Symptom expression and accumulation of potato virus Y (PVY°) and
potato leaf roll virus in thirteen potato cultivars
Jari Valkonen and Eerik Mäkäräinen
Valkonen,J.& Mäkäräinen,E. 1993.Symptom expressionand accumulation of potato virusY(PVY°)and potato leaf roll virusinthirteen potato cultivars.Agric.
Sci.Finl.2: 33-40.(Dept.PI. Product.,FIN-00014 Univ.Helsinki,Finland.) Necrotic local lesionsdeveloped incvs.Matilda,Ostara,Record,Satuma, Stina, Hank- kija’s (Hjan)Tanu andHjanTimo and localringspotsin OlympiaandSieglinde(Siikli) followingsap inoculation with theordinarystrain of potato virusY(PVY°).Secondar- ily infected cvs. Ostara,Pito, Siikli and HjanTimodevelopedleafdrop. No infected progenywasproduced by Matilda, SaturnaandHjanTanu.Incontrast,Bintje,Puikula and Sabinadeveloped neither local lesionsnorsystemic necrosis,but showed mosaic symptoms following primary andsecondaryinfection byPVY°. The ELISA absorb- ancevalues for potato leafroll virus (PLRV)inOstara, Pito and Saturnawereless than
10%of thosein the PLRV-infected Siikli. The ELISA values forPLRV in Olympia, Stina, HjanTanu andHjanTimowerenotsignificantlydifferent from those of Siikli.
The severityof the symptoms did not correlate with the concentration ofPLRV inthe potatoes.
Key words:hypersensitivity,virusaccumulation,virusresistance,potato
Introduction
Potato virus Y (PVY) and potato leaf roll virus (PLRV) are the economically most important viruses ofpotato(Solarium tuberosumL.)in Euro- pe, causing yield losses of upto80%(DeBokx and vander Want 1987).InScandinavia,PVY ismore important than PLRV (UMAERUSet al. 1979,KURP-
PA 1983).In Finland, the tobacco vein necrosis strain of PVY (PVYN) seems to be dominating, whereas the ordinary strain of PVY
(PVY°)
is the onemostcommonly encountered in otherparts of Europe (Kurppa 1983, De Bokx and van der Want 1987).Resistance inpotatotoviruses is important in the control of viral diseases. Thetypesof resistance to PVY are hypersensitivity and extreme resistance
(ROSS 1986). Localized hypersensitivity is ex- pressedas necrotic local lesions in the sap-inocu- lated leaves. Systemic hypersensitivity results in necrosis in thetop orthetopleaves of the PVY-in- fected plant. Ross (1986) defined extremeresist- ance as intensified local hypersensitivity. We de- fineextremeresistanceasthetype where the virus concentrationremains extremely low inaninfected plant. The latter definition not only includes the type of resistance in which the multiplication of PVY is reduced(Barkerand Harrison 1984),but also the possibility that the low virus concentration in plant tissue is due to the inhibited cell-to-cell spread of the virus(Valkonen etal. 1991).
The threecomponents of resistancetoPLRV in potato are restriction of virus multiplication, resist- ance to infection and inhibition of virusmovement
Agric. Sei.Fin!. 2 (1993)
from foliage to tubers (Barker and Harrison 1985, 1986, Barker 1987). The first component was considered to be the most important by Barker and Solomon (1990), who detected re- striction ofPLRV accumulation in somecultivated potato genotypes. Furthermore, extremeresistance to PLRV has been found in a few wild potato species (Jones 1979,Brown etal. 1984,Brown
1991, Valkonen etal. 1992).
The localpotatoproduction in Finland is mainly basedon Finnish, Swedish, German and Dutch cul- tivars.However,the informationabout thetypeand level of resistancetoPVY and PLRV in thepotato cultivars commonly grown in Finland is limitedand data has been obtained mostly from the field and not from experiments performed under controlled conditions (Kurppa 1983, Stegemann and Schnick 1985,Kurppaand Hassi 1989). Deter- mination of thetype and level of virus resistance in the above potato cultivars was considered impor- tantfor the strategic planning of thepotatobreeding programme for virus resistance initiatedrecently as a joint project of the Department of Plant Produc- tion, University ofHelsinki, and the Institute of Plant Breeding, Agricultural Research Centre of Finland. The present study was undertaken to test the symptom expression and accumulation of
PVY°
and PLRV in 13potato cultivars grown in FinlandasPVY°
and PLRVarethemostimportant viruses ofpotato worldwide,PVY°
usuallycauses more severe symptoms in potato than does PVYN (De Bokx and van der Want 1987),and PLRV may becomemoreimportant also in Scandinaviaas aconsequence of the forecastedwarming up of the climate(Carter 1992).Material and methods Viruses
One isolate each of PLRV and
PVY°
wasobtained from fieldgrown potatoes in Sudan and England, respectively (El-Amin et al. 1990, Gibsonet al.1990). These virus isolates were used,because the above isolate of
PVY°
is well-characterized (GIB-SON etal. 1990, Valkonenetai. 1991, 1992)and resembles the Finnish
PVY°
isolate YSFII ofKurppa (1983) in terms of biological properties, and because PLRV occurs only sporadically in Finnish potato fields and no local isolates of the virus were available.
PVY°
was maintained in Nicotiana tahacumcv.Samsun and PLRV inpotato cv.Sieglinde.Antisera
The antibodies and alkaline phosphatase conjug- ated antibodies toPVY and PLRV were obtained from Böhringer.
Plants and growing conditions
Virus-tested seedpotatoes ofcvs. Bintje, Ostara, Record and Satuma (Dutch), Olympia and Sieg- linde (Siikli) (German), Matilda,Sabina and Stina (Swedish), and Pito, Puikula, Hankkija’s (Hjan) Tanu and Hjan Timo(Finnish) (Stegemann and SCHNICK 1985) were obtained from the Finnish Seed Testing Institute, Helsinki. The tuberswere planted intopots21cmin diameter and filled with amixture of steam-sterilizedpeatand washed sand (10:1 v/v)in the beginning of July 1991, and the pots were then sunk by halfway into asand bed in anaphid-proof nethouse. The plantswerewatered daily and fertilized weekly with NPK fertiliser.
Late blight (Phytophtohora
infestans
(Mont.) deBary)wascontrolled by applications ofmethalaxyl, mancozeb and maneb. The average photoperiod was 16h and the mean temperatures at day and nightwere26°C and 16°C,respectively. The tubers wereharvestedatthe end of September and driedat room temperature for two days. After storage of four monthsat 4°C,five randomly taken progeny tubers per plantwereplanted intopotsand grown in glasshouse as described above. The photoperiod wasextendedto 18 h by illumination with fluores- cent lamps. Themean temperaturesduring day and nightwere asabove.
Agric.Sei.Fin!. 2(1993)
Virus inoculation part of the stem with its axillary bud, cutting the stem lengthways, sidegrafting aninfected scion of cv.Siikli in place of the removed bud and binding it with Parafilm. The growth of the infected scion was taken as an indication of a successful graft union and the scions were removed three weeks after grafting.
PVY°
was sap-inoculated by grinding leaves of infected tobaccoat lg/sml of distilledwaterwitha pestle and mortar, and rubbing the extract onto carborundum-dusted leaves of the potato plants.Two plants perpotatocultivarwereinoculated with PVY . The two oldest leaves of four shoots per plant were inoculatedatthe emergence of the fifth leaf of the shoot (stage of developmentno. 305) (Jefferies and Lawson 1991), whereas all other shoots werecutoff.
Virus detection
The local and systemic symptoms were visually observed 14 and 28 days after sap inoculation of
PVY°.
The secondary symptoms werevisually ob- served and the litres ofPVY°
and PLRV were determined by a double antibody sandwich en- zyme-linked immunosorbent assay (DAS-ELISA) (Clark 1981) in the progeny plants atthe appear- anceof flower buds (stage of developmentno.410) (Jefferies and Lawson 1991). For DAS-ELISA, the uppermostfully expanded leaves of the plants PLRV was graft-inoculated, which is themostsensitive method oftesting the resistancetoPLRV (Swiezynski etal. 1989).Two plants per cultivar were inoculated at the emergence of the seventh leaf of the shoot (stage of developmentno. 307) (Jefferies and Lawson 1991). Four shoots per plant were left growing; all other shootswere cut off. Two of the four shootswere then graft-inocu- lated with PLRV by removingaleaf from the upper
Fig.
1.
Local lesionsfollowingsap inoculation withPVY°.From the left: cvs.Matilda, Record,Siikliand Stina.Agric. Sei.Finl. 2(1993)
Table 1. Symptoms due toprimary infection (sap-inoculated PVY°),and symptoms and ELISA absorbance values (A405)
±standard deviation due tosecondaryinfectionbyPVY°,asdetermined at the time of flower initiation.
Primary infection symptoms Secondaryinfection no.of tubers
infected
local systemic symptoms A405values
Bintje O M 5/5 M,S,R 1.61 ±0.13
0.02±0.00 1.3610.22 1.61 ±0.17 1.7210.11 1.7210.13 1.4510.18 1.5510.30 0.0310.01
1.0710.50 0.28 NRS,VN
RS
VN, IVN VN,CS
0/5 O
Matilda Olympia Ostara
4/5 M
NS O 2/5 M.LD
M, LD M.R O
O
VN,CS 5/5
Pito Puikula Record
Sabina Satuma Siikli
M 4/5
NS O 3/5 M
o M 5/5 M
NRS,VN RS
VN 0/5 O
NRS,VN VN, IVN
2/5 M, LD
M, LD NRS,VN
NRS
1/5
Stina HjanTanu
HjanTimo
O
o 0/5
O 0.0310.01
1.3210.29
NRS 4/5 M.LD
non-inoculated
cultivars 0.02+0.01
O =nosymptoms CS =chlorotic spots IVN =interveinal necrosis LD =leaf-drop
M =mosaic
NRS=necroticringspots NS =necrotic spots R =rugosity S =stunting VN =vein necrosis RS =ringspots
were sampled in duplicate, extracted at lg/3ml of buffer and pipetted onto ELISA plates (Greiner Labortechnik) in aliquots of 200 pi. Absorbances wererecorded at 405 nm (A405) after developing the colour reaction for 1 h using p-nitrophenyl as substrate.
Statistical analysis
Analysis of variance was used for the statistical analysis of theA405 values and calculations of the least significant differences (LSD) when appropri- ate (Steeland Torrie 1981).
Results
PVY°
Necrotic local lesionswere observed incvs. Ma- tilda, Ostara, Record, Satuma, Stina,Hjan Tanu and Hjan Timo following sap inoculation with
PVY°
(Table 1),whereas Olympia and Siikli developed ring spots that remained green while the leaves turned yellow (Fig. 1).Bintje, Pito, Puikula and Sabina showedno symptoms in the sap-inoculated
leaves.
Systemic vein necrosisfollowing primary infec- tion was observed in Matilda, Olympia,Pito, Sa-
Agric. Sei.Fint.2(1993)
turna,Siikli and Stina(Table 1).Systemic mosaic symptoms without necrosis were observed in Bintje, Puikula andSabina, whileOstara, Record, Hjan Tanu and Hjan Timowerefree of any.
Systemic mosaicsymptoms wereobserved in the progenyplants oftenof the 13 cultivars following secondary infection by
PVY°
(Table 1).Further- more,Ostara, Pito,Siikli and Hjan Timo developed leaf drop, Bintje was heavily stunted and the growth of Puikula waspoor.Incontrast, the prog- eny ofMatilda, Saturna, Stina and Hjan Tanuwere free ofsymptomsandPVY°
accordingtoELISA.PLRV
All the tested progeny tubers of each cultivar graft- inoculated with PLRV were infected with PLRV according to ELISA. However, the cultivars fell into three groups according to the PLRV litres (P=0.01)(Fig.2).Olympia.Siikli, Stina,Hjan Tanu and Hjan Timo yielded high PLRV litres (A
405
>1.33),Matilda,Puikula and Record yielded moder- ate PLRV litres (0.90<A405< 0.96), andOstara, Pito and Saturna yielded low PLRV litres(A405<
0.66). The PLRV litres of Bintje ranged between low and moderate and those of Sabina between moderate and high (LSDo.oi= 0.34 for the A405
values).
PLRV caused yellowing of the leaves and leaf roll in Matilda, Ostara, Siikli and Hjan Timo,yel- lowing of the leaves without leaf roll in Olympia, Puikula,Saturna and Hjan Tanu, and yellowing of the leaf margins in Pito. Symptomless infection by PLRV wasdetected in Bintje, Record,Sabina and
Stina.
Discussion
Hypersensitivity reactions wereobserved in 10 of the 13 cultivars following infection with
PVY°.
Expression of hypersensitivity isuseful, assuch a response inducedbyavirusorvirus strain inaplant
Fig.2.Absorbance values (A405) (ELISA) and their least significant differences atarisk level of1%(LSDo.oi=
0.34) for detection ofPLRV inthefoliageof the progeny ofpotato cultivarsgraft-inoculatedwithPLRV.The dilution of the sap fromcv.Siikliwas4 - 4000-fold and that of the other cultivars 4-fold. A405 values for uninfected cultivarsweresimilar to those for the 4000-fold dilution of the sap of PLRV-infectedcv.Siikli.
Agric.Sei.Finl.2 (1993)
canreduce the systemic spread of the virus(Fritig etal. 1987).Indeed, no infected progeny wasde- tected in Matilda,Saturna and Hjan Tanu and few infected progeny tubers were produced by Stina.
These cultivars reacted by local necrosistoprimary infection of
PVY°.
Similar resultswereobtained in Matilda byKurppaand Hassi (1989).There are few studies on the genetic control of hypersensitivity to PVY in potato. Cockerham (1970) identified four genes which control the hy- persensitivity toall strains of PVY ina number of Solarium spp., of which the genes in S. chacoense Bitt. and S. demissum Lindl. acted also against po- tatovirus A(PVA). Jones (1990) suggested that the strain group specific hypersensitivity to
PVY°
in certainpotato genotypes wascontrolled byasingle dominant gene, Nytbr, which possibly originated in Kalahdin and a Scottish potato clone 11-79 (Davidson 1980), and by another dominant gene of unknown origin. Our results donotallow further comparisons of the genes for hypersensitivity to PVY between the cultivars ofour study and those reported by Cockerham(1970)and Jones(1990), as those comparisons would require inoculations with other strains of PVY and PVA.However, the hypersensitivitytoPVY°
in HjanTimo, acultivar derived from the cross Friihnudel x Kalahdin (Varis 1975),is presumably controlled by the gene Nytbr from Kalahdin.The progenies of Bintje, Puikula and Sabina werealmost 100% infected with PVY0.The plants generated high PVY litres and showedsevere mo- saic symptomswithout necrosis. The susceptibility of these cultivarstoPVY hampers their cultivation in Southern Scandinavia. Therefore, Puikula and Sabina aremainly grown in the northernmostareas, where the spread of PVY is reduced duetothe low populations of the PVY transmitting species of aphids (Umaerus et al. 1979, Sigvald 1984,
KurppaandRajala 1986).
The incidence of PLRV-infected progeny tubers wassimilarly high in all of the cultivars.However, there werevariations in the PLRV concentrations between the progenies of cultivars. Theconcentra- tion of PLRV in cultivars suchas Ostara, Pito and Saturna was less than 10% of that in Siikli. Re- stricted accumulation of PLRV is useful inpotato
as it potentially reduces the spread of PLRV by aphids (Barker and Harrison 1986).
The severity of thesymptomsand theconcentra- tions of PLRV didnotstrictly correlate in thepotato genotypes tested. For example, apparent leaf roll and yellowing symptoms wereobserved in Ostara which exhibited the lowest PLRV concentration of all the cultivars. In contrast, no symptoms were observed in Stina which generated highconcentra- tions of PLRV. The severity ofsymptoms observed in the PLRV-infected progeny ofpotato correlates rather with the physiological stage of the mother plantsatthe time ofinoculation than with the PLRV concentrations in the progeny plants (Barker and Harrison 1986,Barker and Woodford 1987).
The valuabletypes of virusresistance,i.e. hyper- sensitivityto
PVY°
and restricted accumulation of PLRV, were incorporated in e.g. Ostara and Sa-tuina. However, nocultivar exhibitedextremere- sistance to PVY and PLRV. Genes for extreme resistancearethe obvious choicetobe incorporated in new cultivars (Jones 1990),because extreme resistance actsagainstmost, ifnot all strains ofa virus and efficiently reduces the virus transmission by aphids(Jones 1979, 1990, Gibsonetal. 1990).
Furthermore, virus strains cabable of overcoming extreme resistance are not common in the field (Jones 1985, Ross 1986). Extreme resistance to PVY, PLRV and PVA is incorporated in the wild potato species of the Etuberosa group, e.g. S. hre- videos Phil. (VALKONEN etal. 1992), and it has recently been transfered tosomecultivatedpotato genotypes(Pehu etal. 1990, Williamsetal. 1990, Xuetal.,unpublished results). The resultssuggest higher acceptability and productivity formostcul- tivars of thepresentstudy in Southern Scandinavia and otherpartsof Europe, where PLRV and
PVY°
arethemostdisasterous viruses inpotato, if genes forextremeresistancetoPLRV and PVY could be incorporated into them.
Acknowledgement. The financial assistance to J. Valkonen (grant 2777/501) from the Finnish Ministry ofAgriculture and Forestry is gratefully acknowledged.
Agric. Sei.Finl.2(1993)
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Agric. Sei.Fin!.2(1993)
Manuscriptreceived November1992 Jari Valkonen
Eerik Mäkäräinen
Department ofPlant Production P.O. Box27(Viikki)
FIN-00014 UniversityofHelsinki,Finland
SELOSTUS
Perunan Y-viruksen japerunankierrelehtisyysviruksen oireetja pitoisuus kolmessatoista perunalajikkeessa
Jari Valkonen ja Eerik Mäkäräinen Helsingin yliopisto
Perunan Y-virus (PVY) ja perunankierrelehtisyysvirus (PLRV) ovat haitallisimmat perunaa infektoivat virukset maailmanlaajuisesti, jaerityisesti PVYonmerkittävä satotap- pioiden aiheuttaja Pohjoismaissa. Kolmentoista Suomessa yleisesti viljeltävän perunalajikkeen kestävyyttä PVY:n yleistä rotua (PVY°) jaPLRV:ta vastaan tutkittiin, sillä käytettävissä ei ollut rittävästi tietoakyseisten lajikkeiden PVY-ja PLRV-kestävyydestäperunan virusresistenssijalos- tusohjelmansuunnittelunperustaksi.
PVY:lla saastutettuihin lehtiin muodostui kuoliolaikkuja lajikkeissa Matilda,Ostara,Record, Satuma, Stina,Hankkijan (Hjan) Tanu ja Hjan Timo,kun taasOlympiassa ja Sieg- lindessa (Siikli) havaittiin rengaslaikkuja. Saastuttamat- tomissaylälehdissähavaittiin(systeemistä) suonikuoliota la- jikkeissa Matilda, Olympia, Pito, Saturna, Siikli ja Stina.
Näissälajikkeissa ilmeniN-geenien tuottamaahypersensitii- visyyttäPVY°:tavastaan.Sensijaan Bintjessä,Puikulassaja Sabinassa ilmeni systeemistä viherkirjoa ilman kuoliota.
PVY:n saastuttamista mukuloista kasvaneiden Ostaran, Pidon, Siiklinja Hjan Timon taimien alalehdet kuolivat ja
jäivätroikkumaan varresta. PVYdlasaastutettujen Matildan, Satuman ja Hjan Tanun mukuloistayksikään ei tuottanut virussaastuneitataimia,kun taas muiden PVYdla saastutettu- jen lajikkeiden mukuloista kasvaneissa taimissa ilmeni viher- kirjoa, ja taimissa havaittiin korkeita PVY-pitoisuuksia
ELISA-testeissä.
Kaikkilajikkeet, jotkaoli saastutettu PLRVdIa varttumalla PLRV-infektoitunut perunanverso tutkittavan lajikkeeseen, tuottivat PLRV-saastuneita mukuloita. Saastuneista muku- loista kasvaneiden taimienPLRV-pitoisuusvaihtelilajikkeit- tainsiten, ettäseolialhainen Ostarassa, PidossajaSaturnassa, keskinkertainen Matildassa,PuikulassajaRecordissa, jakor- kea Olympiassa, Siiklissä, Stinassa,Hjan Tanussa ja Hjan Timossa.Bintjen PLRV-pitoisuus sijoittuioli alhaisenjakes- kinkertaisen välille ja Sabinan keskinkertaisen jakorkean välille. PLRV aiheutti lehtien kellastumista ja kiertymistä Matildassa,Ostarassa, Siiklissäja Hjan Timossa, pelkästään lehtien kellastumista Olympiassa, Puikulassa, Sallimassa ja Hjan Tanussa jalehtien reunojen kellastumista Pidossa.
PLRV-saastuneessa Bintjessä, Recordissa, Sabinassa jaSti- nassaei havaittu oireita. Siten lajikkeiden PLRV-pitoisuuden jaoireiden voimakkuuden välillä ei ollutjohdonmukaistariip- puvuutta.
Ostarassa jaSaturnassa yhdistyivät yliherkkyys (hypersen- sitiivisyys) PVY°:ta vastaan ja alhainen PLRV-pitoisuus, mikä voi merkittävästi ehkäistäPVY°:njaPLRV;nleviämis- tänäissälajikkeissa.Sensijaanmissäänlajikkeessaei havaittu äärimmäistä R-geenien tuottamaa kestävyyttä PVY:ta ja PLRV:ta vastaan.
Agric. Sei.Fint.2(1993)