Resistance to Sclerotinia sclerotiorum in wild Brassica species and the importance of Sclerotinia subarctica as a Brassica pathogen

Brassica crops are of global importance, with oilseed rape (Brassica napus) accounting for 13% of edible oil production. All Brassica species are susceptible to sclerotinia stem rot caused by Sclerotinia sclerotiorum, a generalist fungal pathogen causing disease in over 400 plant species. Generally, sources of plant resistance result in partial control of the pathogen although some studies have identified wild Brassica species that are highly resistant. The related pathogen S. subarctica has also been reported on Brassica but its aggressiveness in relation to S. sclerotiorum is unknown. In this study, detached leaf and petiole assays were used to identify new sources of resistance to S. sclerotiorum within a wild Brassica ‘C genome’ diversity set. High‐level resistance was observed in B. incana and B. cretica in petiole assays, whilst wild B. oleracea and B. incana lines were the most resistant in leaf assays. A B. bourgeai line showed both partial petiole and leaf resistance. Although there was no correlation between the two assays, resistance in the detached petiole assay was correlated with stem resistance in mature plants. When tested on commercial cultivars of B. napus, B. oleracea and B. rapa, selected isolates of S. subarctica exhibited aggressiveness comparable to S. sclerotiorum indicating it can be a significant pathogen of Brassica. This is the first study to identify B. cretica as a source of resistance to S. sclerotiorum and to report resistance in other wild Brassica species to a UK isolate, hence providing resources for breeding of resistant cultivars suitable for Europe.     

Non‐pathogenic rhizosphere bacteria belonging to the genera Rhizorhapis and Sphingobium provide specific control of lettuce corky root disease caused by species of the same bacterial genera

Lettuce corky root (CR) is caused by bacteria in the genera Rhizorhapis, Sphingobium, Sphingopyxis and Rhizorhabdus of the family Sphingomonadaceae. Members of this family are common rhizosphere bacteria, some pathogenic to lettuce. Sixty‐eight non‐pathogenic isolates of bacteria obtained from lettuce roots were tested for control of CR caused by Rhizorhapis suberifaciens CA1^T and FL1, and Sphingobium mellinum WI4^T. In two initial screenings, 10 isolates significantly reduced CR induced by one or more pathogenic strains on lettuce seedlings in vermiculite, while seven non‐pathogenic isolates provided significant CR control in natural or sterilized field soil. Rhizorhapis suberifaciens FL11 was effective at controlling all pathogenic strains, but most effective against R. suberifaciens CA1^T. The other selected isolates controlled only pathogenic strains belonging to their own genus. In a greenhouse experiment, a soil drench with selected biocontrol agents (R. suberifaciens FL11, Sphingomonas sp. NY3 and S. mellinum CA16) controlled CR better than seed treatments or application of alginate pellets. In microplots infested with R. suberifaciens CA1^T, seed treatment with R. suberifaciens FL11 provided complete control and a soil drench with FL11 significantly reduced the disease. Pathogenicity tests with FL11 on 23 plant species in 10 families resulted in slight yellowing on roots of lettuce and close relatives; similar yellowing appeared on some roots of non‐inoculated lettuce plants. This research showed that biocontrol agents can be genus‐specific. Only one isolate, FL11, provided more general control of various pathogenic strains causing CR even in field soil in pots and microplots.     

Incidence and distribution of ‘Candidatus Phytoplasma prunorum’ and its vector Cacopsylla pruni in Spain: an approach to the epidemiology of the disease and the role of wild Prunus

‘Candidatus Phytoplasma prunorum’ is the causal agent of the European stone fruit yellows (ESFY) disease. This phytoplasma affects wild and cultivated species of Prunus to different degrees, depending on their susceptibility. ‘Candidatus Phytoplasma prunorum’ is present in the four regions of Spain surveyed in this study (Aragon, Catalonia, Extremadura and Valencia) with a variable incidence. Results showed that ‘Ca. Phytoplasma prunorum’ was detected in all of the cultivated Prunus species studied, except P. avium and P. dulcis, and was widespread in Spain. The most affected species was P. salicina, with symptoms including early bud break and blooming, leaf curling and yellowing, collapse, and a major decrease in production. In some plots in the Baix Llobregat area of Barcelona province (Catalonia), the incidence of ESFY on P. salicina was as high as 80%. The insect vector, Cacopsylla pruni, was present in all four of the regions studied, with the highest captures in yellow sticky traps in Catalonia on P. mahaleb and in Extremadura in peach orchards. In Baix Llobregat, large populations of C. pruni were present on infected P. mahaleb bushes, and with high infection rates. This was a key factor in the local pathogenic cycle that caused a major ESFY outbreak in the nearby P. salicina orchards. In the Ebro valley (Lleida and Aragon) and Valencia, the surveys showed very low incidences of the disease and low C. pruni populations.     

Observed changes in soyabean growth and seed yield from Abutilon theophrasti competition as a function of carbon dioxide concentration

Soyabean (Glycine max) was grown at ambient and projected levels of atmospheric carbon dioxide (+250 μmol mol^?1 above ambient) over two field seasons with and without the presence of a weed, Abutilon theophrasti, to quantify the potential effect of rising atmospheric carbon dioxide concentration on weed–crop interactions and potential yield loss in soyabean. Under weed‐free conditions, elevated CO₂ resulted in stimulations in soyabean seed yield and associated components, including pod number. At an approximate density of 6 plants m^?2, A. theophrasti competition resulted in a significant reduction (?40%) in soyabean seed yield. Although differences in seed yield reduction by A. theophrasti were observed as a function of year, the relative decrease in seed yield with A. theophrasti biomass did not differ in response to CO₂. Although careful weed management will be necessary if CO₂‐induced increases in seed yield for soyabean are to be achieved, these data suggest that soyabean seed yield may be more resilient in competition with A. theophrasti as a function of rising atmospheric levels of carbon dioxide.     

The roles of light and pericarp on seed dormancy and germination in feral Raphanus sativus (Brassicaceae)

The timing of seed germination may determine the success of a weed species in an agroecosystem, and its expression is modulated by environmental conditions, but also by seed physiology and anatomy. The aims of this study were to investigate the roles of light, pericarp, dry storage and cold stratification on seed dormancy and germination in feral radish, a troublesome agricultural weed in temperate zones of the Americas that reduces crop yields. To this end, we used isolated intact pods and extracted seeds to test germination over time under contrasting temperature, light and storage conditions. Here, we showed that fresh seeds were non‐dormant, but that light and the presence of the pericarp reduced germination, especially under low temperatures. The pericarp reduced the final water content absorbed by seeds inside pods and decreased absorption/dehydration rates. The pericarp showed several small lignified cell layers in the endocarp, and x‐ray images displayed the lack of space between the partially embedded seed and the endocarp. Dry storage and cold stratification were ineffective in breaking the dormancy imposed by the pericarp. The apparent requirement for darkness and the mechanical restriction of the pericarp may have the potential to induce dormancy, spreading the timing of seed germination over a more extended period and hindering the control of feral radish.     

Contribution of both lignin content and sinapyl monomer to disease resistance in tobacco

As a major component of the cell wall, lignin has been suggested to play an important role in the plant defence response to various pathogens. However, how lignin is involved in plant pathogen interaction is still unclear. Here, a series of transgenic tobacco lines were cultivated with a range of differences in lignin content and composition. Evaluation of pathogen resistance in these plants indicated that lower total lignin content aggravated the severity of tobacco black shank and bacterial wilt diseases, while increased sinapyl lignin (S) alleviated the disease symptoms. The regression analysis indicated both lignin content and S lignin were positively correlated with disease resistance. These two factors had additive effects, exhibiting stronger correlation with disease resistance when they were combined. Neither guaiacyl lignin (G) nor S/G ratio showed close correlation with disease resistance. The expression of pathogenesis‐related protein genes PR2 and PR3 was induced after pathogen inoculation. However, the up‐regulation of PR2 and PR3 was not associated with a disease resistance‐induced increase in lignin content. These data collectively suggest that both total lignin content and S lignin are main factors that contribute to the basic defence response in tobacco.     

Rhizoctonia solani: taxonomy,population biology and management of rhizoctonia seedling disease of soybean

Rhizoctonia solani, the most important species within the genus Rhizoctonia, is a soilborne plant pathogen with considerable diversity in cultural morphology, host range and aggressiveness. Despite its history as a destructive pathogen of economically important crops worldwide, our understanding of its taxonomic relationship with other Rhizoctonia‐like fungi, incompatibility systems, and population biology is rather limited. Among the host of diseases it has been associated with, seedling diseases inflicted on soybean are of significant importance, especially in the soybean growing regions of North America. Due to the dearth of resistant soybean genotypes, as well as the paucity of information on the mechanisms of host–pathogen interactions and other molecular aspects of pathogenicity, effective management options have mostly relied upon a combination of cultural and chemical control options. The first section of this review summarizes what is currently known about the taxonomy and systematics, population biology and molecular genetics of R. solani. The second section provides an overview of the pathology and management of rhizoctonia root and hypocotyl rot of soybean, a seedling disease of importance in North America.     

Evidence for Lettuce big‐vein associated virus as the causal agent of a syndrome of necrotic rings and spots in lettuce

Lettuce big‐vein associated virus (LBVaV, genus Varicosavirus) was shown to be responsible for characteristic necrotic symptoms observed in combination with big‐vein symptoms in lettuce breeding lines when tested for their susceptibility to lettuce big‐vein disease (BVD) using viruliferous Olpidium virulentus spores in a nutrient film technique (NFT) system. Lettuce plants showing BVD are generally infected by two viruses: Mirafiori lettuce big‐vein virus (MiLBVV, genus Ophiovirus) and LBVaV. New mechanical inoculation methods were developed to separate the two viruses from each other and to transfer both viruses to indicator plants and lettuce. After mechanical inoculation onto lettuce plants MiLBVV induced vein‐band chlorosis, which is the characteristic symptom of BVD. LBVaV caused a syndrome of necrotic spots and rings which was also observed earlier in lettuce plants inoculated in the NFT system, resembling symptoms described for lettuce ring necrosis disease (RND). This observation is in contrast with the idea that LBVaV only causes latent infections in lettuce. De novo next‐generation sequencing demonstrated that LBVaV was the only pathogen present in a mechanically inoculated lettuce plant with symptoms, providing evidence that LBVaV was the causal agent of the observed necrotic syndrome and thus fulfilling Koch’s postulates for this virus. The necrotic syndrome caused by LBVaV in lettuce is referred to as LBVaV‐associated necrosis (LAN).     

The hotter the weather,the greater the infestation of Portulaca oleracea: opportunistic life‐history traits in a serious weed

Portulaca oleracea, an r‐strategist, is one of the world's most troublesome weeds. During hot seasons, P. oleracea frequently becomes monodominant in choy sum (Brassica parachinensis) fields in Guangzhou city, southern China. Here, we studied the seasonal dynamics of P. oleracea's germinable soil seedbank, population density and above‐ground biomass in choy sum fields that had been cultivated continuously for several years. Using P. oleracea seeds collected from these fields, we tested seed dormancy, survival and germination, seedling growth and generation time. Portulaca oleracea occurred at high levels during the hot season, but its occurrence was low, and the germinable soil seedbank was much greater during the cold season. The weed's opportunistic characteristics allowed it to avoid freezing and to proliferate during optimal conditions. Portulaca oleracea's generation time was very flexible, as short as 31 days during the hot season, but longer than 100 days during the colder season. Seed dormancy tended to be shorter when the seeds were stored at a higher temperature. At a temperature of 35°C, both seed germination and seedling growth showed advantages over those of choy sum. Storage for one year at a temperature of ?20°C or burial in a paddy field did not significantly reduce P. oleracea seed germination. Nevertheless, seed storage at a temperature of 15°C and soil coverage of 0.5 cm on top of the seeds significantly constrained seed germination. ‘Stale seedbed’ and/or coverage of the surface with soil are recommended during the hot season. However, rotation of rice and upland crops is not an efficient method for managing Portulaca oleracea infestation.     

The threat of root‐knot nematodes (Meloidogyne spp.) in Africa: a review

Meloidogyne species pose a significant threat to crop production in Africa due to the losses they cause in a wide range of agricultural crops. The direct and indirect damage caused by various Meloidogyne species results in delayed maturity, toppling, reduced yields and quality of crop produce, high costs of production and therefore loss of income. In addition, emergence of resistance‐breaking Meloidogyne species has partly rendered various pest management programmes already in place ineffective, therefore putting food security of the continent at risk. It is likely that more losses may be experienced in the future due to the on‐going withdrawal of nematicides. To adequately address the threat of Meloidogyne species in Africa, an accurate assessment and understanding of the species present, genetic diversity, population structure, parasitism mechanisms and how each of these factors contribute to the overall threat posed by Meloidogyne species is important. Thus, the ability to accurately characterize and identify Meloidogyne species is crucial if the threat of Meloidogyne species to crop production in Africa is to be effectively tackled. This review discusses the use of traditional versus molecular‐based identification methods of Meloidogyne species and how accurate identification using a polyphasic approach can negate the eminent threat of root knot nematodes in crop production. The potential threat to Africa posed by highly damaging and resistance‐breaking populations of ‘emerging’ Meloidogyne species is also examined.     

Effects of temperature and light on seed germination of ephemeral plants in the Gurbantunggut Desert,China: implications for vegetation restoration

Seed germination is a key transitional stage in plant life cycle and is strongly regulated by temperature and light. Therefore, research on the effects of temperature and light on seed germination is extremely meaningful for vegetation restoration, especially in desert ecosystems. Seeds of 28 ephemeral plants collected from the Gurbantunggut Desert of China were incubated at different temperatures (5°C/1°C, 15°C/5°C, 20°C/5°C, 25°C/10°C and 30°C/15°C) in 12-h light/12-h darkness or continuous darkness regimes, and the responses of seed germination to temperature and light and the germination speed were studied in 2016. Results showed that seed germination percentage of the 28 ephemeral plants significantly differed to temperature and light. We classified the studied plants as the following groups based on their responses to temperature: 1 low temperature responsed plants, 12 moderate temperature responsed plants, 7 high temperature responsed plants, 4 non-responsed plants and 5 plants of no germination. It should be noted that Corispermum lehmannianum Bunge is sensitive to both moderate and high temperatures. There were 4 groups of plant in response to light, i.e., 7 light responsed plants, 10 dark responsed plants, 6 light non-responsed plants and 5 plants of no germination. Based on seed germination speed of the 28 ephemeral plants, we divided them into 4 patterns of germination, i.e., very rapid, moderately rapid, moderate and slow. Combining variations of temperature, precipitation and sand dune types in the study area, we suggested that very rapid and moderately rapid germinated plants could be used to moving sand dunes in early spring during vegetation restoration, moderate germinated plants could be used to semi-fixed sand dunes in late autumn, and slow germinated plants could be used to sand plain in summer. Thus, seedling establishment and vegetation restoration would be improved by considering seed germination characteristics of these ephemeral plants in the Gurbantunggut Desert, China.     

Sorghum (Sorghum bicolor) as a bean intercrop or rotation crop contributes to the survival of bean root rot pathogens and perpetuation of bean root rots

Root rots (RR) are the main cause of declining bean (Phaseolus vulgaris) production in southwestern Uganda. Here, beans are mainly intercropped/rotated with maize (Zea mays), sorghum (Sorghum bicolor), sweet potato (Ipomoea batatas), potatoes (Solanum tuberosum) and garden peas (Pisum sativum). These crops also suffer from RR and bean RR pathogens have been isolated from some of these crops. This study aimed to determine the extent of RR on maize, sorghum and peas, and their potential to contribute to the survival of bean RR pathogens. Therefore, experiments were carried out in bean RR‐infested farmers’ fields as well as soils inoculated with bean RR pathogens (Pythium spp. and Fusarium spp.) under screen house conditions and a susceptible bean cultivar served as a control. High RR incidence/severity scores were recorded in beans and sorghum in both farmers’ fields and screen house experiments. The high field RR incidence/severity in sorghum correlated with the screen house scores. This study shows that RR is also a problem to other crops, especially sorghum, warranting attention. The findings also imply that sorghum plays a potential role as an alternate host to bean RR pathogens, increasing inoculum density of bean RR pathogens and potentially negatively impacting the bean RR problem. Intercropping or rotating beans with sorghum in this region is not recommended. However, maize was RR‐resistant and therefore appropriate as an intercrop/rotational crop to beans in the system. A holistic rather than commodity approach is recommended for managing RR in this cropping system.     

The reliability of leaf bioassays for predicting disease resistance on fruit: a case study on grapevine resistance to downy and powdery mildew

This study was designed to assess the reliability of grapevine leaf bioassays for predicting disease resistance on fruit in the field. The efficacy of various grapevine quantitative trait loci (QTLs) for conferring resistance to downy and powdery mildew was evaluated in bioassays and in a 2‐year field experiment for downy mildew. The resistance genes studied were inherited from Muscadinia rotundifolia (Rpv1 and Run1) and from American Vitis species through cv. Regent (QTLRgP and QTLRgD). In bioassays, genotypes carrying Run1 blocked powdery mildew development at early stages. Genotypes combining Run1 with QTLRgP displayed no greater level of resistance. For downy mildew, genotypes carrying Rpv1 and/or QTLRgD were more resistant than the susceptible cv. Merlot, and showed a high level of leaf resistance in the field (<10% severity). Disease levels on bunches were much higher than those on leaves, with a high variability between Rpv1 genotypes (1–48%). A Bayesian decision theory framework predicted that an OIV‐452 threshold of 5 in leaf bioassays allowed accurate selection of grapevine genotypes (P = 0·83) with satisfactory disease severity on bunches. Therefore, this study validates that the use of early bioassays on leaves, as currently performed by grapevine breeders, ensures a satisfactory level of resistance to downy mildew of bunches in the field.     

Molecular characterization of Ralstonia solanacearum strains from Ethiopia and tracing potential source of bacterial wilt disease outbreak in seed potatoes

Bacterial wilt, caused by Ralstonia solanacearum, is emerging as a major threat to potato production in Ethiopia, reaching epidemic proportions in the Chencha district recently, with a prevalence of 97% of potato fields in 2015. The recent disease outbreak in the district coincided with a significant introduction of seed potatoes. This research was therefore initiated to genetically characterize the pathogen so as to trace its source, identify its relationship with outbreaks in the rest of the country, and make intervention recommendations. Ralstonia solanacearum isolates were sampled both from seed and ware potato fields in Chencha and from seed potato fields in production regions suspected of being potential sources of the pathogen. Multiplex PCR and phylogenetic analysis of partial endoglucanase gene sequences identified all of the isolates as phylotype IIB sequevar 1. VNTR sequence analysis distinguished 11 different haplotypes, nine of which were unique to the Chencha district. However, one of the haplotypes was common to all seed potato producer regions of Ethiopia except for the Shashemene area. The unique and diverse VNTR haplotypes of the pathogen in Chencha indicates that it is well established in the district. When a geographical map of the VNTR haplotypes was superimposed with the main cross‐regional seed potato distribution pattern of the country, it became evident that the pathogen was being disseminated via latently infected seed from the Holeta‐Jeldu area in the Central Highlands of Ethiopia. Identification of largely uninfected highland districts and multiplication of high‐grade seed potato exclusively in those districts should be given priority.     

Testing and modelling the potential of three diploid plants in Poaceae as a new pathosystem to investigate the interactions between cereal hosts and cereal cyst nematode (Heterodera avenae)

Cereal cyst nematode (CCN), Heterodera avenae, is one of the most important pathogens of wheat worldwide, and causes significant yield losses. Research on CCN–wheat interactions is hampered by the lack of an effective model pathosystem. This study investigated the potential of the model cereal Brachypodium distachyon (Bd21‐3) and diploid wheat 2A (G1812) and 2D (AL8/78) as model hosts for CCN. Nematode infection analysis showed that although some CCN penetrated Bd21‐3 roots, these nematodes failed to develop to the later developmental stages or form cysts, indicating B. distachyon is not a host for CCN. A strong burst of reactive oxygen species (ROS) within Bd21‐3 roots infected with CCN was induced 3 days after infection and the expression of seven ROS‐producing genes was significantly increased. In contrast, CCN completed its life cycle in both diploid wheat 2A and 2D, and formed normal syncytia in these hosts. Although CCN developmental processes within both diploid wheat 2A and 2D were very similar to those in the susceptible control, the number of cysts formed on diploid wheat 2D was less than those formed on diploid wheat 2A and the susceptible control, indicating that diploid wheat 2A was a more suitable host for CCN than 2D. This is the first report of a potential new pathosystem for CCN–host interactions using diploid wheat.     

Development and validation of a set of standard area diagrams to aid in estimation of spot blotch severity on wheat leaves

This study aimed to develop and validate a standard area diagram (SAD) set to quantify the severity of spot blotch, caused by Bipolaris sorokiniana, on wheat leaves. The proposed SAD set includes images of leaves with 11 distinct disease severities (0·1, 1, 5, 10, 20, 30, 40, 50, 60, 70 and 83%). The SAD set was validated by 12 raters without experience in evaluating plant disease. Lin's concordance correlation analysis of estimated versus actual disease severity (based on image analysis) showed that precision and accuracy improved for all raters using the SAD set in contrast to assessments made without it. The SAD set improved accuracy (coefficient of bias, C_b = 0·88 and 0·99, without and with the SAD set, respectively) and agreement (Lin's concordance correlation coefficient, ρ_c = 0·81 and 0·96 without and with the SAD set, respectively) of the estimates of severity. The severity estimates were also more reliable when using the SAD set (coefficient of determination, R² = 0·76 unaided and R² = 0·92 with the SAD set, and intra‐class correlation ρ = 0·79 without the SAD set and ρ = 0·95 using the SAD set). The SAD set proposed in this study will improve the accuracy and reliability of estimates of spot blotch severity on wheat leaves.     

Analysis of resistance to witches’ broom disease (Moniliophthora perniciosa) in flower cushions of Theobroma cacao in a segregating population

This study evaluates resistance to witches’ broom disease in flower cushions of Theobroma cacao under field conditions. The aim was to determine optimal inoculation methods to evaluate the disease incidence using flower cushions in the field. A segregating mapping population of 580 trees (cultivar TSH 1188 × CCN 51) was analysed under two field conditions: high and low inoculum levels (in different years), corresponding respectively to trees with or without dried witches’ brooms hanging on the trees and producing basidiocarps. The number of newly formed cushion brooms in each tree was counted by the conventional method, and also the healthy and infected flower cushions in three 30 cm‐long regions along the trunk and the two main branches. The field inoculation methods discriminated between genotypes, with a 26% increase in disease incidence by Moniliophthora perniciosa at high inoculum. Two different segregation patterns were also observed: 27:27:9:1 under low, and 27:9:9:9:3:3:3:1 under high inoculum potential. It was also determined that at least 20 flower cushions were needed to accurately determine the percentage of infection. These methodologies allowed identification of the extreme phenotypes in this mapping population, and can therefore facilitate the detection of sources of resistance to witches’ broom disease.     

Biological control of Pythium,Rhizoctonia and Sclerotinia in lettuce: association of the plant protective activity of the bacterium Paenibacillus alvei K165 with the induction of systemic resistance

The soilborne fungi Sclerotinia sclerotiorum, Rhizoctonia solani and the oomycete Pythium ultimum are among the most destructive pathogens for lettuce production. The application of the biocontrol agent Paenibacillus alvei K165 to the transplant soil plug of lettuce resulted in reduced S. sclerotiorum, R. solani and P. ultimum foliar symptoms and incidence compared to untreated controls, despite the suppressive effect of the pathogens on the rhizosphere population of K165. In vitro, K165 inhibited the growth of S. sclerotiorum and R. solani but not P. ultimum. Furthermore, the expression of the pathogenesis‐related (PR) gene PR1, a marker gene of salicylic acid (SA)‐dependent plant defence, and of the Lipoxygenase (LOX) and Ethylene response factor 1 (ERF1) genes, markers of ethylene/jasmonate (ET/JA)‐dependent plant defence was recorded. K165‐treated plants challenged with P. ultimum showed up‐regulation of PR1, whereas challenge with R. solani resulted in up‐regulation of LOX and ERF1, and challenge with S. sclerotiorum resulted in up‐regulation of PR1, LOX and ERF1. This suggests that K165 triggers the SA‐ and the ET/JA‐mediated induced systemic resistance against P. ultimum and R. solani, respectively, while the simultaneous activation of the SA and ET/JA signalling pathways is proposed for S. sclerotiorum.     

Elucidation of the infection process of Ustilaginoidea virens (teleomorph: Villosiclava virens) in rice spikelets

This study reports an efficient inoculation protocol that allowed cytological analysis of the infection process of the rice false smut pathogen Ustilaginoidea virens. Examination of serial semithin and ultrathin sections of infected spikelets showed that the primary infection sites for the pathogen were the upper parts of the three stamen filaments located between the ovary and the lodicules. The stigma and lodicules were also occasionally infected to a limited extent. The pathogen infected the filaments intercellularly and extended intercellularly along the filament base. The host cells were degraded gradually. The pathogen did not penetrate host cell walls directly and did not form haustoria. In the balls the ovary remained alive and was never infected. This suggests that the pathogen is a biotrophic parasite that grows intercellularly in vivo.