, la crescita delle piante è regolare così come lo sviluppo e la maturazione del fiore; tuttavia i tegumenti di ovuli e semi sono viola. Sorprendentemente, la sovra-espressione di MYB10.1 non solo ha fatto aumentare la sintesi di antociani, ma ha anche fortemente alterato lo sviluppo fiorale in linee di tabacco transgeniche. In piante di tipo I, la sovra-espressione di MYB10.1 porta alla riduzione dell’espressione di NtMYB305, gene necessario per lo sviluppo del fiore, simile, ma non ortologo, a MYB10.1. Inoltre, MYB10.1 induce l'espressione di geni coinvolti nella biosintesi (NtAOS) e segnalazione (NtJAZd) dell’acido jasmonico (JA), oltreché di quella dell’etilene (NtACO). Inoltre, NtNCE1, gene di cui è noto il coinvolgimento nella formazione della ghiandola del nettario, è represso nella sua trascrizione, che è noto essere regolata da NtMYB305. Al contrario, piante transgeniche di tipo II hanno mostrato risultati opposti per NtMYB305, NtAOS, NtJAZd e NtNEC1. Pertanto, si può concludere che la sovra- espressione di MYB10.1 di pesco nel tabacco non solo regola la biosintesi dei flavonoidi nelle parti riproduttive ma colpisce anche altri processi come lo sviluppo vegetativo e riproduttivo. Questi nuovi risultati promuoveranno ulteriori indagini per chiarire i molteplici ruoli dei TF di tipo R2R3-MYB. Inoltre, le maggiori e migliori conoscenze sulla regolazione della biosintesi degli antociani aiuteranno i coltivatori a produrre frutta più colorata introducendo nuove cultivar di pesco con più alto livello di antiossidanti.

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Tipo di EPrint:Tesi di dottorato
Relatore:Trainotti, Livio
Dottorato (corsi e scuole):Ciclo 26 > Scuole 26 > SCIENZE DELLE PRODUZIONI VEGETALI
Data di deposito della tesi:13 Gennaio 2014
Anno di Pubblicazione:31 Gennaio 2014
Parole chiave (italiano / inglese):Anthocyanin, bHLH, Prunus persica, MYB, tobacco, trangenic plant, transcription factor
Settori scientifico-disciplinari MIUR:Area 05 - Scienze biologiche > BIO/01 Botanica generale
Area 07 - Scienze agrarie e veterinarie > AGR/03 Arboricoltura generale e coltivazioni arboree
Struttura di riferimento:Dipartimenti > Dipartimento di Agronomia Animali Alimenti Risorse Naturali e Ambiente
Codice ID:6244
Depositato il:24 Apr 2015 16:17
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