Melampsora Medusae

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Melampsora Medusae Europejska i Śródziemnomorska Organizacja Ochrony Ro ślin PM 7/93(1) Organisation Européenne et Méditerranéenne pour la Protection des Plantes Diagnostyka Diagnostic Melampsora medusae Zakres stosowania Niniejszy standard opisuje protokół diagnostyczny dla Melampsora medusae .¹ Zatwierdzenia i nowelizacje Zatwierdzony we wrze śniu, 2009 roku. Wprowadzenie Grzyb Melampsora medusae Thümen jest jednym ze sprawców rdzy topoli. Głównymi gatunkami w Europie powoduj ącymi rdz ę w uprawach topoli s ą Melampsora larici-populina i Melampsora allii-populina. Wszystkie trzy gatunki obficie wytwarzaj ą na li ściach urediniospory, co mo że prowadzi ć do przedwczesnych defoliacji i ograniczenia wzrostu. Po kilku latach silnych infekcji i powtarzaj ących si ę defoliacji, patogen mo że sta ć si ę przyczyn ą wi ększej podatno ści drzew na zamieranie, a nawet powodowa ć śmier ć drzew młodszych. Rdza powodowana przez grzyby z rodzaju Melampsora jest bardzo powszechn ą chorob ą topoli, której skutkiem mog ą by ć powa żne straty ekonomiczne w uprawach komercyjnych, z powodu powstawania i rozprzestrzeniania si ę nowych patotypów M. larici-populin a, M. medusae pochodzi z Ameryki Północnej, sk ąd rozprzestrzenił si ę na inne kontynenty. W XX wieku poza USA, Kanad ą i Meksykiem patogen rozprzestrzenił si ę w Europie (patrz poni żej), Ameryce Południowej (Boliwia, Brazylia, Chile), Południowej Afryce (Republice Południowej Afryki, Zimbabwe), Azji (Japonia), Oceanii (Australia, Nowa Zelandia). W regionie EPPO pojawienie si ę M. medusae odnotowano na ograniczonym obszarze w Belgii, Francji i Portugalii (EPPO, 1997). Gospodarzami pierwszorz ędowymi, na których wytwarzane jest stadium telium grzyba są drzewa nale żą ce do rodzaju Populus oraz ich miesza ńce. Gospodarzami drugorz ędowymi, na których grzyb wytwarza stadium ecjum są drzewa iglaste z rodzaju Larix (modrzew), Pseudotsuga (daglezja) i Pinus (sosna). Jednak że, w regionie EPPO grzyb nigdy nie został stwierdzony na żadnym z gospodarzy drugorz ędowych (Pinon, 1986; EPPO/CABI, 1997). ¹Użyte w standardach EPPO nazwy odczynników lub wyposażenia nie znacz ą wykluczenia innych, które równie ż mog ą by ć stosowane. © 2009 OEPP/EPPO, Biuletyn OEPP/EPPO 39, 328–336 Dotychczas w Europie opisano osiem gatunków z rodzaju Melampsora pora żaj ącycyh topole (Pinon, 1973; Cellerino, 1999). Jakkolwiek tylko trzy z nich: M. larici-populina , M. allii-populina, M. medusae są patogeniczne dla topoli nale żą cych do sekcji Aigeiros i Tacamahaca , tj. Populus nigra , P. deltoides , P. trichocarpa ich mi ędzygatunkowych miesza ńców (Frey i wsp., 2005). W Europie wi ększo ść uprawianych topoli stanowi ą miesza ńce P. x euramericana (P. deltoides x P. nigra ) i P. x interamericana (P. trichocarpa x P. deltoides ). Ponadto, Shain (1988) udowodnił istnienie dwóch form specjalnych w obr ębie gatunku M. medusae i nazwał je M. medusae f. sp. deltoidae i M. medusae f. sp . tremuloidae, ze wzgl ędu na ich pierwotnego gospodarza, odpowiednio P. deltoidae i P. tremuloidae. Ani dyrektywa Unii Europejskiej ani EPPO nie rozró żnia tych dwóch form specjalnych, tylko odnosi si ę do M. medusae. Niemniej jednak, od czasu gdy M. medusae został odnotowany w Europie tylko na topolach nale żą cych do sekcji Aigeiros i Tacamahaca i maj ąc na uwadze specjalizacj ę zwi ązan ą z ro ślin ą żywicielsk ą, mo żna stwierdzi ć, że do tej pory odnotowano w Europie tylko obecno ść M. medusae f. sp. deltoidae (Pinon, 1991; Pinon & Frey, 2005). Pi ęć innych gatunków, M. larici-tremulae , M. magnusiana , M. pinitorqua , M. rostrupii i M. pulcherrima , są tylko patogeniczne w stosunku do gatunków z sekcji Populus dawniej Leuce ) tak że nazywanych „osik ą”: P. alba , P. tremula oraz ich miesza ńców (Frey i wsp., 2005). W literaturze naukowej cztery pierwsze taksony s ą czasem poł ączone w jeden kompleks M. populnea (Pers.) P. Karst, ale aktualnie w oparciu o filogenez ę wielogenow ą, kompleks ten mo że by ć wyra źnie podzielony na ró żne grupy (Feau i wsp., 2009). Od czasu jak „osikowe” gatunki nie s ą powszechnie uprawiane w celach komercyjnych i rosn ą w Europie głównie jako dzikie lub ozdobne drzewa, obecno ść tych pi ęciu gatunków Melampsora na P. alba , P. tremula i ich miesza ńcach ma znacznie mniejsze znaczenie ekonomiczne (Frey i wsp., 2005) ni ż mogłoby mie ć wprowadzenie do Europy M. medusae f. sp . tremuloidae. Ponadto, warto odnotowa ć, że M. medusae f. sp . deltoidae tworzy miesza ńce mi ędzygatunkowe z innymi gatunkami z rodzaju Melampsora. Pierwszy M. medusae – populina został odkryty w 1991 w Nowej Zelandii (Spiers & Hopcroft, 1994) mi ędzygatunkowy mieszaniec M. medusae i M. larici-populina a nast ępnie znaleziony w Afryce Południowej (Frey i wsp., 2005). Drugi, M. x columbiana , mi ędzygatunkowy mieszaniec M. medusae i M. occidentalis, został odkryty w 1995 na północno – zachodnim Pacyfiku Stanów Zjednoczonych (Newcombe i wsp., 2000). Chocia ż żaden z tych miesza ńców nie został do tej pory odnotowany w regionie EPPO, szczególn ą uwag ę nale ży zwróci ć w rejonach, gdzie wyst ępuje M. medusae i M. larici-populina. To żsamo ść Nazwa: Melampsora medusae Thümen Nazwy makrocyklicznych stadiów rozwojowych: Caeoma faulliana Hunter (stadium ecjo), Uredo medusae Thümen (stadium uredio). Synonim: Melampsora albertensis J.C. Arthur Stanowisko taksonomiczne: Fungi: Basidiomycota: Uredinales Nazwa zwyczajowa: Rdza topoli Informacje dotycz ące taksonomii i nazewnictwa: patrz sekcja wprowadzenie Komputerowy kod EPPO: MELMME Kategoria fitosanitarna: EPPO A2/74 UE I/AI © 2009 OEPP/EPPO, Biuletyn OEPP/EPPO 39, 328–336 Wykrywanie Objawy Objawy chorobowe na ro ślinie żywicielskiej (stadium telium) wywołane przez grzyby, sprawców rdzy topoli, pochodz ących z Europy ( M. allii-populina, M. larici-populina ) s ą trudne do odró żnienia od objawów spowodowanych przez M. medusae i nie s ą wystarczaj ące dla postawienia prawidłowej diagnozy. Objawy chorobowe na ro ślinie żywicielskiej, na której wytwarzane jest stadium telium (topola). Wczesnym objawem infekcji wywołanej przez grzyby z rodzaju Melampsora, sprawców rdzy, bez wzgl ędu na gatunek, s ą żółtawe plamy (uredinia wypełnione mas ą urediniospor, fot. nr 1) na spodniej stronie li ścia wzdłu ż nerwu (fot. nr 2). Uredinia mog ą by ć rozproszone na powierzchni liści lub w skupiskach, tak że cała powierzchnia wygl ąda jak posypana żółto- pomara ńczowym pudrem (fot. nr 3). W sprzyjaj ących warunkach klimatycznych, podczas całego sezonu wegetacyjnego tworz ą si ę liczne bezpłciowe stadia z urediniami i urediniosporami. Po pierwotnym zaka żeniu, w warunkach du żej wilgotno ści urediniospory są łatwo przenoszone przez wiatr w obr ębie całej korony drzewa i w pobli że podatnych drzew. Przy silnej infekcji uredinia mog ą obejmowa ć cał ą powierzchni ę li ścia, nadaj ąc mu złoty wygl ąd i powodowa ć cz ęś ciow ą lub całkowit ą defoliacj ę. Pó źnym latem do połowy jesieni po obu stronach li ści pojawiaj ą si ę telia (skupienia zarodników), małe, br ązowe do czarnych wygl ądem przypominaj ące krosty. Telia zawieraj ą ogromn ą liczb ę grubo ściennych zarodników spoczynkowych (teliospor), które zimuj ą na opadłych li ściach i kiełkuj ą nast ępnej wiosny w podstawk ę, na której tworz ą si ę bazidiospory. Poło żenie telium jest zró żnicowane i zale żne od gatunku. M. medusae i M. allii-populina tworzy telia głównie pod epiderm ą, podczas gdy u M. larici-populina obserwuje si ę je mi ędzy kutikul ą a komórkami epidermy. Fot. nr 1 Uredinia grzyba z rodzaju Melampsora na spodniej powierzchni li ścia topoli (dzi ęki uprzejmo ści P. Frey, INRA Nancy, Fr). © 2009 OEPP/EPPO, Biuletyn OEPP/EPPO 39, 328–336 Fot. nr 2 Typowe objawy rdzy na li ściu topoli (uredinia) spowodowane przez grzyb z rodzaju Melampsora (dzi ęki uprzejmo ści P. Frey, INRA Nancy, Fr). Fot. nr 3 Młoda topola silnie zainfekowana przez grzyb z rodzaju Melampsora - sprawc ę rdzy (dzi ęki uprzejmo ści P. Frey, INRA Nancy, Fr). Poniewa ż M. medusae jest gatunkiem heteroecjalnym, jak M. larici-populina i M. allii- populina, potrzebuje gospodarza drugorz ędowego, na którym wytwarza stadium ecjum do zimowania w stadium seksualnym. Nie ma dowodów, że w klimacie umiarkowanym M. medusae prze żywa w stadium uredinium. Jednak że, uwa ża si ę, że M. medusae zimuje w © 2009 OEPP/EPPO, Biuletyn OEPP/EPPO 39, 328–336 Australii w stadium bezpłciowym dzi ęki tzw. „zielonemu mostowi ” który tworz ą zimozielone topole (Wilkinson & Spiers, 1976). W krajach EPPO grzyb M. medusae jest odnotowany tylko na topoli i do tej pory nie był nigdy obserwowany na potencjalnych gospodarzach drugorz ędowych. Objawy chorobowe na ro ślinie żywicielskiej, na której wytwarzane jest stadium ecjum (drzewa z rodzaju Larix, Pseudotsuga, Pinus ). Bazidiospory infekuj ą wiosn ą gospodarzy drugorz ędowych zaka żaj ąc igły i produkuj ąc jasno żółte plamy (spermogonia) na górnej powierzchni. W ci ągu kilku dni, po plazmogamii pojawi ą si ę na dolnej stronie li ści jasno żółte uwypuklenia (ecja). Struktury te najlepiej rozwijaj ą si ę na tegorocznych igłach, ale mo żna je obserwowa ć, chocia ż rzadko na szyszkach drzew iglastych lub na młodych p ędach. Ecja produkuj ą du żą liczb ę żółtych ecjospor, uło żonych w ła ńcuchy. Wiosn ą przenoszone przez wiatr ecjospory infekuj ą podatne ro śliny z rodzaju Populus, nast ępnie grzyb tworzy uredinospory i w ten sposób zamka swój cykl życiowy. Procedura pobierania próbek Poniewa ż stadium ecjum grzyba M. medusae nigdy nie zostało odnotowane na drzewach iglastych w regionie EPPO, procedura pobierania próbek ogranicza si ę do drzew topoli, gdzie prawdopodobie ństwo wykrycia grzyba b ędzie najwy ższe. Poniewa ż, wymiary, kształt i inne morfologiczne cechy teliospor M. medusae nie wykluczaj ą innych rodzimych gatunków Melampsora na topoli, powinny by ć zbierane tylko li ście topoli, na których znajduj ą si ę urediniospory. Inspekcje najlepiej przeprowadzi ć wczesnym latem, kiedy symptomy choroby są najbardziej widoczne i ilo ść urediniospor na najwy ższym poziomie. W celu analizy, zainfekowane li ście mog ą by ć bezpo średnio przesyłane do laboratorium.
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