Evolution of a pathogen population in host mixtures: rate of emergence of complex races

 The development of a pathogen population in a crop varietal mixture was studied with an epidemic simulator based on the model EPIMUL. The pathogen population was composed of simple races, able to develop on only one genetic component of the mixture, and a complex race, which developed on all mixture components. The complex race was modelled with a very low initial frequency compared to simple races, to simulate the emergence of a complex race in the field. Several successive epidemics were simulated as if the pathogen population reproduced on the same plot for several years. The development of the complex race on the simulated plot was either focal or uniform. The effects of the cost of virulence, of density dependence and of differential adaptation to host genetic background on the simple race-complex race competition were studied. Experimentally measured values of the cost of virulence and differential adaptation were incorporated into the model, and both factors were shown to greatly reduce the increase in frequency of the complex race over time. Density dependance also influenced race competition, but mainly for high values of the parameter. Our results suggest that the cost of virulence is probably not the only mechanism that may influence the simple race-complex race competition in host mixtures. In our simulations, differences in the spatial distribution of the initial inoculum between parasite races led to large differences in their final frequencies. Thus, more investigations, including randomized disease distributions, would be of interest to judge the potential importance of spatial effects in the field. Received: 4 October 1996 / Accepted: 3 January 1997     

Detection of Cochliobolus heterostrophus races in South China

Cochliobolus heterostrophus is the causal pathogen of the southern corn leaf blight (SCLB). There are three known races: race O, race C and race T. To determine which C. heterostrophus races comprise the field population in southern China and to assess diversity of these strains in terms of virulence, 200 isolates from diseased plants were collected in nine provinces/municipalities. All were race O, that is, no race T or race C isolates were found. Sixty race O isolates that sporulated well were chosen for further analysis. Virulence was measured using the integral optical density (IOD) of leaf lesions on four maize inbred lines. UPGMA cluster analysis of AFLP markers was applied to the 60 race O isolates plus control race O, T and C strains. Phylogenetic distribution, geographic location and virulence were not correlated. These results can provide valuable information for guidance in early warning and disease control.     

Pathogen populations evolve to greater race complexity in agricultural systems--evidence from analysis of Rhynchosporium secalis virulence data

Fitness cost associated with pathogens carrying unnecessary virulence alleles is the fundamental assumption for preventing the emergence of complex races in plant pathogen populations but this hypothesis has rarely been tested empirically on a temporal and spatial scale which is sufficient to distinguish evolutionary signals from experimental error. We analyzed virulence characteristics of ≈ 1000 isolates of the barley pathogen Rhynchosporium secalis collected from different parts of the United Kingdom between 1984 and 2005. We found a gradual increase in race complexity over time with a significant correlation between sampling date and race complexity of the pathogen (r(20) = 0.71, p = 0.0002) and an average loss of 0.1 avirulence alleles (corresponding to an average gain of 0.1 virulence alleles) each year. We also found a positive and significant correlation between barley cultivar diversity and R. secalis virulence variation. The conditions assumed to favour complex races were not present in the United Kingdom and we hypothesize that the increase in race complexity is attributable to the combination of natural selection and genetic drift. Host resistance selects for corresponding virulence alleles to fixation or dominant frequency. Because of the weak fitness penalty of carrying the unnecessary virulence alleles, genetic drift associated with other evolutionary forces such as hitch-hiking maintains the frequency of the dominant virulence alleles even after the corresponding resistance factors cease to be used.     

The significance of biological races of Ditylenchus dipsaci and their hybrids

Pure cultures of six races of Ditylenchus dipsaci derived from a single female, namely, lucerne race (LR), red clover race (RCR), white clover race (WCR), narcissus race (NR), tulip race (TR) and oat race (OR) were inoculated into eight plant species growing in pots. Onion was host to all six races, and tulip to all except RCR, whereas lucerne was susceptible only to LR and slightly to WCR. Hyacinth was not very susceptible to any race. The TR and OR were the most polyphagous. Severe symptoms were not necessarily associated with large nematode populations. Parthenogenesis did not occur. Ten fertile hybrids between races were produced and the host range of five of these was tested. On average the hybrids multiplied less than their parent races and their host ranges showed no relationship to those of their parent races. RCR and TR inoculated together into tulip produced significantly fewer nematodes than did TR alone and more than RCR alone. Mixed populations of races occur in nature which are probably a mixture of parental races, their hybrids and back-crosses. Repeated back-crossing of the hybrids with parental types and the slower multiplication of the hybrids is an explanation for the variation in host range of known races and the failure to record new races.     

HOST-PATHOGEN INTERACTIONS IN NATURAL POPULATIONS OF LINUM MARGINALE AND MELAMPSORA LINI: I. PATTERNS OF RESISTANCE AND RACIAL VARIATION IN A LARGE HOST POPULATION

Populations of wild flax, Linum marginale and its associated rust fungus Melampsora lini growing at Kiandra, New South Wales, Australia, were sampled during the 1986–1987 growing season. Thirteen different races of M. lini were detected in a sample of 96 isolates. The distribution of isolates was uneven: race A comprised 73% of the samples; race N, 8%; and race H, 5%; while the remaining races were represented by only one or two samples. The dominance of race A increased over the course of the growing season, comprising 67% of the early season samples and increasing to 78% for those collected late in the season. The overall diversity of the pathogen population decreased late in the growing season, but this trend was not statistically significant. The average virulence of individual isolates of the pathogen population increased during the growing season. This trend was most pronounced among the minor races, where the mean number of differential hosts infected increased from 4.58 for early season samples to 5.12 and 5.08 for mid and late season samples, respectively. In contrast to the virulence pattern in the pathogen, the L. marginale population displayed a more even distribution of resistance. In a sample of 67 plants 10 resistance phenotypes were detected from their pattern of resistance/susceptibility to seven pathogen isolates. No phenotype had a frequency that exceeded 30%. Resistance phenotypes were randomly distributed on both a population level and on a fine scale.     

Relative Virulence of Meloidogyne incognita Host Races on Soybean

Sensitivity and host efficiency of susceptible (''Lee 68'', ''Coker 156'') and resistant (''Bragg'', ''Centennial'', ''Forrest'', ''Lee 74'') soybean (Glycine max (L.) Merr.) cultivars for races of Meloidogyne incognita (Mi) were determined in greenhouse experiments. Eight Mi populations collected from the southeastern United States were utilized. All Mi races reproduced readily on Lee 68 and Lee 74 and moderately on Forrest and Bragg. Coker 156 exhibited resistance to races 1 and 2, and some race 3 populations, but was very susceptible to certain race 3 and 4 populations. Reproduction of all races was lowest on Centennial. Forrest and Centennial shoot growth was not significantly suppressed by any race. There were no distinct differences in virulence between races except for a race 3 population which reproduced readily on all cultivars, stunting their growth. Considerable variation in reproduction existed within races 1 and 3.     

Sequence variations in alleles of the avirulence gene avrPphE.R2 from Pseudomonas syringae pv. phaseolicola lead to loss of recognition of the AvrPphE protein within bean cells and a gain in cultivar-specific virulence

The bean halo blight pathogen, Pseudomonas syringae pv. phaseolicola ( Psph ), is differentiated into nine races based on the presence or absence of five avirulence ( avr ) genes in the bacterium, which interact with corresponding resistance genes, R1–R5 , in Phaseolus vulgaris . The resistance gene R2 is matched by avrPphE , which is located adjacent to the cluster of hrp genes that are required for pathogenicity of Psph . Although only races 2, 4, 5 and 7 are avirulent on cultivars with R2 (inducing the hypersensitive response; HR), homologues of avrPphE are present in all races of Psph . DNA sequencing of avrPphE alleles from races of Psph has demonstrated two routes to virulence: via single basepair changes conferring amino acid substitutions in races 1, 3, 6 and 9 and an insertion of 104 bp in the allele in race 8. We have demonstrated that these base changes are responsible for the difference between virulence and avirulence by generating transconjugants of a virulent race harbouring plasmids expressing the various alleles of avrPphE. Agrobacterium tumefaciens -directed expression of avrPphE from race 4 in bean leaves induced the HR in a resistance gene-specific manner, suggesting that the AvrPphE protein is alone required for HR induction and is recognized within the plant cell. The allele from race 6, which is inactive if expressed in Psph , elicited a weak HR if expressed in planta , whereas the allele from race 1 did not. Our results suggest that the affinity of interaction between AvrPphE homologues and an unknown plant receptor mediates the severity of the plant's response. Mutation of avrPphE alleles did not affect the ability to colonize bean from a low level of inoculum. The avirulence gene avrPphB , which matches the R3 resistance gene, also caused a gene-specific HR following expression in the plant after delivery by A. tumefaciens .     

The genetic relationship between races of Bremiae lactucae and cultivars of Lactuca sativa

The reactions of lettuce cultivars to physiologic races of Bremia lactucae are interpreted in terms of a gene-for-gene relationship between pathogen and host. The hypothesis takes into account the parentage of cultivars and the origins of their resistance, the characteristics of the resistance reactions and data available from detailed genetical analysis of various race/cultivar combinations. Cultivars are classified with respect to ten postulated resistance genes and B. lactucae races are defined by the virulence genes present. The practical significance of these studies is discussed in relation to both future lettuce breeding programmes and to the choice of cultivars available to counteract any given local race situation.     

Modelling the evolution of common cuckoo host‐races: speciation or genetic swamping?

Co‐evolutionary arms races have provided clear evidence for evolutionary change, especially in host–parasite systems. The evolution of host‐specific races in the common cuckoo (Cuculus canorus), however, is also an example where sexual conflict influences the outcome. Cuckoo females benefit from better adaptation to overcome host defences, whereas cuckoo males face a trade‐off between the benefits of better adaptation to a host and the benefits of multiple mating with females from other host‐races. The outcome of this trade‐off might be genetic differentiation or prevention of it by genetic swamping. We use a simulation model to test which outcome is more likely with three sympatric cuckoo host‐races. We assume a cost for cuckoo chicks that express a host adaptation allele not suited to their foster host species and that cuckoo males that switch to another host‐race experience either a fitness benefit or cost. Over most of the parameter space, cuckoo male host‐race fidelity increases significantly with time, and gene flow between host‐races ceases within a few thousand to a hundred thousand generations. Our results hence support the idea that common cuckoo host‐races might be in the incipient stages of speciation.     

Serological Comparison of Two Races of Fusarium oxysporum f. sp. lupini

Water-soluble mycelial antigens from two physiological races (2 and 3) of Fusarium oxysporum L. sp. lupini were compared by tandem-crossed immunoelectrophoresis. When antiserum against race 3 was tested some 50 antigens were detected. The two races had apparently almost identical antigenic patterns differing only in one antigen specific to race 3. This specific antigen might be related to the virulence of this fungus.     

Acclimation of Plasmopara halstedii (sunflower downy mildew) in relationship with trade-off between virulence and aggressiveness in a local pathogen population

The acclimation in relationship with virulence cost was analysed for seven Plasmopara halstedii (sunflower downy mildew) isolates including five progeny isolates of several races descending from two parental isolates of races 100 and 710. Aggressiveness criteria were analysed in one sunflower inbred line showing a high level of quantitative resistance. Isolates of races 100 and 3xx were characterised with shorter latent period and higher sporulation density than isolates of races 7xx. All isolates showed high percentage infection values and caused a large reduction in seedling size except for one isolate involved in dwarfing. The seven isolates were divided, according to their virulence and aggressiveness, into two main groups as more aggressive isolates of the 100 and 3xx races which do not overcome the sunflower differential host D3, and less aggressive isolates of 7xx races which can overcome D3. Consequently, the 100 and 3xx avirulent races had a virulence cost measured by differences in aggressiveness (from 45.5 to 76.3%) compared to 7xx virulent races carrying unnecessary virulence gene.     

Pathogenicity and its alternation with both morphological and genetic variation in Plasmopara halstedii (sunflower downy mildew)

Morphological, pathogenic and genetic variation was studied in seven Plasmopara halstedii (sunflower downy mildew) isolates of several races using five singlezoosporangium isolates per pathogen isolate. Aggressiveness criteria were analysed in one sunflower inbred line showing a high level of quantitative resistance. Genetic relationships were detected between the single zoosporangium isolates using 12 expressed sequence tags (EST)-derived markers. Analysis of the five single zoosporangium isolates for P. halstedii isolates showed variability within pathogen isolates for all aggressiveness criteria, but not for all pathogen isolates. Isolates of races 100 and 3xx were characterised with shorter latent period and higher sporulation density than the isolate of races 7xx. All pathogen isolates showed high percentage infection values and caused a large reduction in seedling size except for one isolate involved in dwarfing. There was no relation between zoosporangia form or size and race virulence profiles or aggressiveness criteria. There was no intra-genetic variation for all pathogen isolates, but it was observed an important genetic variation between single zoosporangium isolates of all races. No correlation was detected between pathogenicity traits and EST genotypes.     

Characterization of the Virulence Phenotypes of Heterodera glycines in Minnesota

Knowledge of the virulence phenotypes of soybean cyst nematode, Heterodera glycines populations is important in choosing appropriate sources for breeding resistant cultivars and managing the nematode. We investigated races of 59 H. glycines populations collected from 1997 to 1998 and races and HG Types of 94 populations collected in 2002 from soybean fields across southern and central Minnesota. In the 1997 to 1998 samples, race 3 was predominant and represented 78% of the populations. The remaining populations were 11.9% race 1, 1.7% race 4, 6.8% race 6, and 1.7% race 14. In the 2002 samples, the populations were classified as 15.3% race 1, 77.6% race 3, 2.4% race 5, 3.5% race 6 and 1.2% race 9. Percentage of 1997 to 1998 populations with female indices (FI) higher than 10 were 10.2% on Pickett 71, 3.4% on Peking, 13.6% on PI 88788, 3.4% on PI 90763, 1.7% on PI 209332, and 1.7% on PI 437654. Percentage of 2002 populations with FI >10 was 1.1% on Peking, 17.0% on PI88788, 14.9% on PI 209332, 33.0% on PI 548316, 11.7% on Pickett 71, and 0% on the other three indicators, PI 90763, PI 437654, and PI 89772. The line PI 548316 was relatively susceptible to the Minnesota H. glycines populations and may not be recommended for breeding resistant cultivars in the state. There was no noticeable change of frequencies of virulence phenotypes in response to the use of resistant cultivars during 1997 to 2002 in Minnesota except that FI increased on the PI 209332.     

Distribution of genetic variability in populations of two chromosomal races of Dichroplus pratensis (Melanoplinae, Acrididae) and their hybrid zone

We examined, through allozyme electrophoresis, the genetic structure of populations of the acridid grasshopper Dichroplus pratensis from two chromosomal races (Northern and Southern) and their hybrid zone in Argentina. No fixed alleles for any particular race were found, although genetic differentiation among parental races was significant (0 = 0.044, 95% CI: 0.004-0.068). Hybrid populations are genetically more similar to the Southern race (0 = 0.008, 95% CI: -0.005-0.018) than to Northern ones (0 = 0.018, 95% CI: 0.002-0.030). Differential viability or fertility of hybrids, or asymmetry in mating preferences in favour of one particular cross would cause a higher proportion of matings between hybrid individuals and those from the Southern race. This would explain the high genetic similarity between those groups, in spite of their geographical vicinity with northern race populations.     

Host-Pathogen Interactions : XX. BIOLOGICAL VARIATION IN THE PROTECTION OF SOYBEANS FROM INFECTION BY PHYTOPHTHORA MEGASPERMA F. SP. GLYCINEA

Phytophthora megasperma f.sp. glycinea, which causes soybean (Glycine max) root and stem rot, exists as several races which differ in their ability to infect a range of soybean cultivars. A glycoprotein-rich fraction (Fraction I) isolated from fungal culture fluid protects soybean seedlings from infection with compatible races. In an early study (13), seedlings were protected only by Fraction I purified from incompatible races. In 1979, seedlings were better protected by Fraction I isolated from incompatible races than by Fraction I isolated from compatible races. In 1980, seedlings were protected equally well by Fraction I from incompatible and compatible races. Materials similar in composition to Fraction I did not protect seedlings from infection. No cause could be identified for the apparent change, during the 3-year period, in the race specificity of the protection assay. Variability in the bioassay prohibited further purification or characterization of Fraction I components that protect seedlings from infection.     

水稻品种对美国稻瘟病小种IB-33、IB-45和IE-1的抗性遗传

1995年10月至1997年11月,在美国阿肯色大学水稻研究推广中心,用水稻品种LA110和Jasmine-85与水稻品种Teqing、Katy、Mars、LaGrue和Newbonnet进行不完全双列杂交,对其杂交后代和亲本用美国3个主要稻瘟病菌小种(以下简称小种)IB-33、IB-45和IE-1进行接种鉴定和遗传分析研究.结果表明:亲本LA110、Jasmine-85、Teqing抗所有3个小种.Katy抗小种IB-45和IE-1,感小种IB-33.Mars抗小种IE-1,感小种IB-33和IB-45.LaGrue感所有3个小种.Newbonnet抗小种IB-45,感小种IB-33和IE-1.所有抗病亲本的抗病基因,其F1分别对相应小种呈现显性抗病性.抗病亲本杂交,LA110与Jasmine-85对小种IB-33,LA110与Teqing、Jasmine-85对小种IE-1,及Jasmine-85与Teqing对小种IE-1,是等位的抗病基因.LA110与Teqing对小种IB-33,及Jasmine-85与Teqing对小种IB-33,分别存在三对独立遗传的显性抗病基因.LA110与Teqing、Katy、Newbonnet、Jasmine-85对小种IB-45,Jasmine-85与Teqing、Katy、Newbonnet对小种IB-45,LA110与Katy、Mars对小种IE-1,Jasmine-85与Katy、Mars对小种IE-1,分别存在两对独立遗传的显性抗性基因.抗病亲本LA110或Jasmine-85与感病亲本Mars对小种IB-33,抗病亲本LA110与感病亲本Mars对小种IB-45,具有两对显性互补抗病基因,当两对显性抗病基因同时存在时,表现出抗性.抗病亲本LA110或Jasmine-85与感病亲本Katy、LaGrue、Newbonnet对小种IB-33,抗病亲本LA110与感病亲本LaGrue对小种IB-45,抗病亲本Jasmine-85与感病亲本Mars、LaGrue对小种IB-45,抗病亲本LA110或Jasmine-85与感病亲本LaGrue、Newbonnet对小种IE-1,分别存在一对显性抗病基因.两个亲本正、反交的遗传表现一致.本文也讨论了LA110、Teqing和Jasmine-85三个抗病品种在美国水稻抗病育种中利用的可能性.     

Pathotypic and molecular evolution of contemporary population of Puccinia striiformis f. sp. tritici in Egypt during 2016–2018

The contemporary races of Puccinia striiformis f. sp. tritici (Pst) in Egypt during 2016–2018 were differentiated based on virulence and molecular patterns. Virulence patterns based on the reaction of the 17 World/European differential sets carrying stripe rust resistance genes (Yr genes) resulted in ten races including four new (first recorded in Egypt) and six old (previously recorded in Egypt). The new races were identified as 64E0 (virulence [V] Yr4, Su), 0E16 (V Yr8, 19), 66E0 (V Yr4, 7, 22, 23, Su) and 4E130 (V Yr2, 6, 7, 25, HVII), while the old were 0E0 (avirulence), 2E0 (V Yr7, 22, 23), 2E16 (V Yr7, 8, 19, 22, 23), 4E0 (V Yr2, 6), 6E4 (V Yr2, 6, 7, 22, 23, 25) and 70E4 (V Yr2, 4, 6, 7, 22, 23, 25, Su). Cluster analysis differentiated Pst races based on virulence frequency to Yr genes. Simple sequence repeat (SSR) markers were used to detect the molecular polymorphism of the Pst races. Clustering separated the old and new races into two groups, indicating their common ancestry since the new races were very distinct from the old races. Although clustering based on virulence revealed some evolutionary patterns, where the new races 64E0 and 66E0 may have probably evolved from the old races (2E16, 2E0, 6E4, 70E4) and the new race 4E130 may be evolved from the joint race 4E0. However, clustering based on molecular patterns indicated that the new races appear to be genetically distinct and may represent an exotic introduction rather than a mutation in isolates of the old races. A weak association between virulence and molecular patterns revealed that they are independent of each other. The SSR markers did not correspond to the virulences in the pathogen. Further studies on the potential virulence genes of the detected Pst virulences are needed.     

STUDIES ON THE PHYSIOLOGIC SPECIALIZATION OF YELLOW RUST (PUCCINIA GLUMARUM (SCHM.) ERIKSS. & HENN.) IN GREAT BRITAIN

During 1945-8, 223 British collections of yellow rust were determined on Gassner & Straib's differential hosts, with the addition of the wheat variety Wilma. Physiologic races 2, 3, 5, 6, 7, 8 and 17 were obtained from wheat, race 46 from barley, race 33 from Hordeum murinum and race 28 from Agropyron repens. Three new races, 6x from wheat, M from H. marinum and G from Dactylis glomerata were also isolated. High temperatures increase the susceptibility of some differential hosts to certain wheat races, but decrease that of others. Biotypes of races 2, 6 and 8, differing from the 'type' cultures of those races, were isolated.
Of the commoner wheat races, nos. 6 and 8, occurring on many wheats, were widespread, but races 5 and 7, on fewer varieties, were confined to the north of Britain, and races 2 and 3, on very few varieties, to the south. Certain grasses are partially susceptible to some of the cereal races. The uredospores of race 2 germinate less well than those of races 5 or 8, other wheat races being intermediate. The optimum temperature for spore germination of race G is 22-5 C, that of all other races 10–13^oC.
Twelve wheat varieties were inoculated in the field with each of the eight wheat races, and some varieties developed field resistance to certain races. The races attacking a variety the most severely in the field inoculations were usually the races isolated from that variety in collections received.     

Detection of Races 1 and 2 of Fusarium solani f. sp. cucurbitae and their Distribution in Watermelon Fields in Tunisia

A crown, foot and fruit rot of watermelon has been observed in most of the watermelon production areas in Tunisia. A survey conducted from 2000 to 2001 allowed the isolation of 291 isolates which were identified as Fusarium solani. These isolates were identified as F. solani f. sp. cucurbitae (Fsc) and races 1 and 2 characterized on the basis of pathogenicity tests on watermelon seedlings and muskmelon fruits. These results were confirmed by counts of the number of septa in the macroconidia. About 271 isolates were identified as Fsc race 1, 12 isolates were identified as Fsc race 2 and eight isolates were not pathogenic. Race 1 is widely distributed in watermelon production areas in Tunisia and race 2 has a lower incidence but it is present in the north, the middle and southern Tunisian watermelon cropping areas. Additionally, a study to compare the virulence of 122 isolates of Fsc race 1 showed different degrees of virulence among them. This is the first report of Fsc races 1 and 2 in Tunisia.