Co-evolution of potato cyst nematodes and their hosts: implications for pathotypes and resistance1

The co-evolutionary process is believed to have resulted, through interaction of wild populations of potato cyst nematodes and their hosts in geological time, in the resistant hosts now utilized in plant breeding programmes and in nematode populations with genes for resistance-breaking or virulence. It is argued that all such interactions between highly adapted, truly parasitic plant nematodes and their hosts are likely to be governed by gene-for-gene interrelationships. Practical implications of this hypothesis are that only pathotypes (resistance-breaking races) defined against identified resistance genes are scientifically sound and of practical value; that, in the case of potato cyst nematodes, other pathotypes (Ro2, Ro3, Ro5 and Pa2 and Pa3) should be abandoned; and that oligogenically based resistance to potato cyst nematodes, especially important in providing resistance to Globodera pallida, is non-durable. Working definitions of the terms ‘pathotype’, ‘host-race’ and ‘virulence’ are provided.     

Breeding for resistance: breeding for Plum pox virus resistant apricots (Prunus armeniaca L.) in Spain

Plum pox virus , Dideron type (PPV-D), was first detected in Spain in 1984. Since then, it has spread among Japanese plum trees and apricot trees has been extremely rapid in the main producing areas. In Spain, breeding for resistance was the only efficient method for controlling the disease on apricot. Two breeding programmes are currently producing new hybrids resistant to the disease. The main problem encountered by both programmes is the difficult procedure needed for screening the trait that delays the programmes. Nevertheless, more than 8000 seedlings have been produced, two new varieties have been released and several advanced selections are under study. The procedure for screening sharka resistance has varied by techniques and cultivars used which has resulted in different hypotheses about inheritance of the trait. Additionally, several studies on mapping and molecular markers in progress could provide markers for molecular assisted selection that can increase the breeding efficiency.     

Costs and trade-offs associated with induced resistance

Plants resist attack by pathogens and herbivorous insects through constitutive and inducible defences. Based on differences in signalling pathways and spectra of effectiveness, different types of induced resistance have been defined. Systemic acquired resistance (SAR) occurs in distal plant parts following localized infection by a necrotizing pathogen. It is controlled by a signalling pathway that depends upon the accumulation of salicylic acid (SA) and the regulatory protein NPR1. In contrast, induced systemic resistance (ISR) is induced by selected strains of non-pathogenic plant growth promoting bacteria (PGPR). ISR functions independently of SA, but requires NPR1 and is regulated by jasmonic acid (JA) and ethylene (ET). It is generally believed that induced resistance evolved to save energy under pathogen or insect-free conditions, although costs still arise when defences are activated following attack. Costs can arise from the allocation of resources to defence and away from plant growth and development, and there are also ecological costs which result from trade-offs between induced resistance and the plant's interaction with beneficial organisms e.g. mycorrhizal fungi. To date, few studies have examined the costs and trade-offs associated with induced resistance to pathogens. There is a clear need for long-term studies of costs and trade-offs associated with induced resistance in crops under commercial conditions. Without such information, the potential offered by induced resistance is unlikely to be realized.     

Marker-assisted selection for disease resistance in wheat and barley breeding

Marker-assisted selection (MAS) provides opportunities for enhancing the response from selection because molecular markers can be applied at the seedling stage, with high precision and reductions in cost. About each of 50 genes conferring monogenic resistances and hundreds of quantitative trait loci (QTL) for quantitative disease resistances have been reported in wheat and barley. For detecting single-major gene resistance, MAS could be easily applied, but is often not necessary because the resistances are selected phenotypically. In quantitative disease resistances, MAS would be very useful, but the individual QTL often have small effects. Additionally, only a few monogenic resistances are durable and only a few QTL with high effects have been successfully transferred into elite breeding material. Further economic and biological constraints, e.g., a low return of investment in small-grain cereal breeding, lack of diagnostic markers, and the prevalence of QTL-background effects, hinder the broad implementation of MAS. Examples in which MAS has been successfully applied to practical breeding are the wheat rust resistance genes Lr34 and Yr36, the eyespot resistance gene Pch1, the recessive resistance genes rym4/rym5 to barley yellow mosaic viruses, mlo to barley powdery mildew, and two QTL for resistance to Fusarium head blight in wheat (Fhb1 and Qfhs.ifa-5A). Newly identified broad-spectrum resistance genes/QTL conferring resistance to multiple taxa of pathogens offer additional perspectives for MAS. In the future, chip-based, high-throughput genotyping platforms and the introduction of genomic selection will reduce the current problems of integrating MAS in practical breeding programs and open new avenues for a molecular-based resistance breeding.     

Strategy to identify resistance to white mould associated with high yield for irrigated common bean in Brazil

Development of common bean cultivars with partial white mould resistance through breeding techniques has been a challenge in Brazil. As yet, lines/cultivars from breeding programmes have not been investigated for resistance; therefore, this study screened 107 lines/cultivars for their reactions to white mould in 14 preliminary trials conducted under irrigation. Thirteen resistant lines/cultivars (three of Andean origin) and six Mesoamerican cultivars (three intermediately resistant and three susceptible) were selected for further investigation. These lines/cultivars and the resistant control A195 were evaluated in six advanced trials and two straw tests to assess the effectiveness of the screening procedure. In 11 preliminary trials, screenings were performed under moderate/high or higher disease pressure. These pressures occurred in two advanced trials in which, when yields were averaged across moderate/high and high pressures, 10 Mesoamerican lines/cultivars selected for resistance yielded 14%, 23%, and 38% more than intermediately resistant cultivars, A195, and susceptible cultivars, with median disease ratings (1–9 scale) of 4.5, 5.7, 5.7, and 6.7, respectively. In the straw test, three Andean lines/cultivars (A195 included) and two susceptible cultivars in the field were among those with the highest levels of physiological resistance. Thus, field rating under high disease pressure and greenhouse rating did not correlate significantly, suggesting that field trials are critical to evaluating resistance and to identifying high-yielding beans. Therefore, lines/cultivars from breeding programmes assessed in field trials may provide a low cost and fast way to identify high-yielding bean cultivars with partial resistance to white mould in the subtropical southern hemisphere.     

Resistance to powdery mildew (Blumeria graminis f.sp. avenae) in oat seedlings and adult plants

In this work, 165 Avena sativa and Avena byzantina accessions were screened for resistance to powdery mildew caused by Blumeria graminis f.sp. avenae and the defence mechanisms of resistant plants were further characterized. Ten resistant and moderately resistant accessions were selected according to macroscopic assessment. A detailed histological study of selected genotypes showed a range of defence mechanisms, acting alone or in combination, that impeded fungal development at different stages. Since the resistance observed in the collection was scarce, a study of adult plant resistance was carried out in 45 genotypes selected from field trials. Nine oat landraces and two commercial varieties showed very high levels of adult plant resistance. A detailed study of the components of the adult plant resistance revealed a high increase of penetration and post‐haustorial resistance in the fifth compared to the first leaves. Identification of the resistance sources and characterization of underlying defence mechanisms will be useful for future breeding programmes and for further cellular and molecular studies to unravel the genetic basis of resistance, in this species in particular and in cereal–powdery mildew interactions in general.     

Cowpea aphid-borne mosaic potyvirus: A review

Cowpea aphid-borne mosaic potyvirus (CABMV) is a cosmopolitan, economically significant seed-borne virus of cowpea. It can cause a yield loss of 13 - 87% under field conditions depending upon crop susceptibility, virus strain and the environmental conditions. CABMV has spread world-wide through the exchange of virus-infected germplasm material. The virus-infected seed provides the initial inoculum and aphids are responsible for the secondary spread of the disease under field conditions. The virus symptoms vary with the cowpea genotype and virus strain. Excellent sources of resistance are available for the breeding of resistant cultivars. Resistance in cowpea is conferred by either a dominant or a recessive gene. Enzyme-linked immunosorbent assay (ELISA) is the most appropriate method for the detection of the virus in the seed or plant tissue for seed certification programmes. Production of virus-free seed is another control option. Given the significance of the disease and the large area under cowpea cultivation in the world, we review knowledge of various aspects of the disease.     

Predicting quantitative host plant resistance against phoma black stem in sunflower

Phoma black stem is an important disease in sunflower, against which no specific management method is currently deployed in France. Relevant phenotyping methods for quantitative resistance are critical for integration of this trait into breeding programmes. Components of resistance associated with physiological resistance, and morphological traits associated with disease escape were measured on 21 sunflower genotypes under growth chamber (on seedlings), greenhouse (on adult plants), and field conditions, together with disease intensity in the field. Potential predictors were first selected for sensitivity and robustness from mixed model anova s. Analyses involving ranking tests and logistic regressions were then performed to identify predictors for field resistance. The identification of predictors for resistance involved analyses conducted in two broad steps: process‐oriented experiments, and epidemic‐oriented experiments. This stepwise approach departs from many studies aimed at identifying predictors for field resistance, which rely mainly on the computation of correlation coefficients between predictors and measured field disease variables. Predictors for quantitative resistance were identified: (i) lesion length and lesion expansion on stems of plants before flowering stage, and (ii) lesion length on first leaf petioles of seedlings. A high number of leaves and tall plants were associated with disease escape. Control genotypes for susceptibility and quantitative resistance were identified, and implications for breeding and improvements were derived from this work.     

Gene expression profiling of Medicago truncatula roots in response to the parasitic plant Orobanche crenata

Orobanche crenata is a root parasitic weed that is a major constraint for grain and forage legume cultivation in Mediterranean and West Asia. Only moderate to low levels of incomplete resistance of complex inheritance has been identified so far in legume crops, which has hampered genetic and genomic analysis. In the present study, we provide a gene expression profile of roots of the model legume Medicago truncatula in response to infection by O. crenata . M. truncatula accessions SA27774 (complete resistance acting at early penetration stages) and SA4087 (incomplete late acting resistance mediated by necrosis of parasite tubercle) were inoculated with O. crenata seeds in a semi-sterile dish system. Roots were harvested at 15 (first contacts of the parasitism structures with the host roots), 21 (initial stage of parasite tubercle formation on SA4087) and 35 (prior necrosis of well-developed parasite tubercle of on SA4087) days post-inoculation. Array hybridisations revealed several hundred genes up-regulated in response to O. crenata infection. Gene expression patterns suggest that resistance mechanisms activated in both genotypes are temporally and spatially different and resemble those associated with plant resistance to microbial pathogens. Regulated genes identified here represent a comprehensive resource that can be used as a support to breeding strategies for resistance.     

Crop Rotation,Monoculture and Soil Ecology1

Crop rotation and monocultural systems are discussed in relation to disease management and control. The ecological basis of both systems is examined. In crop rotation the leading principle is that in the absence of host-plants parasite populations invariably decline below threshold levels, mostly within a few years. Knowledge of the behaviour of parasite populations in the soil is becoming vast, especially in the case of plant parasitic nematodes. Analysis of soil samples for nematodes and fungi often helps the farmer to plan rotation schemes. From the ecological point of view, crop rotation is interpreted as a system of interrupted monocultures. Monocultural practices cannot be applied so universally as crop rotation, but in a number of cases monoculture is feasible, e.g. with cereals. In the equilibrium situation between host-plant (e.g. cereals), parasite (Gaeumannomyces graminis [Sacc.] v. Arx & Olivier) and the antagonists of the parasite, growth of the crop is satisfactory. These cases illustrate the role of antagonists, and it is proposed to investigate more systematically monocultures in relation to antagonisms. This may lead to more opportunities for biological control of soil-borne diseases.     

In vitro culture of Orobanche ramosa

Orobanche spp. (broomrapes) are holoparasites that subsist on the roots of many important crops and can considerably reduce yield. The control of Orobanche spp. includes physical, chemical and biological methods. Interactions between parasitic angiosperms and their hosts first occur at the level of parasite seed germination. The seeds of all Orobanchaceae germinate in soil under natural conditions only in response to specific chemical exudates from the host plant. This study describes the influence of different plant growth regulators and host plant root exudates on germination and development of calli from Orobanche seeds in vitro . The effect of indole-3-acetic acid, gibberellic acid and kinetin on the germination of Orobanche seeds varied with concentration. These plant growth regulators also affected the period of germination and the structure of calli and protrusions. An in vitro system for the collection of tobacco root exudates was established. Compounds released from the host roots of three different tobacco cultivars were found to provoke high levels of germination of the Orobanche seeds without any period of pre-conditioning. This study developed methods for the investigation of host–parasite interactions and the effect of germination stimulants in Orobanche spp.     

Sources of resistance to septoria tritici blotch and implications for wheat breeding

Twenty-four wheat cultivars and breeding lines were screened for isolate-specific resistance to septoria tritici blotch (STB) caused by 12 isolates of Mycosphaerella graminicola. New isolate-specific resistances that could be used in wheat breeding were identified. Major sources of resistance to STB used in world breeding programmes for decades, such as Kavkaz-K4500, Veranopolis, Catbird and TE9111, have several isolate-specific resistances. This suggests that 'pyramiding' several resistance genes in one cultivar may be an effective and durable strategy for breeding for resistance to STB in wheat. Several cultivars, including Arina, Milan and Senat, had high levels of partial resistance to most isolates tested as well as isolate-specific resistances. Resistance to isolate IPO323 was common, present in all but one of the major sources of resistance tested. This suggests that resistance to IPO323 may be an indicator of varietal resistance to STB in the field.     

Revisiting strategies for reducing the seedbank of Orobanche and Phelipanche spp.

Weedy Orobanche and Phelipanche species are important in Southern and Eastern Europe, the Middle East and North Africa, and have recently been reported in the USA, Australia and some countries in South America. Rather than being controlled, the Orobanche and Phelipanche problem is increasing, both in intensity and in acreage. Large areas of new territory are at risk of invasion, if care is not immediately taken to limit the introduction of parasitic seeds and to educate farmers and others to be alert for new infestations. There is an urgent need to re-evaluate novel integrated Orobanche and Phelipanche management programmes that will allow a better control of the parasite species and limit their distribution. The main obstacle in the long-term management of Orobanche and Phelipanche infested fields is the durable seedbank, which may remain viable for decades in the field. Large quantities of long-lived seeds give the parasite genetic adaptability to changes in host resistance and cultural practices. As long as the seedbank is not controlled, the need to control the parasite will persist whenever a susceptible host is grown in the infested field. We discuss strategies for seedbank reduction, paying particular attention to cultural practices, whereas chemical and biological control methods are covered by other reviews in this issue.     

Breeding for resistance: breeding for Plum pox virus resistant plums (Prunus domestica L.) in Germany

In German plum breeding programmes new varieties tolerant to Plum pox virus (PPV) were initially obtained by selecting hybrids originating from crossings between tolerant old varieties while, later on, sharka tolerant varieties bred in Čačak, Serbia were used as crossing partners. Varieties released from these programmes replaced the sensitive ones in all sharka infected regions in Germany. While all the new tolerant varieties can be infected by the virus, PPV symptoms on the fruits are acceptable. However, environmental factors can weaken the plant, causing them to suffer more from PPV infections and to display worse symptoms on fruits, as did occur during some recent very dry and warm years. A breeding programme at the University of Hohenheim is tackling this problem with genotypes that show a hypersensitive response after infection. These genotypes are completely field resistant to PPV, remaining virus free in the field since they cannot be infected via aphid transmission. They are able to isolate the virus after infection. The first hypersensitive variety was 'Jojo'. Many seedlings originating from crossings with at least one hypersensitive parent are under evaluation. Since the inheritance of the trait 'hypersensitivity against PPV' is very good, combining hypersensitivity with excellent fruit quality and good cropping capacity will be possible soon. In 2005, a breeding programme for hypersensitive Prunus genotypes began at the Technical University of Munich.     

43个中国小麦品种(系)抗叶锈性研究

 选用12个墨西哥叶锈菌生理小种对43个中国小麦品种(系)所携带的抗叶锈病基因进行了推导,在25个品种(系)中推导出6个抗叶锈基因Lr1,Lr10,Lr13,Lr14a,Lr16和Lr26,9个品种(系)对本试验所使有的12个叶锈菌生理小种都表现感病反应,另有9个品种(系)携带未知的抗叶锈基因。在墨西哥2个地点进行的田间成株期抗叶锈性试验表明,12个品种(系)表现慢叶锈性,在将来的抗病育种中有一定的应用价值。     

Domestication affected the basal and induced disease resistance in common bean (Phaseolus vulgaris)

Crop plants exhibit reduced levels of disease resistance, but little is known about the specific resistance mechanisms that are affected by breeding for increased yields. We investigated basal and chemically induced resistance of two wild accessions and four cultivars (including one landrace and three ??modern??, yield-improved cultivars that have been produced by hybridisation and pedigree breeding) of common bean (Phaseolus vulgaris) under greenhouse and field conditions. After treatment with benzothiadiazole, a widely used inducer of systemic acquired resistance, plants were challenged with one of two bacterial pathogens (Pseudomonas syringae pv. syringae and Enterobacter sp. strain FCB1). Basal resistance to Pseudomonas in the wild accessions was significantly higher than in the cultivars. Moreover, benzothiadiazole-treatment elevated resistance to the same pathogen in a wild accession and the landrace, but not in the yield-improved cultivars. Similarly, benzothiadiazole-induced resistance to Enterobacter FCB1 was detected in both wild accessions and the landrace, whereas the same treatment enhanced susceptibility to Enterobacter FCB1 in two of the yield-improved cultivars. Basal resistance to Pseudomonas was highly (but negatively) correlated to induced resistance over all accessions, and basal and inducible resistance to Enterobacter FCB1 were negatively correlated for the cultivars, but not when considering all six accessions. Benzothiadiazole-treatment increased growth rates under pathogen pressure of the wild accessions but not the cultivars. Apparently, the yield-improved cultivars investigated here have lost a considerable part of the basal and induced broad-spectrum disease resistance that characterises their wild relatives and to some degree also the landrace. Two of the yield-improved cultivars even became highly susceptible to infection by an Enterobacter strain that has not yet been described as a pathogen of bean and that is likely to represent a common environmental or phyllosphere bacterium. Future studies should disentangle the effects of domestication on the various layers of plant resistance to pathogens and consider the potential of wild accessions and landraces for future breeding programmes.     

The effect of insecticide application sequences on the control and insecticide resistance status of the peach-potato aphid, Myzus persicae (Hemiptera:Aphididae), on field crops of potato

Experiments were done on commercial potato crops in the UK to investigate the effect of different insecticide sequences on the control and insecticide resistance status of Myzus persicae (Sulzer). The work was done to provide field validation of similar laboratory studies done in 'field simulators'. To ensure adequate aphid populations and to influence the initial resistance status of the aphid population, cultured M. persicae from a clone of known resistance status (esterase R1, kdr heterozygote, non-MACE (modified acetylcholinesterase)) were inoculated into both experiments. Two-spray programmes starting with lambda-cyhalothrin (a pyrethroid insecticide) gave poor control in comparison with programmes starting with pirimicarb (a carbamate insecticide) or pirimicarb-containing mixtures. This concurred closely with the results obtained from single applications in field simulator studies. Treatment sequences containing pymetrozine (a pyridine azomethine insecticide) were also effective, though slower-acting. This again concurs with field simulator studies. The proportions of aphids carrying different resistance mechanisms were largely unaffected by treatment in these experiments. The implications of these results for field control strategies are discussed.     

Genetic resistance for the sustainable control of plant virus diseases: breeding,mechanisms and durability

Plant viruses are important agricultural pathogens, and are responsible for a significant number of commercially relevant plant diseases. There are very few efficient control measures for viral diseases, but the use of genetic resistance appears to be the most promising strategy, often conferring effective protection without additional costs or labour during the growing season, and without damaging the environment. Sources of virus resistance have been identified for most crop species, and many resistant cultivars are already commercially available and of widespread cultivation; however, much remains to be learned about genetic resistance. This review article considers three main aspects that require intense investigation. First, we review the identification of sources of resistance and how plant breeders and pathologists have focused on aspects of the breeding process particularly relevant to viruses, such as germplasm screening and the dissection of resistance phenotypes. Second, we review how molecular mechanisms controlling resistance have been unravelled, looking at case studies where resistance mechanisms are now understood in detail for each stage of the infection cycle. Third, we turn to the durability of resistance in a global context, examining factors that influence durability and how this can be predicted. We conclude with a short discussion of the technological and scientific opportunities provided by recent advances in the field.     

Bacterial wilt resistance in tomato associated with tolerance of vascular tissues to Pseudomonas solanacearum

High populations of Pseudomonas solanacearum were detected in some, but not all stems of bacterial wilt resistant ('CRA 66', 'Hawaii 7996' and 'Caraibo') and susceptible ('Floradel') tomatoes. Latent infection, i.e. spread of P. solanacearum into xylem vessels, was confirmed in Caraibo, Hawaii 7996 and 'CRA 66' (the resistant parent of Caraibo). None of the plants within the resistant cultivars wilted and those cultivars were characterized by tolerance of the vascular tissues to high bacterial densities. In contrast, plants of cultivar Floradel showed consistent symptoms and wilted rapidly, with higher mean bacterial density than resistant cultivars. Bacterial wilt resistance was not associated with resistance to bacterial root invasion but with the capability of the plant to limit P. solanacearum colonization in the stem. The extent of bacterial colonization is proposed as a criterion to quantify tolerance, complementary to absence of external wilt symptoms used in breeding programmes for resistance.     

A model system for rapid selection for resistance and investigation of resistance mechanisms in Fusarium wilt of oil palm

Fusarium wilt ( Fusarium oxysporum f.sp elaeidis ) is the most serious disease of oil palm in Africa. Control measures are dependent upon breeding for resistance, but the lack of a rapid screen for resistance and lack of understanding of the underlying resistance mechanisms has constrained breeding programmes. A novel petiole infusion technique is reported that exploits the ability of the majority of petiole cells to express disease resistance or susceptibility. A clear correlation was found between disease resistance or susceptibility of six clones and external symptoms and fungal colonization in petioles. Antifungal compounds accumulated in resistant but not in susceptible clones in response to inoculation; there was also some evidence of preformed antifungal compounds in resistant clones. Further investigation of resistance mechanisms can be undertaken using this model system. The use of this novel technique, both as a potential rapid replacement for current nursery selection methods (time reduced from 8 months to 8 days) and as an assessment of resistance in individual field palms, is discussed. *