The effect of specific and non-specific resistance in mixtures of barley or genotypes on infection by mildew (Erysiphe graminis f. sp. hordei) and on yield

Summary Preliminary field trials indicated that the yields of mixtures of barley cultivars partially resistant to mildew (Erysiphe graminis f. sp. hordei) were greater than those expected from the mean of their components. Subsequent trials in 1988, 1989 and 1990 demonstrated that such mixtures gave enhanced yield. However, fungicide treatment of mixtures gave significant positive and negative effects on yield. In mixtures with cultivars containing partial resistance there was evidence of a positive relationship between high yield loss due to mildew (low disease tolerance) in cultivars grown in monoculture and their yield advantage in mixtures. The converse resulted in yield disadvantage.     

Wheat grain yield and stability assessed through regional trials in the Eastern Gangetic Plains of South Asia

Improving the level and stability of grain yield is the primary objective of wheat breeding programs in the Eastern Gangetic Plains (EGP) of South Asia. A regional wheat trial, the Eastern Gangetic Plains Yield Trial (EGPYT), was initiated by CIMMYT in collaboration with national wheat research programs in Bangladesh, Nepal, and India in 1999–2000 to identify wheat genotypes with high and stable grain yield, disease resistance, and superior agronomic traits for the EGP region. A set of 21 wheat experimental genotypes selected from a regional wheat screening nursery in South Asia, three improved widely grown cultivars (Kanchan, PBW343 and Bhrikuti), and one long-term cultivar (Sonalika) were tested at 9–11 sites in six wheat growing seasons (2000–2005) in the EGP. The 21 experimental genotypes were different in each year, whereas the four check cultivars were common. In each year, one or more of the experimental genotypes showed high and stable grain yield and acceptable maturity, plant height, and disease resistance compared to the check cultivars. Three improved cultivars have already been commercially released in the region through EGPYT and many germplasm lines have been used in the breeding programs as parents. Identification of wheat genotypes with high-grain yield in individual sites and high and stable yield across the EGP region underlines their value for regional wheat breeding programs attempting to improve grain yield and agronomic performance.     

Barley mildew in Europe: population biology and host resistance

Summary Isolates of the barley mildew pathogen from the air spora over a large part of Europe and from fields of variety mixtures, were tested for virulence against 12 host resistance alleles. Subsamples were tested for their response to triadimenol fungicide and analyzed for 10 DNA loci using RAPD markers and PCR. There was a large range of haplotypes spread over Europe; irregularity in the distribution was probably due mainly to non-uniform use of the corresponding host resistances and fungicides. A large range of variation was also detectable within individual fields. Positive gametic disequilibria distorted the distribution of virulence alleles among haplotypes and reduced the number of haplotypes detectable in the sample. Analysis of the spread of the newly selectedVal3 allele into different European sub-populations indicated that gene flow throughout the population may be rapid for alleles that have a selective advantage.Fungicide resistance was widespread in areas known for intensive use of fungicides for mildew control. Four classes of fungicide response were detectable and particular virulence haplotypes were found to be characteristic for each class.Variety mixtures used in the former German Democratic Republic (GDR) reduced mildew infection, and thus fungicide use, during the years 1984–1991 despite the limited variation in host resistance among the mixtures. A tendency for complex pathogen races to increase in mixture crops was reversed by the large-scale re-introduction of fungicides for mildew control in 1991. The mixture strategy appeared to be more successful than using the same resistance alleles in pure monoculture or combining them in a single host genotype.     

Identification of bread wheat,durum wheat and barley cultivars adapted to dry areas of Southern Italy

Summary Among the cultivars of bread wheat, durum wheat and barley grown in the South of Italy, genetic variation for adaptation to the high temperature and drought stress conditions typical of the Mediterranean environment has been found.The basic data have been extrapolated from 5 years of Italian national network cultivar trials, where 20–30 cultivars were grown in replicated plot trials in 30–50 locations per year, including some where stress strongly affected grain yield.After careful identification of the most representative years and testing sites it was possible to characterise the cultivars on the basis of the grain yield in stress conditions and the Fischer & Maurer (1978) susceptibility index and to find genotypic differences sufficiently repeatable in years.The cultivars giving the best yield under stress associated with low susceptibility indices were in bread wheat: Etruria, Spada, Pandas, Centauro, Oderzo, Costantino and Gladio, in durum wheat: Aldura, Arcangelo, Adamello, Vespro and Capeiti, in barley: Fleuret, Barberousse, Jaidor, Express, Trebbia, Georgie, Dahlia, Criter and Magie.     

Agronomic performance and genetic shifts of genotype mixtures in peanut

Summary Little information is available on the genetic shifts that may take place when seed mixtures of peanut are grown over time. Five peanut lines with distinct seed coat colors, but similar plant types, pod yields, and maturities, were grown in pure stands for four years. A mixture with equal numbers of seed from each of the five lines was grown each year. All mixtures were subsampled and grown in subsequent years without reconstitution. Yield, 100-seed weight, and total sound mature kernel proportions from the mixtures were higher than the pure stand average of the five lines, but yield and total sound mature kernel proportions were lower than the average adjusted for individual line production. The best component line was significantly better than the mixture for pod yield, 100-seed weight, and proportion of sound mature kernels. Intergenotypic competition shifted the mixture composition to increase the proportion of one of the high yielding lines after only two years, while other high yielding lines in the test either maintained or decreased their original proportion. The two lowest yielding lines in pure stand also decreased in proportion over time.Contribution of the Florida Agricultural Experiment Station. Journal Series No. R-00550.     

Adaptation and competition in mixtures of cowpea (Vigna unguiculata (L.) Walp.)

Summary Six pure lines and four mixtures were grown in six environments throughout lowland Papua New Guinea. There were no significant transgressive increases in the grain yield of mixtures above their pure line components, and mixture yields were adequately predicted by the mean of components. In yield stability it was found that the individual buffering of pure lines was of more importance than population buffering, and that the magnitude of population buffering varied with the particular combination of components. Competitive effects in all the mixtures were of the compensating type. Dramatic changes in mixture composition resulting from natural selection precluded their use in local agriculture. The outcome of competition in mixtures was strongly influenced by the growing environment, such that selective index was correlated to general fertility.     

The identification and characteristics of field resistance to lettuce downy mildew (Bremia lactucae Regel)

Summary Eighty-one accessions of three Lactuca species which showed no recognisable race specific resistance to Bremia lactucae when tested in the laboratory as seedlings, exhibited different degrees of susceptibility when exposed to natural field infection. As a group, crisp genotypes had less mildew and a slower rate of disease development than other types of lettuce. Wild forms of Lactuca sativa and Lactuca serriola were particularly susceptible. In a further trial, the low field susceptibility of three lettuce cultivars (Iceberg, Batavia blonde de Paris and Grand Rapids) was confirmed. Disease development on cv. Iceberg was compared to that on the highly susceptible cv. Hilde in experiments where the two cultivars were grown either in close proximity or in isolation. The absolute level of attack on cv. Iceberg depended upon disease pressure and differences between the two types only became apparent approximately 8 wk after sowing.     

The response of field-inoculated wheat cultivars to mixtures of Septoria tritici isolates

Summary Wheat cultivars of diverse genetical background and response to Septoria tritici were inoculated during 2 years in the field with single or mixtures of isolates. Significant reductions in pycnidial coverage were recorded for mixtures of 2 or 5 isolates relative to the virulent isolate ISR8036 under the moderate 1989/1990 epidemic. The interactions between cultivars and all possible combinations among ISR398A1, USR8036 and the 1:1. mixture of the 2 isolates were highly significant. Cultivars exposed to mixtures of isolates expressed differential response in pycnidial coverage compared to the single isolate response. The coverage in the mixtures was significantly less than that of the arithmetic mean between the two isolates. Under the severe 1990/1991 epidemic pycnidial coverage on cultivars inoculated with the mixture of the same 2 isolates did not differ statistically from that of ISR8036, yet, ISR398A1 differed from ISR8036 and the isolate mixture. Losses in 1000-kernel weight for 12 wheat cultivars which were repeated during the 2-trial-years were significantly lower in the isolate mixture relative to that of ISR8036. The suppression of symptoms in isolate mixture relative to the expected expression of the most virulent component may be indicative of differential aggressiveness of isolates regardless of their virulence. The phenomenon may affect screening and selection procedures in breeding for resistance.     

Field variation in a tetraploid progeny derived by selfing a Solanum commersonii (+) S. tuberosum somatic hybrid: A multivariate analysis

A phenotypic diversity index (PDI) was calculated using 10 agronomiccharacteristics recorded in 90 accessions of 2-row spring barleys and 29accessions of 6-row spring barleys grown in the Nordic Region. The PDIranged from 0.0308 to 0.6245 in 2-row barley accessions and from0.0314 to 0.7016 in 6-row barley accessions. The average PDIs were0.2178 and 0.2438 in 2-row and 6-row barley germplasm, which confirmsthat some older cultivars were parents of newer cultivars. The lowest PDIwas between accessions with the same name, which suggest that irrespectiveof their market source, they were the same cultivars. The largest PDI rangesin 2-row barley cultivars within the same country or release decade werecorrelated to the number of accessions in the respective cluster, whichsuggests that phenotypic diversity in this germplasm depends on the numberof cultivars included in the cluster. However, this association was not alwaysobserved in 6-row barley cultivars. The most distinct 2-row cultivars wereArla and Akka from Sweden, whereas Sigur and Tampa from Iceland werethe most distinct 6-row cultivars as determined by both PDI and averagelinkage cluster analysis. This analysis also confirm that the 2-row barleyaccessions Jenny, Triumph, and Vega, which were obtained from twodistinct market sources, and the 6-row barley accessions under the nameAgneta (but from three market sources) were the same. The analysis ofvariance of the PDI indicates that 6-row germplasm may be clusteredaccording to their geographical origin or decade of release, but this was notobserved in 2-row barley germplasm. This research demonstrates theadvantage of PDI to assess variation among breeding pools.     

Conservation of powdery mildew resistance genes in three composite cross populations of barley

Summary Barley powdery mildew was used as a model to evaluate the potential of barley composite cross populations for conservation of disease resistance. The objective was to determine if increases in resistance to powdery mildew could be detected over periods of time in composite cross populations developed in California, where the disease might have had a selective influence on the populations, and the same populations grown in Montana, where no selective influence of powdery mildew was expected. Four isolates of Erysiphe graminis f. sp. hordei were used to monitor the frequencies of plants with specific mildew resistances through early, intermediate and late generations of three composite cross populations (CCII, CCV, CCXII) grown at Davis, California, and Bozeman and Moccasin, Montana. Changes in frequencies of plants resistant to the four isolates were observed between generations in all populations from the three locations. Trends in the frequencies of resistance are discussed in relation to selection pressure applied by E. graminis. It is suggested that associations with gene complexes other than resistance to E. graminis might help to explain the increased resistance observed in these studies.This research was funded in part by U.S. Agency for International Development Contract No. AID/DSAN-C-0024. The authors are grateful to Dr A. L. Kahler for seed of the composite cross populations and to Dr J. G. Moseman for the powdery mildew cultures.Cooperative investigations of the Agricultural Research Service, U.S. Department of Agriculture; the Department of Plant Pathology, Montana State University; and The Montana Agricultural Experiment Station. Journal Series Paper No. 1381 of the Montana Agricultural Experiment Station.     

Involvement of polyamines, diamine oxidase and polyamine oxidase in resistance of barley to Blumeria graminis f. sp. hordei

Polyamine levels and diamine and polyamine oxidase activities have been investigated in the first leaves of barley (Hordeum vulgare L.) in the absence of or following inoculation with conidia of powdery mildew (Blumeria graminis f. sp. hordei). Two cultivars with varying sensitivity to powdery mildew, viz., Chariot (resistant) and Golden Promise (susceptible) were used. The levels of putrescine, spermidine and spermine were found to be higher in the leaves of Chariot than in the leaves of Golden Promise and, with the exception of spermine, were generally higher in both cultivars after inoculation. In inoculated leaves of Chariot, levels of putrescine and spermidine peaked at 9 days and 12 days, respectively. In controls (uninoculated leaves), the activities of these enzymes, and putrescine and spermidine levels also increased but not to the same extent as in inoculated leaves. With Golden Promise, the levels of putrescine and spermidine in the inoculated leaves changed very little over sampling times but were always higher than in the controls. In this cultivar, there was little difference between inoculated leaves and the controls in diamine oxidase activity which reached a maximum value at 9 days post-inoculation. Activity of the bound form of diamine oxidase was low in both the cultivars. Polyamine oxidase was not detected at 3 days after inoculation in either cultivar but activity at fairly low levels was recorded at later times, usually reaching a maximum value at 9 days. The results suggest that polyamine metabolism and diamine oxidase activity in particular may be involved in the mechanism conferring resistance to barley powdery mildew in Chariot.     

Effects of inter-varietal diversity,biotic stresses and environmental productivity on grain yield of spring barley variety mixtures

Varietal seed mixtures tend to increase and stabilize crop yields, yet their application is sparse. Large-scale cultivation of variety mixtures may require a better understanding of how inter-varietal interactions and their interaction with the environment may influence the grain yield of variety mixtures relative to their component varieties. For this purpose, six variety mixtures of spring barley and 14 component varieties were grown in each of 17 trial environments. A total of 28 observed and a priori plant characteristics, including grain yield, disease severity and weed competitiveness, were derived for each component variety in each trial. The relationship between inter-varietal diversity of each characteristic and the mixing effect on grain yield was analysed. Additionally, various types of yield stability were estimated and compared among mixtures and component varieties. One mixture out-yielded all of its component varieties in almost half of the trial environments. Inter-varietal diversity in grain yield potential correlated significantly with mixing effect, as did straw length diversity when weighted with weed pressure. The grain yields of most mixtures were more stable across environments than their component varieties when accounting also for the general response to environmental productivity. Hence, most mixtures adapted slightly better to environmental productivity and were less sensitive to environmental stress than their component varieties. We conclude that the efficacy of variety mixtures may be enhanced by mixing relatively high-yielding varieties differing in responsiveness to environmental productivity.     

Morpho-physiological traits to complement grain yield selection under semi-arid Mediterranean conditions in each of the durum wheat types mediterraneum typicum and syriacum

Summary Some 238 landraces from Algeria and Tunisia representative of the mediterraneum typicum durum wheat type and 265 landraces from Syria and Jordan representing the syriacum type were grown in a semi-arid Mediterranean environment of Northern Syria characterized by moderate drought stress. The germplasm types were compared for mean value, level of variation and relationships with grain yield of various morpho-physiological traits possibly usable for indirect selection of best yielding materials. The syriacum germplasm showed higher yield mainly due to greater earliness of cycle, slightly longer grain filling period, shorter stature, lower early growth vigour and higher drought tolerance expressed by a visual score recorded in another, more stressful environment in the region. It also showed lower variation for all morpho-physiological characters except plant glaucousness, for which it was more variable. The wheat types differed not only for architecture but also for optima of individual morpho-physiological traits required to maximize the yield response in the given environment. Higher yield of syriacum materials was attained at same heading and three to four days delayed maturity with respect to average phenology of three well-adapted control cultivars, and it was favoured by increasing number of kernels per spike, early vigour and drought tolerance. Higher yield of mediterraneum typicum landraces was related to heading and maturity dates approaching those of the control cultivars and to increasing kernel weight, early vigour and drought tolerance. Plant stature hardly affected the yield. Both absent and strong glaucousness could confer a yield advantage in syriacum materials.     

Isolation of wheat mutants with increased resistance to powdery mildew from small induced variant populations

Small variant wheat populations created by induced mutagenesis (n = 69) or adventitious regeneration (n = 66) were intensively screened for an altered response (compared to the parent variety ‘Guardian’) to the causal pathogen of powdery mildew in wheat, Blumeria graminis f. sp. tritici. Intensive field screening following natural infection of replicated plots of wheat lines over two years revealed a total of 13 mutants exhibiting significantly greater resistance than ‘Guardian’: eight from induced mutagenesis (11.6%) of the M2 population and five from adventitious regeneration (7.6%). Complete resistance was identified in two lines, (one (M66) developed following induced mutagenesis, and the other (SC240) by adventitious regeneration). The complete resistance in the induced mutant was stable over two generations and was associated with a high frequency of leaf flecking, and consequently a low grain yield. Resistance in SC240 proved to be unstable; SC240 exhibited complete resistance to powdery mildew in the SC₂ and SC₃ generations, but only 20% of the SC₄ plants were completely resistant, while the remainder were indistinguishable in mildew response to ‘Guardian’. The mildew response of all the SC₅ generation of SC240 was not significantly different from ‘Guardian’. Yield analysis of the thirteen mutants with increased resistance in the presence of powdery mildew indicated that eleven exhibitedgrain yields at least as high as that of ‘Guardian’, while the mutant M19 exhibited a yield significantly higher than that of ‘Guardian’. This revised version was published online in July 2006 with corrections to the Cover Date.     

The role of resistance breeding in the intergrated control of downy mildew (Bremia lactucae) in protected lettuce

Summary Over the last 30 years, six resistance alleles (Dm2, Dm3, Dm6, Dm7, Dm11 andDm16) located in two linkage groups, have contributed to the control of downy mildew in lettuce crops grown under protection (glass or polythene) in northern Europe. More recently, an as yet genetically uncharacterised resistance factor, R18, has also begun to assume importance. The occurrence of the various combinations of these resistance alleles that exist in commercial cultivars has been dictated by the pathotypes ofBremia lactucae used in their selection but also restricted by linkage in repulsion. In the UK, a pathotype ofB. lactucae insensitive to phenylamide fungicides, such as metalaxyl, emerged in 1978 and became prevalent throughout lettuce production areas in subsequent years. The specific virulence of this pathotype was identical to the previously described phenylamide sensitive pathotype NL10 and cultivars carryingDm11, Dm16 or R18 were resistant. Consequently, an integrated control strategy based on the utilisation of metalaxyl on cultivars carryingDm11 provided effective control in UK until 1987 when a new phenylamide insensitive pathotype began to cause problems. The specific virulence of this second pathotype, which was first reported in the Netherlands and France, was identical to the previously described phenylamide sensitive pathotype NL15. Cultivars carryingDm6, Dm16 or R18, but notDm11, were resistant to NL15; consequently an appropriate change in the cultivar recommendations for use in the integrated control strategy was successfully promulgated. It is predicted that variations of this integrated control strategy involving the use of appropriately selectedDm gene combinations may prove effective for some time. This prediction is based on studies of the status of the avirulence loci in the two phenylamide insensitive pathotypes and of the specific virulence characteristics of phenylamide sensitive components of the pathogen population.     

Molecular characterization of genetic diversity in European germplasm of perennial ryegrass

Summary Perennial ryegrass (Lolium perenne L.) is the most important grass species for temperate grassland agriculture. The level and distribution of genetic variation in gene bank ecotype collections is still largely unknown but of great interest for the planning of breeding programs. The objectives of this study were to (i) assess the molecular diversity of Polish ecotypes of perennial ryegrass, and (ii) compare the relationship between this group and German ecotypes and European cultivars investigated previously. A total number of 166 polymorphic marker bands were detected among the 171 individual plants of the 9 Polish ecotypes. In a joint analysis with 9 Polish and 22 German ecotypes, and 22 European cultivars 172 polymorphic RAPD markers could be found. Genetic distance among the Polish ecotypes ranged from 0.31 to 0.51, while for all 53 populations a broader range was detected (0.25–0.67). An analysis of molecular variance (AMOVA) revealed a much larger variation within populations (71%) than among them (29%). The Polish ecotypes contained the highest within population variation (74%). The largest among group difference (15%) was found between the Polish ecotypes versus all other accessions. We conclude that the Polish ecotypes represent a valuable genetic resource for enlarging the genetic variation in the West European germplasm pool of perennial ryegrass.     

Variation in some agronomically important characters in a germplasm collection of beet (Beta vulgaris L.)

Summary A geographically representative selection of germplasm of Beta vulgaris, section Beta has been assessed for characteristics important in sugarbeet breeding, including downy mildew resistance, resistance to aphid colonisation and infection by the beet virus yellow complex. The occurrence of maintainer lines for cytoplasmic male-steriles was also investigated. Desirable qualities were found in some accessions, including nothern European wild vulgaris ssp. maritima and some old multigerm cultivars of fodder beets.     

Screening immature bulbs of daffodil (Narcissus L.) crosses for resistance to basal rot disease caused by Fusarium oxysporum f. sp. narcissi

Summary Immature bulbs were used in two screening trials to identify daffodil genotypes resistant to basal rot disease caused by Fusarium oxysporum f. sp. narcissi. Progenies, having at least one parent with some resistance, were grown in compost infested with chlamydospores of the fungal pathogen, and survivors were recovered at final harvest. Survival rates were very low when one-year old bulbs were used; the greater survival of two-and three-year old bulbs indicates that resistance increased with bulb age. Selection pressure was influenced by inoculum concentration in the compost. Continuous variation of percentage bulb survival between progenies suggests a polygenic mode of inheritance; there was no evidence of maternal inheritance. Parental general combining ability (GCA) was highly significant and accounted for much of the difference in bulb survival between progenies, but non-additive parental effects were also apparent. GCA rankings in the two trials were similar and reflected the resistance of parental cultivars in the field. A three-dimensional graphing procedure was devised to depict goodness of fit of progeny data to the additive model.     

Yield and Persistence from Mixtures of a Winter Hardy and a Verticillium Wilt Resistant Lucerne Cultivar

In British Columbia, Lucerne stand persistence is reduced by inadequate disease resistance, winter hardiness, or both. We evaluated the possibility that persistence could be improved by sowing mixtures of winter hardy and verticillium wilt (VW) resistant cultivars. Our hypothesis was that dry matter yield of mixtures of two Lucerne cultivars could be calculated simply from the proportion of each cultivar in the mixture and their yield in pure stand. A winter hardy cultivar, Beaver, was grown in mixtures with each of three VW (caused by Verticillium albo-atrum Reinke & Berthier) resistant cultivars (WL316, Sparta, or Barrier) in proportions of 0, 0.25, 0.5, 0.75, and 1.0. The yield of a mixture of two cultivars could be calculated from the proportion of each cultivar in the seed mixture and their yields in monoculture. In the absence of a winter where severe injury to Lucerne stands occurred, the VW-resistant cultivar Barrier yielded about 2 t ha⁻¹ year⁻¹ more than the more winter hardy Beaver.     

Spanish barley landraces outperform modern cultivars at low‐productivity sites

Barley landraces from the western Mediterranean area have not been thoroughly exploited by modern breeding. This study aims at assessing the agronomic value of a core collection of lines derived from landraces of Spanish origin and to compare them with sets of successful old and modern cultivars. The agronomic performance of a set of 175 barley genotypes, comprising 159 landrace‐derived lines and 26 cultivars, was evaluated in a series of 10 field trials, carried out over 3 years and several locations. The most relevant trait of the landraces was higher grain yield at low production sites than cultivars, which may be related with better ability to fill the grain under stressful conditions. On the other hand, lateness, excessive plant height and lodging were negative traits frequently found in the landraces. Large genotype‐by‐environment interaction (GEI) for grain yield was detected, related partly with differences between germplasm groups, probably indicating local adaptation. GEI was also associated with the interaction of heading time and powdery mildew resistance with temperature.