Variable response of open-pollinated seedling progeny of avocado to Phytophthora root rot

Phytophthora root rot, caused byPhytophthora cinnamomi Rands, is the most important disease of avocado (Persea americana Miller). In an attempt to identify root rot-resistant rootstocks that could ultimately be used under conditions in southern Florida, we screened open-pollinated progeny of avocado from the National Germplasm Repository in Miami. From 1996 to 1998, a total of 2,355 seedlings from 51 accessions were examined in potting mix artificially infested withP. cinnamomi. Most seedlings developed severe root rot, but tolerance was observed in some families (i.e., progeny of certain accessions). Although the most susceptible families developed mean disease ratings of up to 97% root necrosis, mean ratings for the most tolerant families were less than 60%. There was also a strong relationship between the racial background of the female parent and the tolerance of seedlings. Seedlings of the West Indian race and hybrids between it and the Guatemalan race were significantly more tolerant than those from other parents (P< 0.05). Individuals in several families developed < 50% root necrosis, the arbitrary standard of tolerance in this study. Twelve families accounted for 82% (188 of 229) of the tolerant seedlings, and only two of these did not have a West Indian or Guatemalan × West Indian pedigree. Broad-sense heritability for PRR tolerance was 0.45. This is the first report on the inheritance of PRR tolerance in avocado and on the influence of genotype and racial pedigree under controlled conditions.     

Histopathology of infection and colonization of Quercus ilex fine roots by Phytophthora cinnamomi

Quercus ilex is one of the European forest species most susceptible to root rot caused by the oomycete Phytophthora cinnamomi. This disease contributes to holm oak decline, a particularly serious problem in the ‘dehesas’ ecosystem of the southwestern Iberian Peninsula. This work describes the host–pathogen interaction of Q. ilex and P. cinnamomi, using new infection indices at the tissue level. Fine roots of 6‐month‐old saplings inoculated with P. cinnamomi were examined by light microscopy and a random pool of images was analysed in order to calculate different indices based on the measured area of pathogen structures. In the early stages of invasion, P. cinnamomi colonizes the apoplast and penetrates cortical cells with somatic structures. On reaching the parenchymatous tissues of the central cylinder, the pathogen develops different reproductive and survival structures inside the cells and then expands through the vascular system of the root. Some host responses were identified, such as cell wall thickening, accumulation of phenolic compounds in the middle lamella of sclerenchyma tissues, and mucilage secretion blocking vascular cells. New insights into the behaviour of P. cinnamomi inside fine roots are described. Host responses fail due to rapid expansion of the pathogen and a change in its behaviour from biotrophic to necrotrophic.     

Selection for decreased sensitivity to phosphite in Phytophthora cinnamomi with prolonged use of fungicide

To test the hypothesis that resistance in Phytophthora cinnamomi to control by the fungicide phosphite (phosphonate) would arise in sites with prolonged use of phosphite, 30 P. cinnamomi isolates were collected from a range of sites with different phosphite‐use histories, including phosphite‐treated and untreated avocado orchards, and phosphite‐treated and untreated native vegetation sites. The colonizing ability of these isolates was tested by different inoculation methods against a range of host tissues, treated and untreated with phosphite, including mycelial stem inoculation on clonally propagated Leucadendron sp., mycelial root inoculation of lupin seedlings and zoospore inoculation of Eucalyptus sieberi cotyledons. Isolates from avocado orchards with a long history of phosphite use were, on average, more extensive colonizers of the phosphite‐treated Leucadendron sp., lupin seedling roots and Eucalyptus sieberi cotyledons. These isolates did not colonize untreated plant tissue (Leucadendron sp.) more extensively than isolates from sites with no history of phosphite use and no isolates were resistant to control by phosphite. Analysis of all isolates with microsatellite markers revealed the majority were from a single clonal lineage. Selection for decreased sensitivity to phosphite in planta has taken place within asexual clonal lineages of P. cinnamomi in sites with prolonged use of phosphite.     

Age‐related susceptibility of Eucalyptus species to Phytophthora boodjera

Phytophthora boodjera is a newly described pathogen causing damping off and mortality of Eucalyptus seedlings in Western Australian nurseries. This study evaluated the age‐related susceptibility of several taxa of mallee Eucalyptus to P. boodjera in sterilized washed river sand‐infestation pot trials. Phytophthora cinnamomi and P. arenaria were included for comparison. Seedlings of Eucalyptus taxa were inoculated at 0, 2, 4, 12 and 88 weeks with individual Phytophthora isolates. Pre‐emergent mortality in the presence of Phytophthora was almost 100%. Post‐emergent mortality was 50–100%, depending on isolate, compared to 0% for the control. Mortality was also high for inoculated 1 month‐old seedlings (46–68%) and root length of surviving seedlings was severely reduced. Death from root infection was not observed for seedlings inoculated at 12 and 88 weeks, but they developed root necrosis and reduced root dry weight compared to non‐inoculated controls. Phytophthora boodjera is a pre‐ and post‐emergent pathogen of mallee eucalypts. These eucalypts are susceptible to P. boodjera at all life stages tested, but the mortality rates declined with plant age. Similar results were obtained for P. cinnamomi and P. arenaria. The events leading to its recent appearance in the nurseries remain unknown and further investigations are underway to determine if this is an introduced or endemic pathogen. The approach used here to understand the impact of a Phytophthora species on multiple hosts at different seedling ages is novel and sets a benchmark for future work.     

Expression of defence‐related genes in stems and leaves of resistant and susceptible field pea (Pisum sativum) during infection by Phoma koolunga

The differential expression of 13 defence‐related genes during Phoma koolunga infection of stems and leaves of susceptible versus resistant field pea (Pisum sativum) was determined using qRT‐PCR. Expression, in terms of relative mRNA level ratios, of genes encoding ferredoxin NADP oxidoreductase, 6a‐hydroxymaackiain methyltransferase (hmm6), chalcone synthase (PSCHS3) and ascorbate peroxidase in leaves and stems differed during 6–72 hours post‐inoculation (hpi) and reflected known host resistance levels in leaves versus stems. In comparison to the susceptible genotype, at 24, 48 and 72 hpi, two genes, hmm6 (122.43‐, 206.99‐ and 32.25‐fold, respectively) and PSCHS3 (175.00‐, 250.13‐ and 216.24‐fold, respectively), were strongly up‐regulated in leaves of the resistant genotype, highlighting that resistance against P. koolunga in field pea is governed by the early synthesis of pisatin. At 24 hpi, leaves infected by P. koolunga showed clear differences in expression of target genes. For example, the gene encoding a precursor of the defensin ‘disease resistance response protein 39’ was substantially down‐regulated in leaves of both the susceptible and the resistant genotypes inoculated with P. koolunga. This contrasts with other studies on another pea black spot pathogen, Didymella pinodes, where this same gene is strongly up‐regulated in leaves of resistant and susceptible genotypes. The current study provides the first understanding of defence‐related genes involved in the resistance against P. koolunga, opening novel avenues to engineer new field pea cultivars with improved leaf and stem black spot disease resistance as the basis for developing more effective and sustainable management strategies.     

Phenotyping Castanea hybrids for Phytophthora cinnamomi resistance

Castanea sativa is susceptible to Phytophthora spp., a serious root pathogen causing ink disease, while C. crenata and C. mollissima show resistance to infection. Interspecific controlled crosses were established for introgression of resistance genes from the resistant species into the susceptible C. sativa, and two mapping populations were created. Phytophthora cinnamomi resistance of each progeny was evaluated by root and excised shoot inoculation tests. The number of days of survival after root inoculation was the best discriminator of resistance to P. cinnamomi while the percentage of shoots with internal lesions was the symptom most associated with survival. The lesion progression rate in the excised shoot inoculation test was strongly and negatively correlated with survival in the root inoculation test. The excised shoot inoculation test appears to be a reliable approach for screening the resistance of chestnut genotypes to P. cinnamomi. Strong genetic correlations were obtained between survival and ink disease symptoms and among symptoms, indicating that common or linked genes might influence resistance to P. cinnamomi. The most resistant genotypes selected from this study will be tested for other commercial variables, such as ease of vegetative propagation and stock–scion compatibility.     

Variability in infection and reproduction of Meloidogyne javanica on tomato rootstocks with the Mi resistance gene

The response of 10 commercial or experimental tomato rootstocks with the Mi resistance gene to an initial inoculum of a Mi‐avirulent population of Meloidogyne javanica was determined in pot tests conducted in spring and summer. In a field test, the rootstocks were subjected to continuous exposure to high initial population densities (2050 ± 900 second‐stage juveniles (J2) per 250 cm³ soil) of the nematode. The presence of the Mi locus in the resistant rootstocks and cultivars was confirmed using the PCR co‐dominant markers REX‐1 and Mi23. Nematode infectivity (egg masses) and reproduction (eggs g^?1 root) were highly variable in the spring tests. Rootstocks PG76, Gladiator and MKT‐410 consistently responded as highly resistant, with nematode multiplication rate (Pf/Pi) < 1 and reproduction index (RI) < 10%, and they were as efficient as standard resistant tomato cultivars at nematode suppression. The relative resistance levels of rootstocks Brigeor, 42851, 43965, Big Power and He‐Man varied depending on the susceptible standard used for reference or the duration of the test. Rootstocks Beaufort and Maxifort were susceptible to M. javanica (Pf/Pi > 50 and RI > 50%). Rootstocks PG76 and He‐Man, and the resistant tomato cv. Caramba showed high levels of resistance in the test conducted in summer, whereas MKT‐410 and 42851 and the resistant tomato cv. Monika were moderately resistant. In the field, seven rootstocks showed high levels of resistance and one (He‐Man) showed an intermediate level, whereas Beaufort and Maxifort were susceptible.     

Implication of peroxynitrite in defence responses of potato to Phytophthora infestans

In this study peroxynitrite (ONOO^?) is proposed as an important player in defence responses during the interaction of potato (Solanum tuberosum) and the oomycete pathogen Phytophthora infestans. The potato–avr P. infestans model system exhibited a transient programme of boosted ONOO^? formation correlated in time with the burst of nitric oxide (NO) and superoxide during the first 6 h post‐inoculation (hpi). The early ONOO^? over‐accumulation was not accompanied by TPx gene expression. In contrast, the compatible interaction revealed a 24 h delay of ONOO^? formation; however, an enhanced level of NO and superoxide correlated with TPx up‐regulation was recorded within the earlier stages of pathogen infection. Peroxynitrite over‐accumulation in the susceptible potato coincided with an enhanced level of protein tyrosine nitration starting from 24 hpi. Surprisingly, the nitroproteome profile of the resistant potato did not show any visible difference after inoculation, apart from one band containing subtilisin‐like protease‐like proteins, which appeared 48 h after pathogen attack. An additional pharmacological approach showed that treatment of the susceptible genotype with ONOO^? followed by inoculation with P. infestans contributed to slowing down of the colonization of host tissues by the pathogen via a faster and stronger up‐regulation of the key defence markers, including the PR‐1 gene. Taken together, the results obtained indicate that a precise control of emitted NO and superoxide in cooperation with thioredoxin‐dependent redox sensors in sites of pathogen ingress could generate a sufficient threshold of ONOO^?, triggering defence responses.     

Production of avocado trees infected with Phytophthora cinnamomi under different management regimes

Avocado root rot is the most important disease of this fruit crop worldwide. This pathology may be caused by several biotic and abiotic agents, with the oomycete Phytophthora cinnamomi being the pathogen more frequently associated with poor phytosanitary conditions. There are disease control methods available that can reduce disease severity and allow plants to recover; however, they are not consistently and promptly applied. In addition, only chemical products are used by farmers as the preferred management method. This research aimed to evaluate different root rot management strategies in a commercial orchard. Data suggest that individual control methods are not as effective as when they are applied in combination, as in the T8 treatment (metalaxyl + mancozeb applied in drench; injection of potassium phosphite to each plant stem; potassium silicate applied in drench; addition of a layer of organic mulch and incorporation of 10 kg of composted substrate, both applied to the ground around the base of each tree). Using this strategy, the area under the disease progress curve for the avocado root rot was reduced by up to 68.6%, and the extra‐quality avocado fruit class increased by as much as 44% compared to the diseased control plants (T0) (P < 0.01). With the combined treatment T8, farm income showed a 9.5‐fold increase, probably due to an increase in the percentage of viable roots by up to 9.4‐fold, which would have improved nutrient and water uptake.     

A quantitative PCR assay for accurate <Emphasis Type="Italic">in planta</Emphasis> quantification of the necrotrophic pathogen <Emphasis Type="Italic">Phytophthora cinnamomi</Emphasis>

A reliable method for measuring disease progression is important when evaluating susceptibility in host—pathogen interactions. We describe a sensitive quantitative polymerase chain reaction (QPCR) assay that enables quantitative measurement of in planta DNA of the necrotrophic pathogen, Phytophthora cinnamomi, that avoids problems caused by variation in DNA extraction efficiency and degradation of host DNA during host tissue necrosis. Normalization of pathogen DNA to sample fresh weight or host DNA in samples with varying degrees of necrosis led to overestimation of pathogen biomass. Purified plasmid DNA, containing the pScFvB1 mouse gene, was added during DNA extraction and pathogen biomass was normalized based on plasmid DNA rather than host DNA or sample fresh weight. This method is robust and improves the accuracy of pathogen measurement in both resistant (non-host A. thaliana–P. cinnamomi) and susceptible (host Lupinus angustifolius–P. cinnamomi) interactions to allow accurate measurement of pathogen biomass even in the presence of substantial host cell necrosis.     

Effects of pre‐ and post‐treatment with ethephon on gum formation of peach gummosis caused by Lasiodiplodia theobromae

Peach gummosis, caused by Botryosphaeria spp. fungi, is the process of gum accumulation and exudation in plants. Ethephon (2‐chloroethylphosphonic acid) has profound effects on plants, including enhanced production of secondary metabolites and regulation of plant diseases. This study investigates the effects of application of ethephon before and after inoculation with Lasiodiplodia theobromae on gum formation. Gum formation was promoted by ethephon treatment prior to pathogen inoculation, but inhibited by ethephon applied after the pathogen. The inhibitory effect was counteracted by 1‐methylcyclopropane, which is an ethylene signal inhibitor. 1‐methylcyclopropane also promoted gum formation. Exposure of three isolates of Botryosphaeria to ethephon inhibited mycelial growth. Both treatment methods increased the sugar content at 12 and 24 h post‐inoculation (hpi). However, the sucrose, glucose and fructose contents were significantly higher in shoots with ethephon post‐treatment (application of ethephon after the pathogen inoculation) than those in shoots with ethephon pre‐treatment (application of ethephon prior to pathogen inoculation) at 48 and 72 hpi. The expression of two putative senescence‐related genes, SEN2 and SEN4, were significantly enhanced in pre‐ and post‐treated shoots with ethephon at 24, 48 and 72 hpi. Ethephon application also up‐regulated expression of the pathogenesis‐related protein PR4 while down‐regulating PR1a and PR10. The results show that ethephon has a dual function in regulating gum formation by affecting both the peach shoots and the pathogen.     

<Emphasis Type="Italic">Lupinus luteus</Emphasis>, a new host of <Emphasis Type="Italic">Phytophthora cinnamomi</Emphasis> in Spanish oak-rangeland ecosystems

Phytophthora cinnamomi is an aggressive pathogen on Lupinus luteus (yellow lupin), causing root rot, wilting and death of this crop, common in oak-rangeland ecosystems ('dehesas') in south-western Spain. The oomycete, the main cause of Quercus decline in the region, was isolated from roots of wilted lupins in the field. Artificial inoculations on four cultivars of L. luteus reproduced the symptoms of the disease, both in pre- and post-emergence stages, recovering the pathogen from necrotic roots. These results suggest the potential of yellow lupin as inoculum reservoir for the infection of Quercus roots. This is the first report of P. cinnamomi as root pathogen of L. luteus.     

Colonization of lettuce cultivars and rotation crops by Fusarium oxysporum f. sp. lactucae,the cause of fusarium wilt of lettuce

The severity of fusarium wilt is affected by inoculum density in soil, which is expected to decline during intervals when a non‐susceptible crop is grown. However, the anticipated benefits of crop rotation may not be realized if the pathogen can colonize and produce inoculum on a resistant cultivar or rotation crop. The present study documented colonization of roots of broccoli, cauliflower and spinach by Fusarium oxysporum f. sp. lactucae, the cause of fusarium wilt of lettuce. The frequency of infection was significantly lower on all three rotation crops than on a susceptible lettuce cultivar, and the pathogen was restricted to the cortex of roots of broccoli. However, F. oxysporum f. sp. lactucae was isolated from the root vascular stele of 7·4% of cauliflower plants and 50% of spinach plants that were sampled, indicating a greater potential for colonization and production of inoculum on these crops. The pathogen was also recovered from the root vascular stele of five fusarium wilt‐resistant lettuce cultivars. Thus, disease‐resistant plants may support growth of the pathogen and thereby contribute to an increase in soil inoculum density. Cultivars that were indistinguishable based on above‐ground symptoms, differed significantly in the extent to which they were colonized by F. oxysporum f. sp. lactucae. Less extensively colonized cultivars may prove to be superior sources of resistance to fusarium wilt for use in breeding programmes.     

Identification and expression pattern of a glutathione S‐transferase in Echinochloa crus‐galli

Plant glutathione S‐transferase (GST) forms a major part of the herbicide detoxification enzyme network in plants. A GST cDNA was isolated from Echinochloa crus‐galli and characterised. The gene, designated EcGST1 (E. crus‐galli GeneBank no: JX518596 ), has a 684 bp open reading frame predicted to encode a 25 kD protein. Sequence alignment showed that EcGST1 is a GST homologue. Its expression in response to quinclorac treatment was monitored in seedlings (leaves and roots) and adult plants (leaves, roots, stems and seeds) of quinclorac‐resistant (R) and susceptible (S) biotypes of E. crus‐galli. EcGST1 expression was 1.5–3 times greater in the R plants than in the S plants. However, after exposure to quinclorac, the difference in the expression levels of EcGST1 in R plants, compared with S plants, increased to a ratio of 6–10. Enhanced EcGST1 levels should enable greater quinclorac detoxification following quinclorac stimulation in R plants. GST‐based metabolism may be partially responsible for resistance to quinclorac in E. crus‐galli. The results suggest a new resistance mechanism for this R biotype in Chinese rice fields.     

Investigation of rice blast development in susceptible and resistant rice cultivars using a gfp‐expressing Magnaporthe oryzae isolate

In this study, an isolate of Magnaporthe oryzae expressing the green fluorescent protein gene (gfp) was used to monitor early events in the interaction of M. oryzae with resistant rice cultivars harbouring a blast resistance (R) gene. In the resistant cultivars Saber and TeQing (Pib gene), M. oryzae spores germinated normally on the leaf surface but produced morphologically abnormal germ tubes. Germling growth and development were markedly and adversely affected in leaves of these resistant cultivars. Penetration of host cells was never seen, supporting the idea that disruption of germling development on the leaf surface might be one of the resistance mechanisms associated with Pib function. Thus, this particular R gene appeared to function in the absence of host penetration by the fungal pathogen. Confocal laser scanning microscopy of M. oryzae‐infected susceptible rice cultivars showed the dimorphic growth pattern that is typically observed during the biotrophic and necrotrophic stages of leaf colonization in susceptible cultivars. The suitability of the gfp‐expressing M. oryzae isolate for further research on R‐gene function and identification of resistant genotypes in rice germplasm collections is discussed.     

Involvement of a vascular hypersensitive response in quantitative resistance to Ralstonia solanacearum on tomato rootstock cultivar LS‐89

Ralstonia solanacearum causes bacterial wilt disease in Solanaceae spp. Expression of the Phytophthora inhibitor protease 1 (PIP1) gene, which encodes a papain‐like extracellular cysteine protease, is induced in R. solanacearum‐inoculated stem tissues of quantitatively resistant tomato cultivar LS‐89, but not in susceptible cultivar Ponderosa. Phytophthora inhibitor protease 1 is closely related to Rcr3, which is required for the Cf‐2‐mediated hypersensitive response (HR) to the leaf mould fungus Cladosporium fulvum and manifestation of HR cell death. However, up‐regulation of PIP1 in R. solanacearum‐inoculated LS‐89 stems was not accompanied by visible HR cell death. Nevertheless, upon electron microscopic examination of inoculated stem tissues of resistant cultivar LS‐89, several aggregated materials associated with HR cell death were observed in xylem parenchyma and pith cells surrounding xylem vessels. In addition, the accumulation of electron‐dense substances was observed within the xylem vessel lumen of inoculated stems. Moreover, when the leaves of LS‐89 or Ponderosa were infiltrated with 10⁶ cells mL^?1 R. solanacearum, cell death appeared in LS‐89 at 18 and 24 h after infiltration. The proliferation of bacteria in the infiltrated leaf tissues of LS‐89 was suppressed to approximately 10–30% of that in Ponderosa, and expression of the defence‐related gene PR‐2 and HR marker gene hsr203J was induced in the infiltrated tissues. These results indicated that the response of LS‐89 is a true HR, and induction of vascular HR in xylem parenchyma and pith cells surrounding xylem vessels seems to be associated with quantitative resistance of LS‐89 to R. solanacearum.     

Characterization of resistance against the olive‐defoliating Verticillium dahliae pathotype in selected clones of wild olive

Verticillium wilt of olive is best managed by resistant cultivars, but those currently available show incomplete resistance to the defoliating (D) Verticillium dahliae pathotype. Moreover, these cultivars do not satisfy consumers' demand for high yields and oil quality. Highly resistant rootstocks would be of paramount importance for production of agronomically adapted and commercially desirable olive cultivars in D V. dahliae‐infested soils. In this work, resistance to D V. dahliae in wild olive clones Ac‐13, Ac‐18, OutVert and StopVert was assessed by quantifying the fungal DNA along the stem using a highly sensitive real‐time quantitative polymerase chain reaction (qPCR) protocol and a stem colonization index (SCI) based on isolation of V. dahliae following artificial inoculations under conditions highly conducive for verticillium wilt. Ac‐13, Ac‐18, OutVert and StopVert showed a symptomless reaction to D V. dahliae. The mean amount of D V. dahliaeDNA quantified in stems of the four clones ranged from 3.64 to 28.89 pg/100 ng olive DNA, which was 249 to 1537 times lower than that in susceptible Picual olive. The reduction in the quantitative stem colonization of wild olive clones by D V. dahliae was also indicated by a sharp decrease in the SCI. Overall, there was a pattern of decreasing SCI in acropetal progression along the plant axis, as well as correlation between positive reisolation and quantification of pathogen DNA. The results of this research show that wild olive clones Ac‐13, Ac‐18, OutVert and StopVert have a valuable potential as rootstocks for the management of verticillium wilt in olive.     

Response of selected South Australian native plant species to Phytophthora cinnamomi

Thirty‐seven South Australian native plant species from 11 families, including 15 threatened species in the state (of which six are listed as threatened under the federal Environment Protection and Biodiversity Conservation Act 1999) were assessed for response to infection by Phytophthora cinnamomi. Seedlings, 3–6 months old and grown in a greenhouse, were inoculated by placing infested pine wood plugs in the potting mix, maintained in moist conditions and assessed for mortality and disease symptoms for between 3 and 10 months. Thirty species were found to be susceptible, of which nine were highly susceptible, 15 moderately susceptible and six slightly susceptible. Three species were found to be resistant and results for four species were inconclusive. Six of the 15 threatened, rare or locally endangered species tested (Eucalyptus viminalis var. viminalis, Correa aemula, C. calycina, Olearia pannosa ssp. pannosa, Pomaderris halmaturina ssp. halmaturina and Prostanthera eurybioides) were moderately susceptible, while two (Allocasuarina robusta and Pultenaea graveolens) were highly susceptible. Significant populations of at least five of the threatened species susceptible to the disease are located close to confirmed or suspected Phytophthora‐infested areas or growing in areas conducive for P. cinnamomi. An effective management strategy is therefore required to avoid extinction of such species due to infection by the phytophthora dieback pathogen.     

Screening tomato and cucurbit rootstocks for resistance to Verticillium dahliae*

Seedlings of tomato (19) and cucurbits (33), previously selected as rootstocks for commercial cultivars, were evaluated for their resistance to verticillium wilt under glasshouse conditions. Disease scoring was based on foliar symptoms, but in plants with no visual or ambiguous symptoms the presence of the pathogen was confirmed by isolations or PCR using specific DNA primers. Five of the cucurbit rootstocks showed resistance to infection, 11 were moderately infected, 11 were susceptible, while the remaining six were very susceptible. In general, Cucurbita pepo (courgette, pumpkin) and Lagenaria siceraria tolerated infection, Cucumis melo rootstocks were susceptible and Citrullus vulgaris was the most susceptible. When inoculated with race 1 of the pathogen, one of the tomato rootstocks was very resistant, three moderately resistant, nine tolerant, ten susceptible and one very susceptible. Pathogenicity tests with race 2 showed that none of the rootstocks exhibited high tolerance, but that there was some variation in susceptibility. Research is under way to evaluate the reaction of commercial cultivars to verticillium wilt when grafted onto resistant rootstocks.