Υπογένος Piperites (Fr.) Kauffman 3.1.1

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Υπογένος Piperites (Fr.) Kauffman 3.1.1 ΓΕΩΠΟΝΙΚΟ ΠΑΝΕΠΙΣΤΗΜΙΟ ΑΘΗΝΩΝ ΤΜΗΜΑ ΓΕΩΠΟΝΙΚΗΣ ΒΙΟΤΕΧΝΟΛΟΓΙΑΣ ΕΡΓΑΣΤΗΡΙΟ ΓΕΝΙΚΗΣ ΚΑΙ ΓΕΩΡΓΙΚΗΣ ΜΙΚΡΟΒΙΟΛΟΓΙΑΣ ΕΚΤΟΜΥΚΟΡΡΙΖΙΚΟΙ ΜΥΚΗΤΕΣ ΤΟΥ ΓΕΝΟΥΣ LACTARIUS PERS. ΣΤΗΝ ΕΛΛΑΔΑ ΜΑΡΙΝΑ Γ. ΤΡΙΑΝΤΑΦΥΛΛΟΥ ΕΡΕΥΝΗΤΙΚΗ ΕΡΓΑΣΙΑ ΣΤΑ ΠΛΑΙΣΙΑ ΤΟΥ ΠΡΟΓΡΑΜΜΑΤΟΣ ΜΕΤΑΠΤΥΧΙΑΚΩΝ ΣΠΟΥΔΩΝ «ΒΙΟΤΕΧΝΟΛΟΓΙΑ ΚΑΙ ΕΦΑΡΜΟΓΕΣ ΣΤΗ ΓΕΩΠΟΝΙΑ», ΚΑΤΕΥΘΥΝΣΗ «ΜΟΡΙΑΚΗ ΟΙΚΟΛΟΓΙΑ ΚΑΙ ΑΝΙΧΝΕΥΣΗ ΓΕΝΕΤΙΚΑ ΤΡΟΠΟΠΟΙΗΜΕΝΩΝ ΟΡΓΑΝΙΣΜΩΝ» ΕΠΙΒΛΕΠΩΝ: ΖΕΡΒΑΚΗΣ ΓΕΩΡΓΙΟΣ, ΕΠΙΚΟΥΡΟΣ ΚΑΘΗΓΗΤΗΣ ΓΠΑ ΑΘΗΝΑ 2012 1 2 ΓΕΩΠΟΝΙΚΟ ΠΑΝΕΠΙΣΤΗΜΙΟ ΑΘΗΝΩΝ ΤΜΗΜΑ ΓΕΩΠΟΝΙΚΗΣ ΒΙΟΤΕΧΝΟΛΟΓΙΑΣ ΕΡΓΑΣΤΗΡΙΟ ΓΕΝΙΚΗΣ ΚΑΙ ΓΕΩΡΓΙΚΗΣ ΜΙΚΡΟΒΙΟΛΟΓΙΑΣ ΕΚΤΟΜΥΚΟΡΡΙΖΙΚΟΙ ΜΥΚΗΤΕΣ ΤΟΥ ΓΕΝΟΥΣ LACTARIUS PERS. ΣΤΗΝ ΕΛΛΑΔΑ ΜΑΡΙΝΑ Γ. ΤΡΙΑΝΤΑΦΥΛΛΟΥ ΕΡΕΥΝΗΤΙΚΗ ΕΡΓΑΣΙΑ ΣΤΑ ΠΛΑΙΣΙΑ ΤΟΥ ΠΡΟΓΡΑΜΜΑΤΟΣ ΜΕΤΑΠΤΥΧΙΑΚΩΝ ΣΠΟΥΔΩΝ «ΒΙΟΤΕΧΝΟΛΟΓΙΑ ΚΑΙ ΕΦΑΡΜΟΓΕΣ ΣΤΗ ΓΕΩΠΟΝΙΑ», ΚΑΤΕΥΘΥΝΣΗ «ΜΟΡΙΑΚΗ ΟΙΚΟΛΟΓΙΑ ΚΑΙ ΑΝΙΧΝΕΥΣΗ ΓΕΝΕΤΙΚΑ ΤΡΟΠΟΠΟΙΗΜΕΝΩΝ ΟΡΓΑΝΙΣΜΩΝ» ΕΠΙΒΛΕΠΩΝ: ΖΕΡΒΑΚΗΣ ΓΕΩΡΓΙΟΣ, ΕΠΙΚΟΥΡΟΣ ΚΑΘΗΓΗΤΗΣ ΓΠΑ ΑΘΗΝΑ 2012 3 4 ΕΚΤΟΜΥΚΟΡΡΙΖΙΚΟΙ ΜΥΚΗΤΕΣ ΤΟΥ ΓΕΝΟΥΣ LACTARIUS PERS. ΣΤΗΝ ΕΛΛΑΔΑ ΤΡΙΑΝΤΑΦΥΛΛΟΥ ΜΑΡΙΝΑ ΕΡΕΥΝΗΤΙΚΗ ΕΡΓΑΣΙΑ ΣΤΑ ΠΛΑΙΣΙΑ ΤΟΥ ΠΡΟΓΡΑΜΜΑΤΟΣ ΜΕΤΑΠΤΥΧΙΑΚΩΝ ΣΠΟΥΔΩΝ «ΒΙΟΤΕΧΝΟΛΟΓΙΑ ΚΑΙ ΕΦΑΡΜΟΓΕΣ ΣΤΗ ΓΕΩΠΟΝΙΑ», ΚΑΤΕΥΘΥΝΣΗ «ΜΟΡΙΑΚΗ ΟΙΚΟΛΟΓΙΑ ΚΑΙ ΑΝΙΧΝΕΥΣΗ ΓΕΝΕΤΙΚΑ ΤΡΟΠΟΠΟΙΗΜΕΝΩΝ ΟΡΓΑΝΙΣΜΩΝ» ΑΘΗΝΑ 2012 ΤΡΙΜΕΛΗΣ ΕΞΕΤΑΣΤΙΚΗ ΕΠΙΤΡΟΠΗ: Ζερβάκης Γιώργος (επιβλέπων) Ταμπακάκη Αναστασία (μέλος) Γεωργακόπουλος Δημήτρης (μέλος) 5 6 ΠΡΟΛΟΓΟΣ Με την ολοκλήρωση της μεταπτυχιακής ερευνητικής μου εργασίας, νιώθω την ανάγκη να ευχαριστήσω κάποιους ανθρώπους που συνέβαλαν, ο καθένας με τον τρόπο του, στην πραγματοποίηση της. Θα ήθελα αρχικά να ευχαριστήσω ιδιαίτερα τον Επίκουρο καθηγητή κ. Γιώργο Ζερβάκη που με επέλεξε για την εκπόνηση της μεταπτυχιακής μου εργασίας και που μου έδωσε τη δυνατότητα να ασχοληθώ και σε μεταπτυχιακό επίπεδο με το ερευνητικό αντικείμενο που ιδιαίτερα αγαπώ, αυτό της συστηματικής των μακρομυκήτων. Τον ευχαριστώ για την επίβλεψη και καθοδήγηση του σε όλα τα στάδια αυτής της μελέτης καθώς και για τις πολύτιμες παρατηρήσεις και διορθώσεις του κατά τη συγγραφή του κειμένου. Ιδιαίτερα τον ευχαριστώ για την καθοδήγηση του στο κομμάτι της μοριακής ανάλυσης, που αποτελούσε σε πρακτικό επίπεδο, ένα πεδίο νέο για μένα. Ένα μεγάλο ευχαριστώ στη Λέκτορα κ. Ζαχαρούλα Γκόνου- Ζάγκου, η οποία ως η πρώτη μου δασκάλα στο αντικείμενο της συστηματικής των μακρομυκήτων, μου μετέδωσε τον ενθουσιασμό της και έχει θέσει τις βάσεις για την μετέπειτα πορεία μου. Ιδιαίτερα θα ήθελα να την ευχαριστήσω για την ουσιαστική συμβολή της στην παρούσα μελέτη, αφού ένα κομμάτι της πραγματοποιήθηκε στο εργαστήριο Μυκητολογίας του Τμήματος Βιολογίας του ΕΚΠΑ, οπότε και είχα την ευκαιρία να τη συμβουλεύομαι συχνά σε ζητήματα ταξινομικής φύσης. Θα ήθελα επίσης να ευχαριστήσω θερμά τον υποψήφιο διδάκτορα Παναγιώτη Δεληβοριά. Η πολύπλευρη βοήθεια του είναι πολύτιμη και οι παρατηρήσεις του πάνω σε ταξινομικά ζητήματα έχουν πάντοτε για μένα ιδιαίτερη βαρύτητα. Σημαντική είναι επίσης η συνεισφορά του στην παρούσα μελέτη, με πολυάριθμες φωτογραφίες ειδών του γένους Lactarius στο πεδίο, με την μετάδοση της εμπειρίας του πάνω στην κατασκευή των plates που βρίσκονται στην εργασία, καθώς και με τις συζητήσεις που είχαμε πάνω σε επιμέρους ταξινομικούς προβληματισμούς που προέκυπταν. Ιδιαίτερα χαίρομαι για την ευχάριστη και εποικοδομητική συνεργασία που είχα με τον διδάκτορα Ηλία Πολέμη, τον οποίο και ευχαριστώ για τις συζητήσεις που είχαμε πάνω σε ταξινομικά θέματα, καθώς και για τις όμορφες φωτογραφίες πεδίου που μου επέτρεψε να συμπεριλάβω στη παρούσα μελέτη. Ένα ευχαριστώ οφείλω επίσης στον κ. Δημήτρη Δήμου για τις φωτογραφίες πεδίου ειδών του γένους Lactarius, και την άμεση ανταπόκριση του, όποτε χρειάστηκα στοιχεία για δείγματα που ο ίδιος είχε συλλέξει και ταυτοποιήσει. 7 Μέσα από την ερευνητική πορεία των παραπάνω ανθρώπων προέκυψε το μεγαλύτερο μέρος του υλικού που μελέτησα και για αυτό η συμβολή τους στην παρούσα μελέτη είναι κάτι παραπάνω από σημαντική. Μέρος, επίσης, του ελληνικού υλικού που εξετάσθηκε έχει συλλεχθεί και μελετηθεί από τον κ. Στέφανο Διαμαντή και τον κ. Ζαχαρία Αθανασίου, τους οποίους και ευχαριστώ θερμά για την παραχώρηση των αντίστοιχων δειγμάτων. Θερμά ευχαριστώ επίσης τους ερευνητές των μυκητολογικών ερμπαρίων του εξωτερικού Annemieke Verbeken (GENT), Georgio Lalli (AQUI) και Giuseppe Venturella (PAL), για την παραχώρηση για τις ανάγκες της παρούσας μελέτης αποξηραμένων δειγμάτων που χρησιμοποιήθηκαν ως υλικό αναφοράς. Σε αυτό το σημείο νιώθω επίσης την ανάγκη να εκφράσω ένα θερμό ευχαριστώ στον υποψήφιο διδάκτορα Χρήστο Φωτιάδη, με την πολύτιμη βοήθεια του οποίου το κομμάτι της μοριακής ανάλυσης των δειγμάτων, που ήταν ένα πεδίο νέο για μένα, ολοκληρώθηκε με επιτυχία. Θα ήθελα να ευχαριστήσω επίσης την Επίκουρη Καθηγήτρια κ. Ευαγγελία Καψανάκη-Γκότση, για την υποστήριξή της και για τη δυνατότητα που μου παρείχε για την επιπρόσθετη μελέτη των δειγμάτων με το μικροσκόπιο τύπου Nomarski, του εργαστηρίου Μυκητολογίας του Τμήματος Βιολογίας του ΕΚΠΑ, καθώς και την διδάκτορα Πυρρή Ιωάννα και την υποψήφια διδάκτορα Σεργεντάνη Αθανασία για την ευχάριστη συνεργασία τους και για την τροποποίηση που έκαναν κάποιες φορές στο πρόγραμμα τους ώστε να διευκολυνθεί η μελέτη μου. Η παρατήρηση επιλεγμένων δειγμάτων σε ηλεκτρονικό μικροσκόπιο σάρωσης δεν θα ήταν εφικτή χωρίς την συνεργασία με τον Καθηγητή κ. Φασσέα, και τη βοήθεια του συνεργάτη του κ. Ψαροκωστόπουλου, τους οποίους και ευχαριστώ. Ευχαριστώ επίσης την Επίκουρη Καθηγήτρια κ. Ταμπακάκη Αναστασία και τον Επίκουρο Καθηγητή Γεωργακόπουλο Δημήτρη για τη συμμετοχή τους στην τριμελή εξεταστική επιτροπή. Θερμά επίσης ευχαριστώ όλους τους συνεργάτες του εργαστηρίου, και ιδιαίτερα τον Γιώργο Κουτρώτσιο για το φιλικό και ευχάριστο κλίμα που πάντοτε δημιουργούσε, κάτι ιδιαίτερα σημαντικό σε έναν χώρο έρευνας και εργασίας. Τέλος, ένα μεγάλο ευχαριστώ θα ήθελα να εκφράσω στους κοντινούς μου ανθρώπους για την στήριξη και αγάπη τους. Ιδιαίτερα ευχαριστώ τον Γιώργο Φουντουλάκη για την υποστήριξη του όλα αυτά τα χρόνια, για τη συντροφιά του και την πολύτιμη βοήθεια του σε όλες τις… μανιταροεξορμήσεις! 8 ΠΕΡΙΕΧΟΜΕΝΑ ΠΕΡΙΛΗΨΗ ................................................................................................................. 11 ABSTRACT ................................................................................................................. 13 Ι. ΕΙΣΑΓΩΓΗ ............................................................................................................... 15 1. Το βασίλειο των Μυκήτων (kingdom Fungi) ...................................................... 15 2. Το φύλο των Βασιδιομυκήτων ............................................................................. 17 2.1. Γενικά - Βιολογικός κύκλος και βασικά χαρακτηριστικά ............................. 17 2.2. Οικολογία ...................................................................................................... 21 2.2.1. Μυκόρριζες ................................................................................................. 21 2.2.2. Εκτομυκόρριζες (Ectomycorrhizal Mycorrhizae - EcM) ........................... 23 2.3. Φυλογενετικές σχέσεις και ταξινομική κατάταξη ......................................... 27 3. Το γένος Lactarius Persoon ................................................................................. 29 3.1. Φυλογενετική θέση του γένους ..................................................................... 29 3.2. Η τάξη Russulales .......................................................................................... 29 3.3. Η Οικογένεια Russulaceae ............................................................................ 32 3.4. Σύστημα ταξινόμησης εντός του γένους Lactarius Pers. .............................. 34 3.5. Εξελικτικές σχέσεις μεταξύ των γενών Russula και Lactarius ..................... 36 3.6. Ποικιλότητα- εξάπλωση του γένους Lactarius ............................................. 38 3.7. Η ποικιλότητα του γένους Lactarius στην Ελλάδα ....................................... 40 4. Οικονομική σημασία - Χρήσεις και εφαρμογές ειδών του γένους Lactarius ...... 41 4.1. Εδώδιμα είδη ................................................................................................. 41 4.2. Τεχνητή μυκόρριζα ........................................................................................ 42 5. Συστηματική Μυκήτων - Βασιδιομυκήτων ......................................................... 42 5.1. Ο κλάδος της Συστηματικής .......................................................................... 42 5.2. Κριτήρια-θεωρήσεις για το είδος στους Μύκητες ......................................... 43 5.3. Συστηματική Βασιδιομυκήτων ...................................................................... 44 9 5.4. Μέθοδοι κλασσικής ταξινόμησης Βασιδιομυκήτων ..................................... 45 5.5. Μοριακή συστηματική Βασιδιομυκήτων ...................................................... 48 6. Σκοπός της μελέτης .............................................................................................. 55 ΙΙ. ΥΛΙΚΑ ΚΑΙ ΜΕΘΟΔΟΙ ........................................................................................ 57 1. Βιολογικό υλικό ................................................................................................... 57 1.1. Αποξηραμένα δείγματα βασιδιοκαρπίων Lactarius ...................................... 57 1.2. Συλλογή νέων νωπών δειγμάτων Lactarius .................................................. 57 2. Καταγραφή μακροσκοπικών χαρακτηριστικών ................................................... 65 3. Μικροσκοπικές παρατηρήσεις ............................................................................
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