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Planthopper Transmission of Phormium Yellow Leaf Phytoplasma

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Planthopper Transmission of Phormium Yellow Leaf Phytoplasma A ustralasian Plant Pathology (1997) 26: 148-154 Planthopper transmission of Phormium yellow leaf phytoplasma L.W. Llefting'", R.E. Beever",C.J. Winksc, M.N.Pearson" and R.L.S.Forster­ AHortResearch, Private Bag 92169, Auckland, New Zealand BSchool ofBiological Sciences, The University of Auckland, Private Bag 92019, Auckland, New Zealand CManaaki Whenua - Landcare Research, Private Bag 92170, Auckland, New Zealand Abstract Phormium yellow leaf(PYL) phytoplasma was transmitted from diseased to healthy New Zealand flax (Phormium tenax) by the native planthopper, Oliarus atkinsoni (Homoptera: Cixiidae). By contrast, transmission was not effected by the introduced passionvine hopper, Scolypopa australis (Homoptera: Ricaniidae). Successful transmission of PYL phytoplasma from New Zealand flax to New Zealand flax by 0. atkinsoni was demonstrated by symptomatology and by polymerase chain reaction (PCR) of the test plants using phytoplasma-specific primers to the 16S rRNA genes. When the salivary glands and the remaining body of the planthoppers used in the transmission studies were tested separately by PCR for the presence of phytoplasma, PYL phytoplasma was detected in 100% of both the salivary glands and the bodies of pre-transmission 0. atkinsoni, and in 44% and 67% of the salivary glands and the bodies of post-transmission planthoppers, respectively. The phytoplasma was not detected by PCR in the whole bodies ofhoppers ofS. australis. Additional keywords: Cordyline australis, vector Introduction (Cumber 1953). Further, Cumber(1953) foundthat 16ofthe 20NewZealandflax seedlings cagedwith Phormium yellow leaf(pYL) isa lethal disease ofthe adult O. atkinsoni developed typical yellow-leaf large tufted lilioid monocotyledons, NewZealand symptoms, whilenoneof20 controlplants showed flax(Phormium tenax J.R etG.Forst.) andmountain any symptoms. A number of further transmission flax (P cookianum LeJolis). PYLwasa contributor experiments, including an experiment using to the eventualdemiseofa locallyimportantnatu­ planthoppers collected froman area free ofyellow­ ral fibre industry based on P tenax (Boyce and leaf, allimplicated 0. atkinsoni asthevectorofPYL Newhook 1953).The diseasewasfirst recognised (Boyce etal. 1953). A searchfor alternativeinsect over 80 years ago, but it was not until 1969when vectors ofPYLwascarriedoutbyCumber(1954b). Ushiyama et al. (1969)observed phytoplasmas in The mealybug Trionymus diminutus (Leonardi), the phloemof diseasedplants and in the proposed leafhopperErythroneura zealandica Myers, and planthopper vector (Oliarus atkinsoni Myers planthopper Oliarus oppositus(Walker) allfailedto (Homoptera: Cixiidae)) byelectron microscopy, that transmit the disease. However, most of these later the etiologyof the diseasewas clarified. More re­ transmission experiments werecomplicatedbytest cently, phylogenetic analysis of the 168 rRNA plants dying from 'sudden die-back', a disease of genes of PYL phytoplasma determined that it unknownetiology (Cumber1954a and b). Further­ belongs to a subgroup of the aster yellows group more, Boyce etal. (1953)reportedmechanicaland (Lieftinget al. 1996), and is verycloselyrelatedto seed transmission of yellow-leaf: in view of our Australiangrapevine yellows phytoplasma, sharing currentknowledge ofphytoplasmas, theseobserva­ 99.5%ofthe 168rRNAgenesequence (padovanet tionsarealmost certainly inerror. Theselaterexperi­ al.1996). ments, carried out before the discovery of Theplanthopper, 0. atkinson i isendemic toNew phytoplasmas (Doi et al. 1967), raisedoubts astothe Zealand where it is confined to Phormium spp. validity ofthetransmission experiments in general. Largenumbers of0. atkinsoni werepresent onNew Thus,therewasa real needto clarifydiseasetrans­ Zealand flax in badly affected yellow-leaf areas missiontaking intoaccountthe proposedetiology. 148 Australasian Plant Pathology Vol. 26 (3) 1997 Since the late 1980s, a syndrome known as or S. australis wereplacedon individual test plants 'sudden decline' has caused widespread death of ininsect-proofcagesfora periodof21 days. SixNew cabbagetrees (Cordyline australis (Forst. f.) Endl.) Zealand flax seedlings and six cabbage trees were in New Zealand (Beever et al. 1996). This tufted, usedwitheach insectspecies. Asa control, sixNew woody, lilioid monocotyledontree is also native to Zealand flax plants and six cabbage trees were New Zealand and frequently grows in association placed in mesh bags (three plants per bag) withno with Phormium tenax. It has been proposed that insects. An additional 30 insects of each species sudden decline is caused by a phytoplasma, whichwerenotused inthetransmission experiments perhaps identical or very closely related to PYL werestoredat -20°C and are referred to as the pre­ phytoplasma, although this has yet to be proven transmission insects. (Beeveretal.1996). At the end of each experiment, the insects that The passionvine hopper, Scolypopa australis remained alive were removed from the plants and Walker (Homoptera:Ricaniidae), introducedfrom stored at -20°C and are referred to as the post­ Australia, isfoundinlargenumbers ona widevariety transmission insects. Theplantsweremaintainedin of native and exotic plants, including Phormium an insect-proofshadehouseand regularlyinspected spp. (Cumber 1966). The distributionofS. australis for symptoms. within NewZealandcloselyparallelsthat ofsudden decline, leading to the suggestionthat it couldbe a Plant DNA extraction A modification of the vector ofthis disease (Beeveret al. 1996). cetyltrimethylanunonium bromide(CTAB) method The main aim of this research was to carry wasused(Ahrensand Seemiiller 1992). FrozenNew out transmission experiments to determine if Zealand flax leaf bases (2 g) were ground in liquid O. atkinsoni is a vector ofPYL phytoplasma. Also, nitrogen with a mortar and pestle. The powder was S. australis was examined as a possibleadditional transferred toa 30mL tubeand mixedwith 10mL of 4' vector ofPYL phytoplasma.Further,attemptswere isolationmedium(100mM Na2HPO 10%sucrose, madetotransmit PYLphytoplasmato cabbagetrees 2%polyvinylpyrrolidone [pVP-40], pH7.6towhich by 0. atkinsoni and S. australis to determineifPYL 0.15%bovineserumalbuminand ascorbicacid to a phytoplasma is involved in the etiologyof sudden final concentrationof 1mM were addedjust before decline. use). The homogenate was centrifuged at 4°C for 5 minat 1500g. The supernatant wasdecantedand recentrifuged at 4°Cfor25 minat 18000g. Thepellet Methods wasresuspendedin 10mL hot (60°C) CTABbuffer (5%CTAB, 100mMTris-HCl [pH8.0], 20mMEDTA, Plant material HealthyP tenax wasgrownfrom 1.4M NaCl, 1%PVP-40) and wasincubated at 60°C seed collected from a vegetatively propagated for 30 min. The lysatewas extracted with an equal descendant of selection 56 shown by Boyce et al. volume ofchlorofonnlisoamyl alcohol (24:1,v/v). (1953) to be susceptible to PYL phytoplasma. Aftercentrifugation, the aqueouslayerwasprecipi­ Healthy seedlings of C. australis weregrownfrom tated with a two-thirdvolume ofice-cold isopropa­ seedcollectedfrom a cuttingfrom a treethat subse­ nol,incubated at -20°C for 111, and then centrifuged quentlydied from suddendecline. Seedlings ofboth at 10 000g for 15min. The pelletwas washedwith species were 1year-old at the time of caging with ice-cold 70% ethanol, dried under vacuum, and insects. resuspendedin 50 J1L TE buffer (10 mM Tris-HCl, 0.1 mMEDTA,pH8.0). Insect samples Adults of O. atkinsoni and S. australis which were feeding on PYL-affected Preparation of insects for PCR New Zealand flax were collected during February 1995from a sitenearKawakawa(Northland) where Dlv'A extraction from insect bodies DNA was an area of about 1 ha of affected plants occurred extractedfrominsectbodiesbythe methodof Good­ naturally. Most of the plants showed yellow-leaf win et al. (1994). Individualinsectsweremacerated symptoms and many were dying. in 125J1Lextractionbuffer(2%CTAB, 1.4MNaCl, 1%PVP-40,0.02MEDTA,0.lMTris-HCLpH8.0)in Insect transmission experiments The day fol­ a microfuge tubeusinga micropestle, madeby melt­ lowing collection, groups often adult 0. atkinsoni ing the tip of a large sizedisposablepipette tip. The Australasian Plant Pathology Vol. 26 (3) 1997 149 remains ofthe insectswereremoved, the tubewas chloroplasts andmitochondria. TheprimersNGF(5'· brieflyvortexed andthenincubatedat65°Cfor5 min. AGG CGGCITGCTGGGTCTT-3')andNGR(5'­ The suspensionwas extractedonce with an equal AGCCATTGTATCACGTT-3') weredesignedto volume ofchloroform:isoamyl alcohol(24:1,v/v), amplify a 500bpfragment ofthe 16SrRNAgeneof and the nucleic acids were precipitated byadding allphytoplasmas (M.T.Andersen, personalcommu­ l/lOvol. 3Msodiumacetate, pH5.2, and2.5 vol. ice­ nication)and wereusedto test for the presence of cold ethanol and incubating at -20°C for at least PYLphytoplasma in bothplant and insectsamples. 1h. Aftercentrifuging for15min at 13000g, thepel­ PCRamplification was performed witha 20 ul, re­ let was washed with 70% ice-coldethanol, dried action mixture containing 4 ul, (10to50ng)oftem­ undervacuum, and resuspended in 15ul, TEbuffer. plateDNA,20 J.lM ofeachprimer,200 J.lM dNTPs, , 1.25mMMgC12 and I U ofTaqDNApolymerase Dissection ofthe salivary glands The salivary withrecommended buffer (Amplitaq, Perkin-Elmer) gland systemof the planthopperslies in the head or 1.75U of Expandl" High FidelityPCR System and in the fore-part ofthe thorax and isbilaterally withrecommended buffer containing 1.5mMMgC12 symmetrical. The insects werefirstbriefly dipped in (Boehringer Mannheim). Amplification
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