The integrated concept of disease resistance: A new view including horizontal and vertical resistance in plants

Summary Horizontal, uniform, race-non-specific or stable resistance can be discerned according to Van der Plank, from vertical, differential, race-specific or unstable resistance by a test in which a number of host genotypes (cultivars or clones) are tested against a number of pathogen genetypes traces of isolatest. If the total non-environmental variance in levels of resistance is due to main effects only differences between cultivars and differences between isolates) the resistance and the pathogen many (in the broad sense) are horizontal in nature. Vertical resistance and pathogenicity are characterized by the interaction between host and pathogen showing up as a variance compenent in this test due to interaction between cultivars and isolates.A host and pathogen model was made in which resistance and pathogenicity are governed by live polygenic loci. Within the host the resistance genes show additivity. Two models were investigated in model I resistance and pathogenicity genes operate in an additive way as envisaged by Van der Plank in his horizontal resistance. Model II is characterized by a gene-for-gene action between the polygenes of the host and those of the pathogen.The cultivar isolate test in model I showed only main effect variance. Surprisingly, the variance in model II was also largely due to main effects. The contribution of the interaction to the variance uppeared so small, that it would be difficult to discern it from a normal error variance.So-called horizontal resistance can therefore be explained by a polygenic resistance, where the individual genes are vertical and operating on a gene-for-gene basis with virulence genes in the pathogen. The data reported so far support the idea that model II rather than model I is the realistic one.The two models also revealed that populations with a polygenic resistance based on the gene-for-gene action have an increased level of resistance compared with the addition model, while its stability as far as mutability of the pathogen is concerned, is higher compared to those with an additive gene action. Mathematical studies of Mode too support the gene-for-gene concept.The operation of all resistance and virulence genes in a natural population is therefore seen as one integrated system. All genes for true resistance in the host population, whether they are major or minor genes are considered to interact in a gene-for-gene way with virulence genes either major or minor, in the pathogen population.The models revealed other important aspects. Populations with a polygenic resistance based on a gene-for-gene action have an increased level of resistance compared to populations following the addition model. The stability, as far as mutability of the pathogen is concerned, is higher in the interaction model than in the addition model.The effect of a resistance gene on the level of resistance of the population consists of its effect on a single plant times its gene frequency in the population. Due to the adaptive forces in both the host and the pathogen population and the gene-for-gene nature of the gene action an equilibrium develops that allows all resistance genes to remain effective although their corresponding virulence genes are present. The frequencies of the resistance and virulence genes are such that the effective frequencies of resistance genes tend to be negatively related to the magnitude of the gene effect. This explains why major genes often occur at low frequencies, while minor genes appear to be frequent. It is in this way that the host and the pathogen, both as extremely variable and vigorous populations, can co-exist.Horizontal and vertical resistance as meant by Van der Plank therefore do not represent different kinds of resistances, they represent merely polygenic and oligogenic resistances resp. In both situations the individual host genes interact specifically with virulence genes in the pathogen. Van der Plank's test for horizontal resistance appears to be a simple and sound way to test for polygenic inheritance of resistance.The practical considerations have been discussed. The agro-ecosystems should be made as diverse as possible. Multilines, polygenic resistance, tolerance, gene deployment and other measures should be employed, if possible in combination.     

PLPKI: A novel serine protease inhibitor as a potential biochemical marker involved in horizontal resistance to Phytophthora infestans

Potato leaves infected with Phytophthora infestans produced a serine protease inhibitor (PLPKI) with specificity for microbial proteases. Sequencing of the first twenty residues at the NH₂‐terminus of the mature PLPKI polypeptide demonstrated that PLPKI is a novel member of the potato protease inhibitor I family. PLPKI inhibited the activity of extracellular proteases produced by two pathogens of potato, P. infestans and Rhizoctonia solani, but was inactive against proteases secreted into the culture media by the binucleate Rhizoctonia N2, a non‐pathogenic fungus for potato. Western blot analysis showed a positive correlation between the levels of PLPKI and the degree of horizontal resistance, showing its highest accumulation in clone OKA 5632.11, which has been described as highly resistant. This correlation, together with the ability of PLPKI to completely abolish the secreted serine protease activity of P. infestans, suggests that PLPKI may have an active role in protecting potato plants from this pathogenic oomycete and that it could be used as a suitable biochemical marker to help breeders in the selection of cultivars with high degree of horizontal resistance.     

Practical breeding for durable resistance to rust diseases in self-pollinating cereals

Summary The rust pathogens of cereals exist as populations of races that differ in their ability to attack various varieties. Varieties that are resistant when first released often become susceptible later due to the spread of previously undetected races but the time taken for this to occur in very variable. It often occurs so rapidly as to curtail the commercial use of otherwise satisfactory varieties.Some varieties, however, are widely grown for many years and remain adequately resistant to the prevalent rust diseases. They may aptly be described as having durable resistance. This durable or long-lasting resistance can be detected without any assumptions about, or detailed knowledge of, whether durability depends on any particular mechanisms of resistance, on various degrees of racespecificity or on many or few genes. Cappelle-Desprez is given as an example of a wheat variety with durable resistance to yellow rust.The most powerful test for the detection of durable resistance occurs when a variety is widely grown commercially for several years. A much weaker test is obtained by growing varieties in small disease nursery plots even when the test is repeated for several years. Usually, resistance which is durable is also partial or incomplete. Often, however, partial resistance of wheat to yellow rust has not been durable. Thus the observation that resistance is partial is not, of itself, a satisfactory criterion for the detection of durable resistance.It is suggested that the most obvious sources of durable resistance for use in breeding programmes are varieties which have been widely grown and have displayed this character. The transfer of such resistance during breeding may be achieved if the creation or incorporation of higher levels of resistance that have not been tested for durability is avoided. It should then be possible to derive resistance from the durably. resistant parent. Methods of achieving this are discussed.     

Diversity of heat-stable genotype specific resistance to Meloidogyne in Maranon races of Lycopersicon peruvianum complex

Accessions of Lycopersicon peruvianum complex and their F₁progenies were screened for genotype specific resistance to Mi-1-avirulent M. incognita and M. javanica biotypes at25 ^°C and at 32 ^°C (a temperature at which Mi-1resistance is not expressed), and to Mi-1-virulent M.incognita at 25 ^°C. All entries of the L. peruvianumChotano-humifusum race accessions LA2157 and LA2334 were resistantto Mi-1-avirulent biotype at 25 ^°C and at 32 ^°C,indicating that the accessions are homozygous for the heat-stableresistance. The L. peruvianum Maranon race accessions LA1626,LA1708, LA2172, LA2185, LA2326 and LA2328 segregated for heat-stableresistance to Mi-1-avirulent biotype. The F₁ progeny tested ofLA392 × LA2157, LA2334 × LA2157, LA2328 × LA2326,LA2328 × LA2185, LA1708 × LA2328 andLA1626 × LA2172 were resistant to Mi-1-avirulent biotype at32 ^°C. There were differences in the segregating accessions andF₁ hybrids for expression of heat-unstable and heat-stable resistanceto Mi-1-avirulent Meloidogyne spp. The L. peruvianumLA392 and LA2163 and L. chilense LA1968, LA1972, LA2404, LA2405,LA2406, LA2748, LA2930, and the L. peruvianum × L.chilense hybrids were homozygous susceptible with all entries testedsusceptible at 32 ^°C. Cuttings of these L. peruvianumaccessions and their F₁ progenies were susceptible to Mi-1-virulent M. incognita biotype at 25 ^°C.     

Race-specific effects in major genic and polygenic resistance of barley to barley leaf rust in the field: identification and distinction

Summary Six partially resistant spring barley cultivars were exposed to four barley leaf rust (Puccinia hordei) races in the field and in the greenhouse. The 24 cultivar-race combinations were tested in field plots of 1.5×1.5 m² in two replications over two years. To reduce the interplot exchange of urediospores each plot was surrounded by winter rye.The level of barley leaf rust varied among cultivars, races and years. In both years the variance for cultivar-race interactions was highly significant and originating largely from the cultivar-race combinations Berac-22. Armelle-22, Armelle-A and Tyra-A. The Berac-22 interaction was towards higher, the other three interactions towards a lower level of barley leaf rust. The reduced rust levels of these three combinations were not due to interactions between the partial resistance of these cultivars and the aggressiveness of the races but to major genes for hypersensitivity not effective to the races 1-2-1 and F, common in Western Europe, but effective against the rare races 22 and A. This was revealed in the greenhouse experiments where all combinations had a susceptible infection type except Armelle-22, Armelle-A and Tyra-A, which showed low infection types in both the seedling and adult plant stages. The urediosori present in the field plots of these three combinations apparently arose from spores derived from other plots; this interplot interchange suggesting partial resistance.The interaction of Berac with race 22 truly was a small race-specific effect within the polygenic, partial resistance of barley to barley leaf rust like the one reported before between Julia and race 18.     

Evaluation of seven Lycopersicon species for resistance to tomato spotted wilt virus (TSWV)

Summary One hundred eighty-eight accessions of Lycopersicon cheesmanii, L. chilense, L. chmielewskii, L. hirsutum, L. parviflorum, L. pennellii, and L. peruvianum were screened for resistance to three isolates of tomato spotted wilt virus (TSWV). All plants in an accession were initially screened for resistance to TSWV using isolate 85–9 from Arkansas. Visual symptoms were used to cull obviously infected plants, followed by enzyme-linked immunosorbent assay (ELISA) to identify uninfected plants. Cuttings were taken from uninfected plants in the first screening and the resulting plants were inoculated with isolates Glox and T-2 from Texas and Hawaii, respectively. No resistance was identified in L. cheesmanii, L. chmielewskii, L. hirsutum, L. parviflorum, and L. pennellii. However, 33 of 63 L. chilense accession produced 91 of 1268 plants that were uninfected with isolate 85–9 and 20 accessions that produced 40 of 257 plants that were not infected with any of the isolates. After screening with isolate 85–9 9 of 12 L. peruvianum accessions tested had 38 plants uninfected and 8 accessions had 25 plants that were not infected with any of the isolates.     

抗植物真菌病害基因及其介导的抗性机理

抗植物真菌病害基因种类繁多,其介导的抗性机理各不相同。在试验研究中应用比较成功的主要有9类,分别是:细胞毒素RNA酶基因、RNA酶抑制剂编码基因、几丁质酶基因、β-1,3-葡聚糖酶基因、核糖体失活蛋白基因、植保素合成酶基因、抗真菌蛋白基因、过氧化物酶基因和病程相关蛋白基因。     

Genetic analysis of late blight resistance in Solanum pimpinellifolium accession PI 270441: Heritability and response to selection

Late blight (LB), caused by Phytophthora infestans, is a destructive disease of tomato (Solanum lycopersicum) worldwide. Currently, there are few commercial cultivars of tomato with resistance to LB, and the disease is mainly controlled by heavy use of fungicides. Due to the emergence of fungicide‐resistant pathogen isolates, there is a concerted effort to identify new genetic sources of resistance and breed new resistant cultivars. A recent screening identified several new tomato accessions with strong resistance to LB. Here, we report on the genetic basis of LB resistance in S. pimpinellifolium accession PI 270441, as determined by generation means analysis and analysis of response to selection, using populations derived from crosses with LB‐susceptible breeding line Fla. 8059. Heritability of LB resistance ranged from 0.76 to 0.78, and the minimum number of genes was estimated 1—few. These results suggest that transfer of LB resistance from PI 270441 to the cultivated tomato should be feasible via a traditional backcross breeding approach. Genetic mapping studies are underway to identify molecular markers associated with resistance in this accession.     

Variation in resistance to the grain aphid, Sitobion avenae (Sternorhynca: Aphididae), among diploid wheat genotypes: Multivariate analysis of agronomic data

A collection of 87 ancient wheat genotypes, 67 Triticum monococcum, 13 Triticum boeoticum, seven Triticum urartu and one cultivar of the modern wheat Triticum aestivum (variety ‘Arminda’) were evaluated for resistance to the cereal aphid Sitobion avenae, the main damaging aphid pest on winter wheat in Europe. The intrinsic rate of natural increase (r_m), which is regarded as a good estimate of the fitness of an aphid population, was used as an indicator for the level of plant resistance. Differentiation of the 88 plant genotypes into four distinct groups was achieved with a cluster analysis of the r_m values. The modern wheat ‘Arminda’ was more susceptible than any of the ancient wheat genotypes tested (r_m= 0.24, i.e. the aphid population doubled every 2.6 days). A second group of 19 plants ranged from relatively susceptible to moderately resistant (0.17 < r_m < 0.21). Fifty–one plants were allocated to a third group and classified as resistant (0.09 < r_m < 0.16). The last group contained 17 genotypes with a high level of resistance where aphid fitness was greatly reduced (0.02 < r_m < 0.09, i.e. the aphid population doubled every 11.4 days or 7.7 days, respectively). Clustering of the accessions into the different phenetic groups did not follow the geographical origin of the wheat genotypes or the species to which they belong. These results show that ancient diploid wheats, all characterized by the genome A, present considerable interest for plant breeding for resistance to S. avenae in modern wheat. The potential use of these strong and partial sources of resistance for introduction of a stable and durable form of resistance to S. avenae in wheat is discussed.     

Seasonal variation in resistance of chrysanthemum cultivars to Frankliniella occidentalis (Thysanoptera: Thripidae)

Seasonal variation in the level of host plant resistance can have important consequences for the repeatability of tests to measure host plant resistance to insect pests. In the present study, the levels of resistance to Frankliniella occidentalis of a susceptible and a partially resistant cultivar of chrysanthemum, Dendranthema grandiflora, were determined throughout the year. Thrips damage, reproduction and adult female survival were determined on excised leaves in Petri dishes under uniform conditions. Strong seasonal fluctuations were observed in these three characteristics. On leaves from plants grown in winter, damage, reproduction, and survival were higher than on leaves from plants grown in summer. Clear differences in resistance were observed between the susceptible and the partially resistant cultivar on leaves from plants grown in winter, while differences disappeared in summer. Damage on both cultivars and survival on the susceptible cultivar were negatively correlated with mean daily solar radiation during plant growth, suggesting that the level of resistance depends on light intensity during plant growth. This was confirmed in an experiment carried out in summertime with shaded and unshaded plants. Leaves from control plants, grown under high light intensity had a higher level of resistance than leaves from shaded plants grown under reduced light intensity. There were clear differences in resistance between the cultivars under shaded conditions (low light intensity), but not under unshaded control conditions (high light intensity). This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance to Pseudomonas solanacearum in the potato: I. Effects of sources of resistance and adaptation

Summary F1 hybrid true potato seed progenies with multiple sources of specific resistance to Pseudomonas solanacearum and adaptation were evaluated under tropical greenhouse conditions for resistance to a race 1 isolate of P. solanacearum. Results indicated that genes for adaptation are involved in conferring resistance to bacterial wilt. The effect of a particular resistant parent clone on the resistance level of its progeny depended on the resistance, adaptation or the combination of both characteristics of its mate. A heat tolerant parent gave a higher frequency of resistant offspring in combination with an ascertained source of resistance. Combining ability was an apparent feature of resistance to bacterial wilt and resistance was highest in the majority of the crosses that had a wide genetic background for both resistance and adaptation. The possible genetic nature of the resistance to bacterial wilt is discussed.     

Inheritance of resistance to damage byThrips tabaci Lindeman (Thysanoptera: Thripidae) in Cabbage

Summary The inheritance of resistance in cabbage (Brassica oleracea var.capitata) to damage caused byThrips tabaci Lindeman was studied in progeny from four crosses between resistant and susceptible inbred lines. In two families sharing the same susceptible parent, the narrow-sense heritability was calculated to be high (>90%), using the Warner method of calculation (Warner, 1952). The other two families, also sharing a susceptible parent, had very low narrow-sense heritability (0–11%). In three of the four families, however, the validity of the Warner calculations were called into question due to significant epistatic interactions. In all four families, the F₁ populations approached or exceeded the susceptibility of the susceptible parent, indicating that susceptibility is generally dominant. The importance of epistasis and dominance suggests that tests of hybrid combinations to determine combining ability for thrips resistance may be required, rather than selecting only on the basis of thrips damage to the inbred lines. The technique of planting wheat upwind from the cabbage test plot did generate adequate levels of thrips pressure, but the infestation was uneven in two of the four blocks, so that reasonable replication of tests is required.     

Levels of host plant resistance to banana weevil, Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae), in African Musa germplasm

Forty-five Musa clones, including endemic and introduced cultivars plus hybrids, were evaluated for resistance against the banana weevil, Cosmopolites sordidus, in a field trial in Uganda. The predominant groups of staple crops, East African highland bananas (Musa spp. AAA) and plantains (Musaspp. AAB), as well as plantain-derived hybrids (AAB × AA), showed the highest levels of susceptibility to this pest. These were followed by dessert bananas (Musa spp. AAA), exotic bananas (Musa spp. ABB) and finally diploids of M. acuminata (AA). Hybrids of banana origin were highly resistant. Some East African highland cultivars, especially brewing types (e.g., Kabula, Bagandeseza, Ediirira), showed intermediate levels of resistance. Among the non-highland bananas, high levels of resistance were observed in Yangambi-Km5 (AAA), Cavendish (AAA), Gros Michel (AAA), Kayinja (ABB, Pisang Awak subgroup), Ndiizi (AB, Ney Poovan subgroup)and Kisubi (Ney Poovan subgroup). The highest resistance was observed in banana hybrids TMB2×7197-2, TMB2×8075-7 and the wild banana Calcutta-4 (AA). These were considered the best sources of resistance for a weevil resistance-breeding programme with the two hybrids commonly used as improved male parents. This revised version was published online in July 2006 with corrections to the Cover Date.