Nonhost plant species as donors for resistance to pathogens with narrow host range. II. Concepts and evidence on the genetic basis of nonhost resistance

Summary It is commonly argued that wild plant species with a very high level of resistance to a micro-organism which is pathogenic on a crop species would be useful donors of resistance. Whether this principle is true if the wild species is a distinct nonhost, remains to be proved. It is assumed that nonhost resistance, if not based on avoidance, rests on a poor adaptation of the micro-organism to general defence mechanisms in the plant. No basic compatibility is achieved. Major-genic hypersensitive host resistance would, according to the concept, be superimposed on basic compatibility. The defence reaction is switched on if a resistance allele recognizes a specific avirulence factor produced by the pathogen. The frequency of the resistance allele would depend on the selection pressure exerted by the pathogen population.In contrast to these concepts, there is evidence that nonhost resistance, at least to formae speciales, rests on an extremely high allele frequency of effective major genes for resistance, rather than on a complex of genes that play part in a general defence system. Especially the high allele frequency and the durability of the nonhost resistance are in conflict to what would be expected if the concepts hold true. It is concluded that part of the theoretical concepts needs to be revised or extended to reconcile them with the evidence. A major-genic base for nonhost resistance would be very interesting from the breeder's point of view, because transfer of such a resistance to a crop would be less hard to accomplish than that of more complex inherited defence systems.     

Resistance to Corynebacterium michiganense in tomato genotypes

Summary The highest level of resistance to Corynebacterium michiganense was found in Irat L3 and Okitsu Sozai 1–20. Resistance was partial and symptomless plants proved to be carriers of the disease. The resistance is at least partially based on a diminished multiplication of the pathogen in the host plant.     

Resistance to plant viruses: Obtaining genes by non-conventional approaches

Summary Plant viruses cause considerable losses to crops and none of the three current approaches to disease control are likely to give the long-term answer. The new concept of non-conventional protection, in which the expression of a viral or virus-related sequence in the plant genome interferes with the virus infection cycle, holds considerable promise for designing new resistance or protection genes. A series of targets in the viral genome is identified and a range of mechanisms for attacking those targets is discussed. There are several current systems which have been proved to give protection, at least to a certain extent, and many systems which are being researched upon for the future. Some of these are described to give a picture of the current situation and of the thinking for the future. The problems of field deployment of the transgenic plants are discussed, especially those associated with the risk to the environment. Various questions which molecular biologists and plant breeders will have to consider include what are desirable characters to have in protection genes and how these new genes should be deployed.     

Wheat x pearl millet hybridization: consequence and potential

Summary Eight grain pearl millet (2n=14) accessions were crossed as male to hexaploid spring wheat cv. Fukuho (2n=6x=42). An average of 80% wheat pistils showed pearl millet pollen tube entry in the ovules, compared to 56% in wheat x maize cv. Seneca 60 cross. Of the 15 embryos, obtained through in vitro immature seed culture from wheat x pearl millet crosses, 3 plantlets were produced and grown to maturity. These three were of the somatic chromosome constitution 2n=42, 21 and 22, respectively. Haploid wheat plant (2n=21) apparently originated from pearl millet chromosome elimination during embryogenesis. The 22 chromosome plant had retained a single pearl millet chromosome at tillering stage, but this chromosome was eliminated from pollen mother cells prior to and also during gamete formation. The significance and potential uses of this wide cross is discussed.     

Resistance to potato virus Y (pathotype 1–2) in Capsicum annuum and Capsicum chinense is controlled by two independent major genes

Summary The genetic basis of the resistance to a pepper strain of potato virus Y (PVY pathotype 1–2) was investigated in two lines: Capsicum annuum Criollo de Morellos 334 and Capsicum chinense PI 159236. F1, backcrosses, and F2 populations were produced using C. annuum Magda as the susceptible parent. Segregation ratios indicated that the resistance in C. annuum Criollo de Morellos 334 is governed by a nuclear, single dominant gene with a very high but incomplete penetrance. On the other hand, the resistant response of C. chinense PI 159236 is associated with the presence of a major recessive gene with an apparently complete penetrance. The failure in recovering or detecting PVY from/in the inoculated leaves of both resistant genotypes is strongly suggestive that both genes are acting by preventing the multiplication of the virus in the leaf tissue. Both mechanisms of resistance closely resemble either an operational immunity or a single-cell hypersensitive reaction to PVY pathotype 1–2. The name Ry 1–2 is suggested for the resistance gene from Criollo de Morellos 334. The gene for resistance to PVY pathotype 1–2 identified in C. chinense PI 159236 may be the same designated et ^c1, which is mentioned to be allelic to the et ^av locus. The phenotypic expression of the et ^av gene was characterized in C. annuum Avelar as a tolerant but not immune or hypersensitive reaction against some potyviruses in Florida. These data suggest that a critical reexamination of the allelic relationships at the locus et should be performed.     

Expression of late maturity α-amylase in wheat containing gibberellic acid insensitivity genes

Summary Two wheat cultivars, Spica and Lerma 52, which consistently produce high levels of -amylase during the later stages of grain development (late maturity -amylase), were crossed with a set of four near-isogenic lines carrying the tall (rht) allele or one of the dwarfing genes Rht1, Rht2 or Rht3 (GA-insensitive alleles). The F₁ and F₂ populations were developed and analysed for grain -amylase and plant height. The Rht3 gene exhibited the strongest influence on plant height and strongly inhibited new -amylase synthesis during the later part of grain ripening. By comparison, Rht1 and Rht2 had a less pronounced effect but still significantly reduced the expression of late maturity -amylase. These observations suggest that gibberellic acid is involved either directly or indirectly in this phenomenon. The implications of the effect of dwarfing genes on expression of late maturity -amylase are discussed in relation to cultivar improvement and to the identification and control of high -amylase germplasm.     

The concept of breeding for uniform and differential resistance and their integration

Summary It has been reported why resistance is needed in crop plants, how it has been searched for, and how long the resistance could be maintained in the host varieties. Uniform and differential resistance have been discussed and integrated within one genetic system. The possible evolution of both polygenes and major genes for resistance is discussed. Polygenes probably are older and produce the materials necessary for a host to show resistance, major genes acting as switching genes. Breeding for disease resistance has been briefly debated in connection with the discussion on host resistance.     

The black currant leaf curling midge,Dasyneura tetensi Rübs.; its host range,and the inheritance of resistance

Summary Established Western European black currant cultivars are susceptible to Dasyneura tetensi but the Scandinavian Sunderbyn II, Kangosfors and Hedda, and 10 out of 33 Russian cultivars showed very slight or no leaf symptoms of midge attack in the field.Of species within the black currant Section Eucoreosma, Ribes dikuscha, R. pauciflorum (probably=syn R. ussuriense), and R. americanum were not attacked, but an accession of R. bracteosum proved very susceptible. Species within other sections of the genus were not attacked.A strong dominant resistance gene in R. dikuscha, designated Dt, appeared to be linked with (a) gene(s) controlling resistance to American gooseberry mildew, Sphaerotheca mors-uvae. Resistance in Sunderbyn II and, probably, in R. pauciflorumwas oligogenic and dominant.     

Chemically induced sterility in wheat for hybrid seed production

Summary The gametocidal effects of RH-531, RH-532, and RH-2956 were studied with three rates and times of foliar application on two spring wheat (Triticum aestivum L.) varieties, Anza and Yecora 70. RH-531 and RH-532 applied at or before booting stage caused rather large reduction in plant height while RH-2956 had a small effect on height. Application of any of the three chemicals on Anza at meiotic stages reduced self-pollination (SP) fertility by 97–99%. If 90% SP fertility reduction is acceptable, the application time could vary from premeiosis to postmeiosis for Anza. Yecora 70 was less sensitive than Anza to gametocide treatments. The lowest SP fertility obtained in Yecora 70 with RH-531, RH-532, and RH-2956 was by treatment at booting (4 kg/ha), at premeiosis (2 kg/ha), and heading (8 kg/ha), and the corresponding SP fertility reductions were 74, 89, and 71%. Pollen stainability-determined by aceto-carmine, IKI, and Alexander's stains-was not affected by gametocide treatment and was not useful in evaluating sterility induced by these chemicals. For the RH-531-treated Anza and Yecora 70 and the RH-532-treated Anza, more than 94% of the seeds produced by open-pollination on treated plants were from self-or sib-pollination as determined by progeny testing. However, RH-532-treated Yecora 70 and RH-2956-treated Anza and Yecora 70 did result in more crossed seeds, with a maximum of 20% outcrossing. It was believed that spike compactness induced by the chemicals and the related poor flower opening were the major limitations for outcrossing. Artificial pollinations made to the test varieties showed that RH-531 induced both male and female sterility and was undesirable for practical use. RH-532, however, could be applied at the booting stage to avoid severe female sterility, and RH-2956 clearly was the best of the three chemicals, with the female fertility very close to those of the untreated checks of both Anza and Yecora 70. Of the three chemicals, RH-2956 gave the highest percentage of hybrid plants in the progeny of hand-or open-pollinated spikes. Its practical use for hybrid seed production, however, will depend largely on improvement of methods to increase cross pollination in wheat.     

Congruity backcrossing as a means of creating genetic variability in self pollinated crops: seed morphology of Phaseolus vulgaris L. and P. acutifolius A. Gray hybrids

Summary Seeds of early generations of three reciprocal congruity-backcross (CBC) pedigrees, developed by backcrossing Phaseolus vulgaris-P. acutifolius hybrids to each of the parent species in alternate generations, exhibited a preponderance of traits (size, shape, color, and pattern) of the cytoplasmic parent. The large size of Red Cloud (V₁), the P. vulgaris parent common to all of the pedigrees, dominated pedigrees with V₁ as the cytoplasmic parent, while the small size and rounded or square shapes of the tepary parents, wild P. acutifolius var. acutifolius PI 263590 or G400445 (A₁₉), wild P. acutifolius var. latifolius PI 406622 (A₁₀), or cultivated P. acutifolius var. latifolius Serowi PI 319443 (A₉), were the majority phenotypes when P. acutifolius was the cytoplasmic parent. Continuing through the second cycle of CBC, that is the second backcross with each of the parent species or the fourth backcross, began an amelioration of the apparent cytoplasmic effect on gene expression, as reciprocal pedigrees became more alike, usually with intermediate expression of parental traits or the appearance of new traits. The large seed size of V₁ was recovered in hybrids with P. acutifolius cytoplasm and the kidney shape of V₁ became rare in hybrids with P. vulgaris cytoplasm. Although the tepary-bean parents represented two subspecies and both cultivated and wild P. acutifolius, the three sets of reciprocal-hybrid pedigrees with P. vulgaris Red Cloud are surprisingly similar. It may be that the exotic parent used to develop a CBC pedigree should be selected more for combining ability in the interspecific cross than for specific economic traits. While the number of generations (six or more) required to produce fertile, intermediate CBC hybrids (that did not require embryo rescue) may preclude routine use of this method by practical plant breeders, the crossability of advanced hybrids with both parental species and the amount of variability apparent in advanced-hybrids progenies suggests that CBC would be valuable for maintaining exotic germplasm in immediately useful forms.     

Evaluation of maize varieties for their potential as breeding populations

Summary A 7-variety diallel of synthetic varieties that have not had selection for yield improvement and a 10-variety diallel of synthetic varieties that either resulted from recurrent selection for yield improvement or from the introgression of exotic germ plasm into adapted varieties were evaluated in six environments for yield and five other agronomic traits. The specific objective was to determine the relative potential of the synthetic varieties as germ plasm sources for breeding programs.Recurrent selection improved population yield in BSSS CO, BSK CO, and Alph CO in the varieties and also in crosses with other varieties. BSSS(R)C6 and BSSS(HT)C7 yielded 11.4% and 16.1% more than BSSS CO, and the variety crosses of BSSS(R)C6 and BSSS(HT)C7 yielded 21.6% and 22.0% more than the variety crosses of BSSS CO. Recurrent selection also improved stalk-lodging resistance for BSSS(R)C6 and BSSS(HT)C7, but root lodging for BSSS(HT)C7 was significantly higher. BS15 had 61.0% higher yields than Alph CO, with 9.3% less stalk lodging. Average heterosis of the 10-variety diallel was 1120 kg/ha (19.6%) and ranged from 800 kg/ha (13.7%) for BSK(S)C5 to 1770 kg/ha (39.4%) for BS12(HI)C5. The heterosis expressed among individual variety crosses ranged from 4.3% for BSK(S)C5 x BSTE(FR)C1 to 37.6% for BSSS(R)C6 x BS12(HI)C5, which approached the yield of the two highest yielding single-cross checks.In the 7-variety diallel of unselected varieties, BSTE CO and Late Rootworm Synthetic were superior as varieties and in crosses for yield and stalk quality. Average heterosis for the 7-variety diallel was 950 kg/ha (18.5%). Stalk lodging was important in the performance of varieties with poor stalk quality.Although recurrent selection improved yield, development of superior breeding populations also will require selection for general agronomic performance. The performance of BS16 and the yields of BSTL and Teozea and their crosses suggest that exotic germ plasm should receive greater attention.Contribution from the Agricultural Research Service, U.S. Department of Agriculture, and Journal Paper No. J-8096 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa 50010. Project No. 1897.     

Interplot interference and the assessment of barley cultivars for partial resistance to leaf rust,Puccinia hordei

Summary The barley cultivars Akka, highly susceptible, and Vada, partially resistant to barley leaf rust, Puccinia hordei, were evaluated for the amount of leaf rust in five experimental field plot situations over three successive years. The field plot situations were: A) plots well isolated from each other by distance and non-leaf rust contributing host plants; B) adjacent plots of 4×41/2 m (18 rows); C) adjacent plots of 4×11/2 m (6 rows); D) adjacent plots of 4×1/4 m (1 row); E) adjacent plots of only one plant (cultivar mixtures).The sporulating leaf area of each plot was measured from samples of 20 tillers taken at random from each plot. In each year the difference in sporulating area between Akka and Vada was large to very large in the absence of interplot interference in the isolated plots, ranging from 150 to 2100 times. In the adjacent plots the partial resistance of Vada was greatly underestimated, 5 to 16 times in the situation B, 14 to 30 times in C, and 75 to 130 times in D and E.Testing lines or cultivars in adjacent plots is the standard procedure in use in breeding programs and in tests of cultivars for their agricultural value. To avoid such under estimation the following procedure is suggested. A few cultivars representing the known range of partial resistance and whose level of partial resistance is well known are evaluated together with the lines and cultivars whose partial resistance has to be assessed. This is demonstrated with a number of cultivars of which resistance values are know from the recommended variety lists for England and Wales. Cultivars have been assessed in Wageningen over four years together with the check cultivars Akka, Sultan, Julia and Vada representing the range of partial resistance with values (on a 1 to 10 scale) of 1, 3–4, 7 and 8 respectively, based on isolated plots experiments.     

Genetics of resistance to five strains of turnip mosaic virus in Chinese cabbage

Summary Inheritance of resistance to turnip mosaic virus (TuMV) strains C1, C2, C3, C4, and C5 in Chinese cabbage (Brassica campestris subsp. pekinensis) was evaluated using monoclonal antibodies. Crosses were made between a resistant line, 0–2, and four susceptible line. Seoul (SE),SSD31 (SS), Cheongbang (CH), and Yaki 1 ho (YA), to determine the inheritance of resistance of 0–2 in different genetic backgrounds. Resistance to TuMV was controlled by a single dominant gene or double dominant genes depending on the strain and cross. The resistance genes of 0–2 were modified by susceptible parents such that a single dominant gene was involved in the SS×0–2 combination, but double dominant genes in the SE×0–2 against TuMV-C3 or TuMV-C5. ELISA tests using inoculated and noninoculated leaves in the same plant suggested that the dominant resistance genes inhibit virus movement rather than virus multiplication.     

Relationship between allelic variation of Glu-1 and Gli-1/Glu-3 prolamin loci and gluten strength in hexaploid wheat

Summary Allelic variation of prolamin loci was examined in the F₂ from crosses between the hexaploid wheat varieties: Cajeme 71, Yécora 70, Ablaca, Anza, Pané 247 and Axona. Different allelic blocks for gliadins and LMW glutenin subunits were determined in Gli-1, Gli-2 and Glu-3 loci. A percentage of recombination of 1.5 ± 0.3 was determined between Gli-A1 and Glu-3 in the F₂ progeny of Yécora 70 x Axona. A significant positive association was found between gluten strength, measured by SDS-sedimentation volume, and the prolamins coded by Anza Gli-D1/Glu-D3 loci and Yécora 70 Gli-A1/Glu-A3 loci. Interactions between non homeologous loci Glu-1 and Gli-1/Glu-3 were also found.     

Life history studies of the leafminer Liriomyza trifolii on susceptible and resistant cultivars of Dendranthema grandiflora

Summary The feeding, fecundity, mortality and rate of development of Liriomyza trifolii (Burgess) on 12 cultivars of Dendranthema grandiflora (Tzvel.) (Syn. Chrysanthemum morifolium (Ramat.)) were determined in no choice experiments. Number of feeding punctures, number of eggs laid and total number of larvae varied between cultivars but these characters were not strongly associated with leafminer population increase. Good indicators of host plant resistance were larval mortality, number of pupae and duration of the larval period. Of these three, larval mortality which could be as high as 98% was most suitable.Based on life history data it was predicted that Liriomyza trifolii cannot survive on the resistant cultivar Penny Lane. This prediction was verified experimentally.     

Wide adaptation: How wide?

Summary Breeding programs aim at improving crop production either within given macroenvironments (for example rainfed vs. irrigated) or in a wide range of growing conditions. The merit of either strategy depends on the range of testing environments and on the definition of stress environment. When environments with average yields of 3–4 t/ha are defined as stress environments, selection for stress conditions can be successfully conducted under optimum conditions. However, when the stress environment has a much lower yield potential (0.5–2.0 t/ha), direct selection in the target environment is the most efficient strategy.A review of data on widely adapted genotypes supports this conclusion.The relative magnitude of heritability in stressed and non-stressed environments is not sufficient to choose the optimum environment for selection, because phenotypic differences can be of opposite sign in different environments.The role of constitutive characters as analytical tools in breeding for yield stability in stress environments is briefly discussed.     

The relationship between powdery mildew (Sphaerotheca fuliginea) resistance and leaf chlorosis sensitivity in cucumber (Cucumis sativus) studied in single seed descent lines

Summary The genetic relation between powdery mildew resistance and sensitivity for leaf chlorosis of glasshouse cucumber was investigated. The powdery mildew resistant, leaf chlorosis sensitive hybrid variety Profito was crossed with the powdery mildew susceptible, non chlorosis sensitive hybrid variety Corona. Forty four F₆ inbred lines of this cross, produced by single seed descent, were tested for powdery mildew resistance (PMR) and leaf chlorosis sensitivity (LCS). PMR and LCS were positively correlated (r=0.59). One or more of the PMR genes probably causes LCS as a pleiotropic effect or is closely linked to LCS, but also other factors, not genetically linked to PMR can cause LCS. Five F₆ lines combined a significantly higher level of PMR than Corona with a significantly lower LCS than Profito. Three of these lines could hardly be distinguished from Corona for the level of LCS.Abbreviations LCS leaf chlorosis sensitivity - PMR powdery mildew resistance - SSD Single Seed Descent     

Effect of genetic blends of dry beans (Phaseolus vulgaris) of different Plant architecture on apothecia production of Sclerotinia sclerotiorum and white mold infection

Summary The determinate Phaseolus vulgaris L. cvs Dark Red Kidney Charlevoix, and near-isogenic Great Northern Nebraska 1 were grown in blends with the indeterminate GN Nebraska 1 in white mold (Sclerotinia sclerotiorum) disease field nurseries. A critical difference between Charlevoix and indeterminate GN Nebraska 1 is that the latter has a greater leaf area closer to the soil surface, and this is associated with increased numbers of apothecia beneath the canopy and higher disease severity. White mold infection and apothecia number/m² beneath the canopy of the blends containing Charlevoix were significantly reduced in comparison with the severely infected, homogeneous, indeterminate GN Nebraska 1. A reduction of white mold infection for the indeterminate GN Nebraska 1 was not observed in blends grown under severe or moderate white mold incidence, but did occur under slight incidence in blends containing 65% and 75% Charlevoix.No significant difference for seed yield occurred between the blends and homogeneous cultivars planted in four experiments under severe, moderate, slight and zero white mold incidence, respectively, except in Experiment 1 under moderate white mold incidence, the blend of 50% indeterminate GN Nebraska 1 and 50% Charlevoix exceeded the yield of the indeterminate GN Nebraska 1. Mean weight of white (GN Nebraska 1) and red seed (Charlevoix) increased and decreased, respectively, in some blends due to the more vigorous growth of the indeterminate GN Nebraska 1.Published as Paper No. 5341, Journal Series, Nebraska Agric. Exp.t Station. Research was conducted un under Project 20–3.     

Inheritance of resistance to powdery mildew (Erysiphe graminis f. sp. hordei) in eleven primitive barley varieties

The inheritance of the powdery mildew resistance of eleven primitive barley varieties was investigated. crosses with varieties with identified resistance genes revealed that at least three different genes are present in a group of six genotypes from Greece (Hor736, Hor847, Hor878, Hor1159, Hor1379 and Hor1873) and one from Turkey (Hor1188). One dominant gene is common to these seve genotypes and in the Greek accessions a second gene is present. The expression of this second gene was respectively dominant in the crosses with Hor736 and Hor1159, recessive in the crosses with Hor878 and Hor1873, and depended on the employed powdery mildew isolate in the crosses with Hor847 and Hor1379. The detected genes differ from the resistance genes of the crossing partners, viz. Ml-al2, Ml-(Ab), Ml-(CP) and Ml-(1402). The accessions from China (Hor824 and Hor4021) each possess two resistance genes which differ from Ml-a12, Ml-(La) and Ml-(CP). The major gene is common to both accessions and is either identical, allelic or closely linked to a gene in the variety Nigrate. The Columbian accession Hor1894 possesses one resistance gene linked to a resistance gene in Nigrate but this gene differs from the Ml-a locus. The variety Palestine (Hor3997) possesses two resistance genes of which one is allelic or closely linked to Ml-(at).     

The importance of Endosperm Balance Number in potato breeding and the evolution of tuber-bearing Solanum species

Summary Endosperm failure is considered the primary reason for the lack of success in intra-and interspecific crosses. The Endosperm Balance Number (EBN) hypothesis is a unifying concept for predicting endosperm function in intraspecific, interploidy, and interspecific crosses. In the EBN system, every species has an effective ploidy (EBN), which must be in a 2:1 maternal to paternal ratio in the endosperm for crosses to succeed. The knowledge of EBN is very useful in the transfer of genes from exotic germplasm, and in the development of new breeding schemes in potato. The paper describes the strategies for introducing 2x(1EBN), 2x(2EBN), 4x(2EBN) and 6x(4EBN) germplasm into the cultivated 4x(4EBN) potato gene pool. A new methodology for producing 4x(4EBN) and 2x(2EBN) chromosome addition lines is also discussed. EBN has evolutionary importance in the origin of tuber-bearing Solanums. The role of the EBN in the origin of diploid and polyploid potato species, and as a barrier for hybridization and speciation of sympatric species within the same ploidy level is demonstrated. The origin of 3x and 5x cultivated tuber-bearing Solanums may also be explained using the EBN concept. EBN has been reported to exist in other plant species: alfalfa, beans, blueberries, rice, soybeans, squashes, tomato, forage legumes, grasses, ornamentals and Datura stramonium. This indicates that EBN may have broad application and could be useful for germplasm transfer and breeding in other crop species.