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    Phyto73n06 875

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    Peology and Epidemiology Comparison of Host Ranges of Peronoselerospora philippinensis and P. sacchari M, R. Bonde and G. L. Peterson Research plant pathologist and biological laboratory technician, respectively. U.S. Department of Agriculture, Agricultural Research Service, Plant Disease Research Laboratory, P.O. Box 1209, Frederick, MD 21701 Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the product by the U.S, Department of ‘Agriculture and does not imply approval to the exclusion of other products that also may be suitable, “The authors thank O. R, Exconde and S.C. Chang for the isolates of Peranosclerospora phillppinensis and P, sacchari, respectively. Accepted for publication & January 1983, AUSTRACT. Bonde, M. R.,and Peterson, L. 1983. Comparison of host ranges of Peronoselerospora philippinensic and. sacchar, Phytopathology 73:87S-878 Seventy-two plant species representing 22 genera within eight tribes of the Gramineae were tested for suscepibilty to systemic colonization alter conigial inoculation with wo solate of Peronosclerspora philppinenss {om the Philippines. All susceptible specce were members ofthe tide Andeopogoneae (genera—Andropagon [one species), Sothriochioa [10 ‘species. Enlafie (ane species], Saccharum fone species), Sehizarhyi [Ihrer species] and. Sorghum [thece species) or the tribe Maydese Philippine downy mildew of maize, caused by Peronosclerospora philiopinensis (Weston) C. G. Shaw, has not been reported in the ‘western hemisphere, but i isa serious problem in the Philippines ‘where disease losses some fields have been 40-40% (7). Although maize varieties resistant to P.philippinensis have been developed in Asia, maize hybrids currently grown in the United Statesare highly susceptible to P.philippinensis, and American breeding lines that are highly resistaat to P sorghi (cause of sorghum downy mildew of maize) inthe United States are highly susceptible to P. hilippinensis (M. R. Bonde, unpublished) ‘Weeds and cultivated erops play a major roe in the perpetuation ‘of the pathogen inthe absence of maize (6). Besides maize (13), P. Philipinensis has been reported to infect Avena sativa L. (6), Euchlaena mexicana Schead. (= Zea mays L. mexicana) (6), E ‘mexicana X Zea may's hybrids (6), Miscanthus japonicus Andress (14, Saccharum officinarum 1. (6). S. spontaneum 1. (14), ‘Tne pateaton css af ni ala mae tayaain pay page car payment Te {Mle must norte be raety mares aaron acorns wih WUSC E ‘ot oto ak ‘1085 The Arwrican Phytopathooaicl Society (geneta—Tripsacum fone species] end Zea [thre species. Not ll secession susceptible to one folie ofthe pathogen showed systemic Somptoms when imoculated with the second isolate in some instances, on) ‘ew plants oan accession developed systemisymploms. The rents were emuarkably simile to our previous host range study with P.scchart rom Taiwan, and the information presented here indicates avery close phylogenetic relationship between ?. hillppinenis and P. sacha. ‘Sorghum arundinacewn (Willd) Stapt (11), 8. bicolor Moench (13), halepense(L.) Pers. (6). and S. propingwum (Kunth) Hitch. 6. ‘As part ofa program to determine the threat of specific foreign downy mildew pathogens to Ameviean agriculture should they spread to the United States, we conducted a study to determine whether P. philippinensis could iniect additional alternative hosts, that might allow the pathogen to overwinter and actas reservoirs of inoculum 10 infeet maize. The information, besides having epidemiologic significance for maize production, also could help resolve the confusion over the taxonomy of Feronoselerospora spp. 2). For this latter purpose, we included plant species that, although not present in the United States, might be useful in differentiating species within the genus Peronosclerospora. The results obtained with P. philippinensis were remarkably similar to those we recently reported foranisolate of P. sacchari from Taiwan a. The purpose of this research was to compare the host ranges of the two pathogens by including recent data and data fram the previous study of the susceptibility of various plant species to P. sacchari. Asfaras weareaware this isthe fist report of host ranges Vol. 73, No.6, 1983875 ‘of P. phillppinensis and P. sacchari isolates from different countries being compared using the same techniques on the same plant accessions MATERIALS AND METHODS. ‘Two cultures of P. philippinensis were obtained from the Philippines—one in 1978 and the other in 1979-—and two cultures fof Pa sacchari were isolated in our laboratory in 1975 and 1977, respectively rom infected sugarcane sets sent from Taiwan, Seeds ‘of test species were planted in pasteurized soil in 10-em-diameter clay pots, Seedlings atthe two- to three-leaf stage were sprayed with A suspension of conidia collected by the technique of Sehimite and Freytag (8) from infected Zea mays Pioneer 3369° or ‘DeKalb XLA3." The inoculum dosage used was 5.0 X 10* conidia per milliliter of suspension, | mi per pot of seedlings. The number of plants per pot depended on plant species. In s few instances, 20-40 plants were present ina pot; however, usually there were less than To per pot. A highly susceptible maize hybrid, either Pioneer 3369 ‘or DeKalb X143, was included as a control in each experiment, Recently planted sugarcane sets (with buds about to break) in IS-em-diameter clay. pots were sprayed with a suspension of conidia at 5.0 x 10" conidia per milliliter, 1 ml per set. All inoculated plants were incubated overnight in a dew chamber at 21-22 C and then placed ina greenhouse for disease development. ‘The normal temperature fluctuation in the greenhouse was 21-28 Cz however, on a few aceasions during the sumer months, it peaked as high as 34 C shortly after noon and returned to the formal temperature range within 3hr. Plants were examined forat Teast 30 days, and usually for 6 wk, after inoculation and were ‘compared with uninoculated controls ‘To verily infection within a plant accession, a few plants displaying systemic symptoms were placed overnight in dew chambers, and thei leaf surfaces were examined for development of conidia and conidiophores typical of P. philippinensis or P. sacchari. (In instances in which only one or a few plants had systemic symptoms, all were placed in the dew chambers). In Addition, leaf pieces displaying systemic symptoms were placed in the whorls of highly susceptible maize seedlings in dew chambers And incubated overnight at an optimum temperature co induce sporulation and subsequent infection of maize. Susceptibility of a plant accession was based on the presence of sporulation andjor reinfection of maize, in addition to symptoms indicating systemic infection All plants lacking systemic symptoms also were placed overnight in dew chambers and examined for the absence of conidia formation to substantiate that they were not infected ‘A few infected plants of each susceptible plant accession were TABLE |, Level af susceptibility of grass accessions susceptible to at lest one isolate of Paronosclerosporaphilpplnenis, and thee susceptibility 10 Peronoslerospora sacchari Proportion of inoculated plants showing systemic symptom Plant specie umber Origin P philppinensis P.saccharh Tribe: Andropogonene “Andropogon gerard Vtm 315686 usa, 3/10 2/20 ‘oshriochioa ambigua ST. Blake 301264 Australia a : "2 barbinods (Lag) Herter 2i60s4 USA’ "7 : 2. decipiens (Hack) C. E. Hub. 301290 Austral 10/34 ° 1. edwardsiana (Gould) Parodi 337509 ‘Argentina 20/21 * 2B lschaemum (L) Keng vat ichaemum 302508 Hungary 35/9 a8 2B leguroides (DC) Piper 202289 Brant 205 oftto 2 perforata (Vin. ex Fourn) Herter 228517 USA, 36/9 1188 1 Sprngheldi (Gould) Parodi 301007 USA, 16/25 193s ‘B. woodrowil (Hook £) A. Camus 732 Iie user 9128 Bulelia fala (RBs) Kee ‘so07e Australi at 3/34 ‘Saccharum oficinarun L. °CP-46-101" usa! 10/10 145 ‘Schizachyrium hiriflorum News 228507 USA, m2 a3 ‘Snierostachyu (Desv. ex Hari) 56269 Argento 3 192 Roseng., Art 10g ',seopariue (Micha) Nash 213875 USA 19. aun ‘Sorghum bicolor (L.) Meese? (drummondif} so2idt Aseria 132. Siar 'S bicolor (gambicun) 302150, Che 2/102 9/30 '. bicolor [hewisont} 0273 Sudan ai Byte '§ bicolor Vapontum) jo2i7a Porugal 2484 a9 bicolor melaleweur] 208708 Algeria 25 2/26 'S bicolor utiforme] 257293, Argentina 249 14/99 'S bicolor (nigricans) 302177 Portugal 139 one '§ bicolor (rion) 196890 Ethionin us “ bicolor acharatum) 302198 Asgentina 639 12/03 'S bicolor [tudanente] usa’ 1/98. 1s) 5 bicolor [echnicur usa sir 16 5 halepense (1) Per USA 2/26 * SS. plumosuam (R. Br.) Reavy." 198999 Australia 46 ~* “Tribe: Maydeae Trpsacunn dot foes (L) L usa 16)38 20/23 Zea diploperennis Is, Deebley & Guzman Mexico gyi 2 “Z mays L sbep. mays 369A" and XLAS USA varia 102) 103 Z moss subsp, meniena (Schad) Is seao7t Mexico 71 99/59 2 perennis(Hiche ) Reeves & Mangels Mexico wis 38 “Re the «wo to theelea stage seedlings ofall azesions, except those of Saccharum officinarum, were sprayed with a conidial suspension of P hilippinenss or P sacchari at 30% 10" conidia permite, | ml per pot of seclings. Data were not obtained *Sugnronne sets (vith buds about to break) were sprayed witha suspension of conidia at 5.0% 10? conidia per milliliter, 1 ml per se “Thetaxanomicsystem described by 1. M. 1. DeWet (3) wa ed for Sorghum bicolor. Thisssstem combined 48 previously separate species with S bolor. “The previous specific epithet aiven in brackets. "Another accession of Sorghiom plumogum not tested agaist P phillppinensis was shown tobe susceptible (5/85) to P. sacchari 876 PHYTOPATHOLOGY saved for later testing for the presence of oospores. Leaf pieces displaying systemic symptoms were collected from upper, middle, and lower leaves of plants with systemic symptoms at or near maturity and were fied ina mixture of absolute ethanol and acetic acid (21, v/v), After at least 18 hr in the fixative, the pieces were cleared for 4 hrin lactophenol, stained 24~72 hr with Sudan 1 in lactophenal (time depending on the specific lot of commercial stain), mounted in lactophenol on glass slides, and observed rieroscopicaly. RESULTS AND DISCUSSION Results of tests for susceptibility f 72 plant species representing 22 genera within eight grass tribes to P philippinensis, in addition to the susceptibility of many of the same plant accessions to P. sacchari are summarized in Tables | and 2and the text below. The dota for each isolate per pathogen were bulked. In several instances, several accessions ofthe same plant species were tested (Tables I and 2). Besides Sorghum bicolor "Tx412" and S. bicolor accession 431392, [8 accessions ofS. bicolor previously considered to belong to separate Sorghum species, and one sorghum-Sudan srass hybrid were tested. A major problem with several grass accessions was the poor germinabilty of thesced, which resulted in ‘only limited numbers of plants, ‘The following plant species were not susceptible to P. philippinensis: Agrostis stoloniferaand Alopecurusaequalis ofthe tribe Agrostideae: Avena abyssinica, A. barbata, A. brevis, A. byzamina, A. fatua, A. longiglamis, A. muda, A. eriantha, A. sativa, A. sterlis, and A. sirigosa of the tribe Aveneae; Bleusine indica of the tribe Chiorideae; Bromus inermis, Festuca rubra ‘subsp. rubra, Poa compressa, and P. nemorails of the tribe Festuceae; Agropyron cristatum, A. repens, and Lolium ‘multiflorum of the tribe Hordeae; and Pantewm miliaceum, P. virgatum, Pennisetum americanum, an P.macrouram ofthe (ibe ‘TABLE2. Plans ofthe tribes Andropogoneacand Maydene not susceptible to Pernoslerosporaphlippinenis and theircorrespondingsuscepsibiies 0 Peronosclerospore saechart Proportion of inoculated plats showing systemie symptoms" Plant species PA number Oris P.philppinenss P.saccharh Tribe: Andropogancae “Andropogon dstachyus L 300696 Hungary a9 0/24 “A terns Miche, 301216 USA’ ovat ota Arthraxon Mspidus (Thun) Mak 225086, Japan 0723 0162 ‘Bothriochloa alia (Witehe) Hen. 337510, ‘Argentina 036 ons "Beaucasica Trin} CE, Hubb. 30712 USA 0738 2/16 Beoucasice 301737 Pakistan ors 4B enartiana (Domin) CE. Hubb, 301308 Australia ois 4 glabra (Roxb) A. Camus 00726 tia oa 4 grahami (Haines) Boe, 2a1498 New Guinea ors 1 hybrida (Gould) Gould 321637 usa 0126 B insculpra (Hochst) A Camus 500730 USA oat B incuba 501436. S. Arion oral B.inermedia (R.Br) A. Camus 00806 Usa or13s ‘intermedia 300857 USA, 0/58 ‘.lsehacraum (L.) Keng var ischacums 301507 aia os ‘B odorata (Lisboa) A. Camus 01682 lia oy6s B pertua(L) A. Camus 50163 ncn oa B porta swiss nia on _ B periwsa 0164s on * B radicans (Lehm.) A. Camvs 369353 Mozambique orias ob Capilipedim venusium (Taw) Bor. sors British Guana 0/26 Heteropogon contort (L.) Beas. die USA 0/28 ‘x Room & Seb Hseontaris amir Inia 0126 Schizachyrium craw (Hack) Woot & Stan 216107 Mesiso : ‘S-condensaums (H.B.K.) Nees 2oiass Brau oj Sorghum ® abmura Parodi 2oree0 0726 '§ Bcofor(L) Moenehs "TXA12" Usa 168 S bicolor 431392 S bicolor (afirorum) 302135 4108 S. bicolor (eautarum passe us Sbleotor fermaumn] 267126, on, S. bicolor [dura] 267108, sia 'S bicolor [euineense] 232857, 216 ‘bicolor [mein 261328 oat ‘bicolor [nervosum) 208710 ont '.alepense [eontroversum] 307268 org ‘.halepense Fniaeuon mss, : '.plumocum (R. Be.) Beau. 220931 Australia . 'S propimgum (Kunth) ithe, s02194 iines 14s 'S puatontfoltam Snowden 20 India 212 'S bersicolor Anders, 260273, Ethiopia 629 '.vertelifforam (Stevd.) Sept 213901 Rhodesia 0136, Tribe: Maydeae Cox lacrymajobl 262 Bras ot ons ‘Seedlings ofall accessions at tho o> to threelea tage were sprayed witha conidial suspension of Pphilippinensis or P. sacchari at 50% 1 conidia pet tlle, Il per pot of seedings, “Date were not obtained "The taxonomic system described by J. M. J. DeWet (S) was used for Sorghum bicolor. This system combined 48 previously recognized species with S. bicolor. The previous specific epithet i given in Brackets Vol.73,No. 6, 1983 877 Panicene. No species ofthese tribes were susceptible to an isolate of P. sacchari (8) “The grass accessions that were susceptible to P. philippinensis (Table I) represented 19 species in six genera in the tcibe ‘Andropogoneae (Andropogon (one species], Borhriochtoa [10 species}, Eulalia [one species}, Saccharum (one species], Schizachyriunt [three species] and Sorghum [three species}) ot four species in two genera in the tribe Maydeae (Tripsacu [one species}and Zea three species). Not all accessions that developed symptoms of systemie infection in response to one isolate of P. ‘Philippinensis did so with the other. ‘Some grass accessions were very susceptible to P. philippinensis (Table 1). These include Borhriochloa ambigua, B. edwardsiana, fone accession of B. isohaemum vat, ischaernun!, and Saccharum officinarum (sugarcane), all with 789} oF higher rates of systemic infection at the imoculum dosage used. ‘Bothriochloa ischaeniun var. ischaemum was represented by fone accession that was not susceptible to P. philippinensis and P. sacchari (one pathogen isolate of each tested), and one accession that was susceptible to both isolates of each pathogen, Eleven accessions of S. bicolor were susceptible to at least one isolate of P. philippinensis, and nine were not susceptible, This difference in susceptibility within S, bicolor probably is partly due to the heterogeneity in the plant species, However, the incidence when present, was always low. 5 bicolor Viaponicum) PI. 302174 was moderately susceptible to both isolates of P. sacchari and somewhat less susceptible to the isolates of P. philippinensts Exconde etal (6) eported that Avena saiva Lin the Philippines is susceptible to P. philippinensis; however, we were unable to infeet any of 11 species of Avena, including four accessions of A. sauiva. Ina further tes, A. sativa cultivar Clintland, besides being inoculated at $ * 10° conidia per milliliter. was inoculated at the dosage 1.0 X 10” conidia per milliliter. None of 298 plants at the Tower dosage or 261 at the higher dosage developed systemic symptoms, and the pathogen could not be recovered, ‘According to results presented here and previously (4), P. philippinensis and P. sacchari are both restricted to host species ‘within the grass tribes Andropogoneaeand Maydeae. Examinatio fof host range results for the two pathogens on specific plant fccessions within these two tribes indicatesa remarkable similarity in their host ranges, OF 26 plant accessions in these tribes susceptible to atleast one isolate of P. philippinensis. 23 also were susceptible to at least one isolate of P. sacchari (Table 1). OF 36 sccessions in the Andropogoneae and Maydeae not susceptible 0 P-philippinensis, 28 also were not susceptibleto P. saechari(Table 2), The differences betwcen pathogen species were similar to the differences between isolates within the same pathogen species. For example, of 21 accessions susceptible to atleast one isolate of P philippinenss, only 13 became infected by the other isolate, Of tight grass accessions susceptible to at least one isolate of B sacchari, seven were susceptible when tested against the other isolate. In a few instances where no infection aecurted, the number of test plants was small; had more plants been available for testing, systemic disease might have developed inat least one ora few plants ‘of a supposedly nonsusceptible combination. Several species of Andropogon, Bothrlochloa,and Schizachyriam that were susceptible in this study are common perennial forage fand wild grasses. in the United States (8). Big bluestem (Andropogon gerardii) and little bluestem (Sehizachyrium 878 PHYTOPATHOLOGY seoparium are pechaps the most prevalent constituents of wild hay inthe prairie states(8). These perennial grasses possibly could serve as reservoir hosts for P. philippinensis or P. sacchart it these pathogens entered the United States and if infected plants could Survive the winter season, Oospores were not detected in leaf tissue of any of the systemically infected plants, even when infected plants were held to maturity, However, the production of oospores by P. sacchari in sugareane has been reported (12). ‘We believe there is # very close phylogenetic relationship between P. philippinensis and P. sacchari. Besides having essentially identical environmental requirements for optimum sporulation, conidial germination, and infection (1,3) and similar shapes of conidia and conidiophores (10) their Rost ranges are femarkably similar, Further studies may indicate the desirability of ‘combining these two species. Infact, Weston (13) suggested in 1920 ‘hat all oriental forms infecting maize may be single species. Ina study of ours not reported here (M. R. Bonde, unpublished), one isolate of P. sacchari that we attempted to test from the Philippines ‘was pathogenetically very different from the isolates of P. hilippinensis irom the Philippines or P. sacchari from Taiwan. It ‘was only weakly pathogenic to maize to the extent that we could ‘ot collect enough conidia fram maize to compare the isolate's host range against those of the other pathogen isolates. The species P. sacchari needs tobe reexamined for homogeneity LITERATURE CITED 1. Bonde, M, R. 1981, Elects ofdew-period temperatureon development fof Philippine dawny mildew of maize and of temperature on conidial termination. (Abst) Phytopathology 71.765, 2, Bonde, MR und Freytag RE, 1979, Host range of an American isolate of Peranoselerosparesorgh, Plant Dis. Rep. 6:650-654, 4 onde, M, Rand Melching, J. S. 1979. Elfects of dew-peiod temperature on sporulation, gevmination of conidia and systernie Infection of maize by Peranotclerospora saecharl. Phytopathology 10841086, 4, Bonde, M.R. and Peterson, GL, 1981. Host range of a Taiwanese isolte of Prronaselerospora sacchari. Plant Dis, 65139-740. 5. De Wet, ) MJ. 1978, Systettatice an evolution of Sorghu see. ‘Sorghum (Gramineae, Ary 1 Bot. 8477-488 6, Exconde,0.R. Elec, 1-V rand Advincul, B.A. 1968, Host range of Solerasparaphlippinensis Weston inthe Philippines. Philipp. ABKe Sensis 7 Exeonde, 0. R., and Raymundo, A, D, 1978, Vield loss eavsed by Philippine corm downy mew. Philipp. Agi, 38115120. 1 Hoover, MM Hein, M_A., Dayton, W. A and Erlanson

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