Inheritance of tolerance ofGlobodera pallida and the relationship between tolerance and resistance toG. pallida in potatoes

Summary The tolerance of and resistance toGlobodera pallida of two potato progenies which segregated for these characteristics were assessed in pots in the glasshouse. The level of tolerance varied widely between genotypes within the progenies, with several genotypes being significantly more tolerant than the intolerant parent. The levels of resistance and tolerance in the progenies were not significantly correlated. No indications were obtained that the ranking of genotypes for tolerance is influenced by the density ofG. pallida in the soil. The results confirm that selection for tolerance in aG. pallida resistance breeding programme is useful and feasible.     

Screening tuber-bearingSolanum spp. for resistance toGlobodera rostochiensis Ro1 Woll. andG. pallida Pa2/3 stone

Summary Accessions of tuber-bearingSolanum spp. related toS. tuberosum subsp.tuberosum were obtained from the German-Dutch collection (Braunschweig, Germany) and the Inter-regional potato collection (Sturgeon Bay, USA). They were screened for resistance toG. rostochiensis Rol andG. pallida Pa2/3. Among 1567 clones from 52 accessions, 135 clones (23 accessions) were resistant toG. rostochiensis. They mainly representedS. andigena, gourlayi, spegazzinii andvernei. Among 1689 clones (74 accessions), 105 clones (32 accessions) were resistant toG. pallida. They representedS. gourlayi, spegazzinii, sparsipilum andvernei. About 25 clones were resistant to both species.     

Monoxenischer Resistenztest für Kartoffelgenotypen gegenüber dem ZystennematodenGlobodera pallida. 3. Anwendung und Prüfung des Tests

Zusammenfassung Zur Resistenzselektion von Kartoffelgenotypen gegenGlobodera pallida (Stone) Beharens (Pa3) wurde ein monoxenischerin vitro-Test entwickelt, der die Auswertung geringer quantitativer Unterschiede in der vermutlich polygen-vererbten Resistenzreaktion erlaubt. Hierzu wurde das aussagekr?ftigste Resistenzkriterium an 30 unterschiedlich resistenten Genotypen bestimmt. An den anf?lligen undG. rostochiensis-resistenten Genotypen entwickelte sich immer ein h?herer Anteil Weibchen als M?nnchen, im Gegensatz zu denG. pallida-resistenten Genotypen, bei denen die M?nnchen überwogen. Eine weitere Unterteilung derG. pallida-resistenten Genotypen war durch das erh?hte Auftreten nekrotisierter Wurzelbereiche und durch Larven mit stagnierender Entwicklung m?glich. Zur Bewertung des Einflusses der Genotypen auf die charakteristische Populationsentwicklung des Nematoden wurde ein Schema mit vier Gruppen erstellt. Die überprüfung der untersuchten Genotypen in einem Topfkulturetest zeigte eine übereinstimmung mit den Ergebnissen aus dem monoxenischen Resistenztests.

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Summary A monoxenicin vitro test, which was developed for selecting resistance in potato genotypes toGlobodera pallida (Stone) Behrens, can distinguish small differences in the resistance reaction, presumed to be polygenically controlled. To establish meaningful criteria for resistance, a number of characters which could be assessed rapidly were determined (number of necrotic root areas, number of invading juveniles but not yet sexually differentiated, number of juveniles developing as males and females, number of juveniles with arrested development). Assessments were made 21, 35 and 49 days after the introduction of 1000 freshly hatched juveniles ofG. pallida Pa3- population ‘Frenswegen’ per test Petri dish. Because of differences in levels of juvenile invasion per genotype (number of necrotic root areas) and differentiated individuals (Table 1), to make genotypic comparisons, it was necessary to consider other resistance criteria along with percentage juvenilie invasion. The ratio of males to females was shown to be the most reliable criterion of resistance. There was a higher proportion of females than males on susceptible, andG. rostochiensis resistant genotypes than onG. pallida resistant genotypes on which males predominated (Table 1). A further subdivision of theG. pallida resistant genotypes was possible using the increased occurrence of necrotic root areas, and juveniles with arrested development. A scheme was constructed for rating the effect of genotypes on the characteristic development of the nematode populations. It contained four groups: susceptible reaction (Figure 1), weakly expressed resistance (Figure 2), expressed resistance (Figure 3) and strongly expressed resistance (Figure 4). Female counts from pot tests with particular genotypes were in general agreement with the results from the monoxenic resistance test (Table 1).

     

A glasshouse test for assessing resistance to early blight (Alternaria solani)

Summary Fourteen genotypes of potato were assessed for resistance toAlternaria solani Sor., both in the glasshouse at the Scottish Crop Research Institute in Scotland and in the field in the Negev, Israel. Glasshouse assessments agreed well with field results, identifying the most resistant and susceptible genotypes. The effects on the glasshouse test of plant age, inoculum concentration, and length of time plants were kept damp after inoculation, were examined. The best agreement with field results was shown by plants inoculated at flowering, and the best discrimination between genotypes was achieved by keeping the foliage damp for 24 h after inoculating with a spore suspension containing 2.5×10³ spores/ml.     

The expression of resistance to the white potato cyst nematode (Globodera pallida) by individual potato seedlings under different inoculation treatments

Summary The resistance toGlobodera pallida of individual potato seedlings from three progenies derived fromSolanum tuberosum ssp.andigena CPC 2802 was assessed using three inoculation treatments. Plants grown from tubers from the seedlings were used to reassess resistance. All the seedling test inoculation treatments over-estimated the proportion of clones classified as resistant compared to the results from tuber-grown plants. However, the mean relative resistance of progenies estimated by both tests showed good agreement, and it is suggested that seedlings are best used for assessing the resistance of progenies of crosses to identify promising combinations rather than individual clones.     

The effect of soil pH on yield losses caused by the potato cyst nematodeGlobodera pallida

Summary The effect of soil pH on yield losses due to potato cyst nematodes (Globodera pallida) was investigated in experiments in the field and in a semi-controlled environment. Through annually liming a reclaimed peat soil with about 22% organic matter, plots with pH levels of 4.5, 5.0, 5.5, 6.0 and 6.5 were obtained. In 1990 potato yields in these plots decreased from 45 t/ha at pH 4.5 to 33 t/ha at pH 6.5, whereas the nematode densitites decreased from about 18 to 9 juveniles per g soil. In a container experiment a strong interaction was recorded between soil pH and the presence of nematodes. Tuber yields were about 11% lower at pH 6.5 than at pH 4.5 in the absence of nematodes, but about 44% lower when an initial population of 27 juveniles per g soil was present.     

The use of single,spaced potato plants to assess their tolerance of damage by potato cyst nematodes

Summary Potato clones and cultivars differ in their ability to tolerate damage by the potato cyst nematodes:G. rostochiensis andG. pallida. This paper reports a simplified field assessment whereby single, spaced plants are grown in heavily infested soils. These trials identified those clones which are either very sensitive to damage or those that grow and yield well in the presence of nematodes. The use of such a screening procedure in a breeding programme is discussed.     

Relationship between potato cyst nematodes and their principal host. IV. Tolerance of potato cultivars and the effect of potato cyst nematode on initial plant growth

Summary Greenhouse experiments on the effects of white potato cyst nematode infestations (Globodera pallida Stone) on initial growth and development of a series of potato cultivars are compared with the results of field experiments on sandy and sandy-peat soils on the effect of nematode density on tuber yield. A simple greenhouse test, assessing root growth response to potato cyst nematode infection, provided a good insight into a cultivar's tolerance performance in the field early in the growing season. As a very limited number of plants is needed for the greenhouse test, screening for tolerance can be conducted in the early stages of a breeding programme.     

The effect of seed-and soilborne inoculum ofColletotrichum coccodes on the incidence of black dot on potatoes

Summary Soil in a potato field naturally infested with black dot (Colletotrichum coccodes) was fumigated with methyl bromide at 126 g m⁻² or left unfumigated. Potato seed tubers (cv. BP1) uninfected, lightly infected (1–25% surface affected), severely infected (26–100% surface affected) and severely infected withC. coccodes but dusted with prochloraz manganese chloride as Octave 2.5% DP at 750 g per 100 kg seed were planted in fumigated and unfumigated soil. When harvested, the incidence of black dot on the progeny of infected seed planted in unfumigated soil was twice that of progeny in fumigated soil, with progeny of uninfected seed having a 68.5 times higher disease incidence in unfumigated soil. Black dot on progeny tubers was reduced by pre-treatment of seed with prochloraz in fumigated soil only. With black dot infested fields, planting disease-free seed or treating seed with fungicides would not decrease disease on progeny tubers.     

Incidence, appearance and development of potato mop-top furovirus-induced spraing in potato cultivars and the influence on yield, distribution in Denmark and detection of the virus in tubers by ELISA

Summary 23 potato cultivars were grown in fields infested with potato mop-top virus and spraing was assessed on tubers at harvest and after storage. Large differences in spraing susceptibility were found between cultivars. Compared with other countries, spraing in Denmark is characterised by a high incidence at harvest, a low increase during storage and a very low proportion of superficial spraing. Decrease in spraing during storage was recorded in two cultivars. DASELISA tests for PMTV in tubers revealted a high reliability. Mapping of PMTV in important Danish potato growing areas showed that the virus is widespread. Occurrence of spraing did not influence total yield or dry matter content. Soil acidity did not influence incidence of spraing but it was more common on coarse-grained soil than on finer sandy soils. PMTV in viruliferous resting spores ofSpongospora subterranea was inactivated by heating to 90°C for 15 minutes.     

Inheritance of resistance toFusarium sulphureum in crosses betweenS. tuberosum potato cultivars measured on field and glasshouse grown tubers

Summary Solanum tuberosum potato cultivars with differing levels of resistance to dry rot caused byFusarium sulphureum were hybridised in a half diallel crossing programme. Glasshouse grown tubers from true seedlings and field-grown tubers of the resulting progenies were assessed for their resistance toF. sulphureum. Statistical analysis revealed differences between the progenies forF. sulphureum resistance on both field and glasshouse grown tubers. On the field grown tubers, differences between progenies were due to differences in the general combining ability (GCA) of the parents. Parental phenotypes and GCA values were highly correlated. Overall. GCA was consistent for resistance or susceptibility, and glasshouse grown progenies showed a good correlation for GCA between years (r=0.85^***), and with the results of the field grown progenies (r=0.61^* in 1998 and r=0.69^** in 1999). It appears that resistance toF. sulphureum is heritable, and that this heritability is better assessed on field grown rather than glasshouse grown tubers, although the two are largely in agreement.     

Synergistic interactions betweenRhizoctonia solani Kühn,Verticillium dahliae Kleb.,Meloidogyne spp. andPratylenchus neglectus (Rensch) Chitwood & Oteifa,in potato

Summary Samples of a sandy soil and a marine clay soil sterilized by steam were put in 55-1 containers insulated with polystyrene and placed outdoors on a brick pavement. Sandy soil was infested singly or in all possible combinations with root-knot nematodes (Meloidogyne spp.) and the fungiRhizoctonia solani andVerticillium dahliae, and the marine clay soil was infested with the root-lesion nematodePratylenchus neglectus and the same fungi to evaluate the effects of these organisms on the yield of potato. The experiments were carried out from 1983 to 1986. Tuber yield was reduced by single infestations of theMeloidogyne spp. andV. dahliae but not significantly byR. solani orP. neglectus. A three-factor interaction: nematode ×R. solani × V. dahliae was found in both experiments.R. solani andV. dahliae showed significant synergistic effects when soil was infested with theMeloidogyne spp. orP. neglectus.     

A method of assessing potato seedling progenies for resistance to the white potato cyst nematode

Summary The resistance expressed by progenies of potato seedlings to the white potato cyst nematode (Globodera pallida) has been assessed on bulk sowings of seedlings and on individual plants raised from tubers, and a positive correlation between the seedling and plant tests has been established. The use of such a seedling test as an addition to other methods of screening is discussed.     

Extreme resistance to infection by potato virus X in genotypes of wild tuber-bearingSolanum species

Summary Clones derived from thirty-one different accessions (nineteen of Argentine origin) belonging to eightSolanum species were screened for resistance to infection by potato virus X strain cp (PVX cp) by mechanical inoculation of plantlets that had been micropropagated in vitro. Estimates of PVX multiplication obtained by enzyme linked immunosorbent assay and slot blot nucleic acid hybridization allowed the identification of resistant clones derived from five accessions belonging toS. commersonii S. oplocense, S. sparsipilum andS. tuberosum andigena. Resistant genotypes supported PVX concentrations 5 to 15 times smaller than did the susceptible control cultivar Spunta. Graft inoculation test confirmed the presence of extreme resistance similar to that conferred by the ‘immunity’ gene X¹ (also called RX_act).     

Reaction toPhytophthora infestans of potato genotypes with race-specific resistance

Summary Phytophthora infestans isolates collected in Poland in 1987–1995 were evaluated on detached leaflets of a differential series consisting of potato genotypes possessing resistance genes R1–R11. On the same differentials was evaluated over three years a highly virulentP. infestans isolate MP 245. Isolate MP 245 and those from natural populations ofP. infestans were seldom or inconsistently pathogenic to R5, R8 and R9. When leaflets of the differentials were infected with MP 245 the lesions were often small and sporulation weak. A statistically significant interaction was found between years and pathogenicity of MP 245 to individual differentials. Attempts to train the isolate MP 245 for pathogenicity to R9 were unsuccessful. The variation in specific resistance of potato genotypes may complicate the evaluation of their resistance toP. infestans, as well as the evaluation of the virulence spectrum of natural fungus populations. This variation may be due to changes in specific pathogenicity of the fungus or in specific resistance of the potato. Some differentials have a possibly useful resistance toP. infestans.     

Erwinia carotovora subsp.atroseptica infection promotes verticillium wilt development in potato in Israel

Summary The interaction betweenErwinia carotovora subsp.atroseptica (Eca) andVerticillium dahliae and its effect on symptom development in potato cultivars showing different degrees of resistance to them was examined over two seasons in irrigated fields in a hot, dry climate. Four cultivars were used: Cara, highly resistant to blackleg and tolerant to Verticillium wilt; Pentland Crown also resistant to blackleg but susceptible to Verticillium wilt; Désirée, moderately susceptible to blackleg but tolerant to Verticillium wilt; and Maris Bard, susceptible to both diseases. Seed tubers were inoculated with Eca immediately after harvest in Scotland, to simulate natural infection, and then planted in Israel in soil naturally infested byV. dahliae. In Verticillium tolerant cultivars, Eca infection increased the rate of colonization byV. dahliae and significantly enhanced symptom development, as assessed by suppression of plant height and wilt symptoms. This report is a portion of the Ph.D. thesis of the senior author in Ben-Gurion University of the Negev.     

Monoxenischer Resistenztest für Kartoffelgenotypen gegenüber dem ZystennematodenGlobodera pallida. 2. Durchführung des Tests mit Nematoden

Zusammenfassung Durch die monoxenischein vitro-Kultivierung k?nnen geringe quantitative Unterschiede in der vermutlich polygen-vererbten Resistenzreaktion von Kartoffelgenotypen gegenüberGlobodera pallida (Stone) Behrens (Pa3) festgestellt und für die Selektion verwendet werden. Grundvoraussetzung hierfür ist die einfache und reproduzierbarein vitro-Kultivierung vonG. pallida. Für die Schlupfaktivierung, die Oberfl?chensterilisation und die Zugabe definierter Larvendichten wurden routinem?ssig durchführbare Verfahren entwickelt, die einen vollst?ndigen Lebenszyklus der Nematoden unter monoxenischen Bedingungen erm?glichen. Die aus dieserin vitro-Population gewonnenen keimfreien Zysten k?nnen für eine vereinfachte Resistenzprüfung eingesetzt werden. Die Testauswertung orientiert sich an der Entwicklung der in die Wurzeln eingedrungenen Larven. Sie kann mit einem Stereomikroskop kontinuierlich beobachtet werden.

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Summary A monoxenicin vitro culture technique can distinguish small differences in the presumed polygenically determined resistance reactions of potato genotypes toGlobodera pallida (Stone) Behrens (Pa3) and can be used in selection. The basis for such a resistance test is the simple and reproducible cultivation ofG. pallida with the aim of achieving the complete life cycle of the nematode under monoxenic conditions. A routine practical technique was developed for hatching the juveniles in a closed system (Figure 1). This was achieved by the introduction of a hatching solution which was a mixture of root washings and 0.6 mmol l⁻¹ picrolonic acid with a pH of 5.9 (=pH value of the nutrient agar). Using special hatching tubes the juveniles from 1000 cysts per tube could be hatched and counted within 2 or 3 days, as shown by experiment (Figure 3). Finally, the juveniles were surface sterilised in a special apparatus and distributed serially to the test genotypes (Figure 2). A density of 1000 juveniles per genotype gave the highest nematode infection (Figure 4) but allowance must be made for the surface sterilisation which in giving a high standard of asepsis killed up to 80% of the juveniles. At the same time especially vital juveniles were selected, thereby guaranteeing that the test was stringent and unequivocal. A variant of the technique used germ-free cysts which had been formed under monoxenic conditions. These were removed and crushed. The hatching effect of the genotypes' roots meant that 500 nematode eggs were sufficient for high infection. The test scores were based on the development within the roots of invading juveniles which could be observed continuously with a stereomicroscope. Males (Figure 5) and females (Figure 6) could be easily distinguished at low magnification (x10) 3–4 weeks and 4–6 weeks after the introduction of surface sterilised juveniles and monoxenically cultivated cysts respectively.

     

Comparison of resistance toFusarium spp. of glasshouse-and field-grown tuber progenies of potato

Summary Field-grown tubers of 22 progenies ofSolanum tuberosum L. generated in a crossing programme involving seven parents differing in resistance toFusarium coeruleum Lib. ex Sacc. andF. sulphureum Schlect. (=F. sambucinum Fuckel, teleomorphGibberella pulicaris (Fr.) Sacc.) were wound-inoculated with a cornmeal + sand culture of each pathogen. Parental genotypes were also included. The mean lesion size of each progeny was compared in 2 years of tests, as well as with published data on glasshouse-grown tubers. ForF. coeruleum there was a high correlation between years as well as with the glasshouse data, but no such correlations were apparent withG. pulicaris. Furthermore, parental and GCA values, as well as progeny means and mid-parent scores, also correlated highly forF. coeruleum but not forG. pulicaris. Glasshouse-grown tubers of 11 wildSolanum spp. were also inoculated with both pathogens. Some resistance to one or other, or both, was apparent, particularly inS. chacoense.     

Resistance to gangrene (Phoma exigua var.foveata) and dry rot (Fusarium solani var.coeruleum) in potato tubers. 2. Evaluation of genotypic differences in pectin-bound magnesium and calcium for screening purposes

Summary The levels of incorporation of Mg and Ca into tuber cell walls and middle lamellae were compared with the intensity of attack byPhoma exigua var.foveata (P.e.f.) (gangrene) andFusarium solani var.coeruleum (F.s.c.) (dry rot) in six genotypes varying in their levels of resistance. The Mg:Ca ratio was higher in those genotypes more resistant toP.e.f., while the cation effect on attack byF.s.c was variable. Although genotype resistance may be improved by extra Mg bridges in the pectin structure the genotypic differences in susceptibilty to gangrene could not be explained by differences in Mg content assessed from alcohol-insoluble solids. Analyses of breeders' selections showed no general relationship between Mg or Ca contents and disease incidence. Therefore, genotypic differences in incorporation of these cations could not be used for screening for improved resistance.