Resistance of wheat to Fusarium spp. in an in vitro seed germination assay and preliminary investigations into the relationship with Fusarium head blight resistance

Resistance of wheat cultivars from the UK, 2003 recommended list were evaluated against Fusarium culmorum, F. graminearum, Microdochium nivale var. majus and var. nivale, F. avenaceum and F. poae using a seed germination assay and related to previously published data on detached leaf assays, using M. nivale, and UK recommended list Fusarium head blight (FHB) resistance ratings. Significant cultivar differences were found after inoculation with all fungal species with the exception of F. poae where the decline in the percentage of seed germinating relative to the controls was small. Correlations of the percentage seed germinating inoculated with the remaining Fusarium spp. and M. nivale were high (r = 0.68; P < 0.01 to r = 0.94; P < 0.001). Overall, F. graminearum caused the greatest reduction with a mean seed germination of 61.7% relative to the controls followed by F. avenaceum (65.5%), M. nivale var. majus (67.2%), F. culmorum (76.6%), M. nivale var. nivale (89.2%) and was least for F. poae (92.5%). The resistance detected in the germination assay was significantly correlated to whole plant FHB resistance ratings (r_s = 0.45; P < 0.05) but was not correlated to partial disease resistance (PDR) components detected using the detached leaf assay, namely, incubation period, latent period and lesion length. The results suggest that while resistances detected in the seed germination and detached leaf assays may in part share a common genetic basis to FHB resistance, resistances detected in both in vitro assays are independent of each other. Resistances detected in the seed germination assay had a lower correlation with FHB resistance ratings against F. culmorum, than those detected by latent period in the detached leaf assay.     

Genetics and population improvement of resistance to bacterial soft rot in Chinese cabbage

Resistance to bacterial soft rot caused by Erwinia carotovora subsp. carotovora is a quantitative trait. The narrow-sense heritability was from 42% to 60% in the studied populations. Griffing's diallel analysis and generation mean analysis indicated that additive genetic effects were most significant, and the resistant sources involved in this study appeared to have different genes or alleles. Recurrent phenotypic selection was used for improving the resistance level by combining different resistance genes from the selected genotypes of Chinese cabbage. After 3 cycles of recurrent selection, the level of resistance increased markedly. When the cycle 3 population was compared with the cycle 0 population, the mean disease severity rating was reduced 38% from 6.3 to 3.9, and the percentage of plant survival increased from 65% to 97% based on the mean of two years experiments using mist-chamber seedling inoculation methods. The improvement of resistance was also demonstrated in field assays. This revised version was published online in July 2006 with corrections to the Cover Date.     

Brassica coenospecies: a rich reservoir for genetic resistance to leaf spot caused by Alternaria brassicae

Development of leaf spot resistant mustard cultivars is a relevant objective in view of heavy crop losses caused by this pathogen. Thirty-eight species belonging to 9 genera, including cultivated and wild allies, of the genus Brassica were evaluated under epiphytotic conditions for two years. Inoculations were done on whole plants (in vivo) and on detached leaves (in vitro). Data on incubation period, number of lesions per leaf, lesion size and leaf area covered by lesions were recorded. Species which never produced disease symptoms throughout the growing period in pots and until 72 hours after inoculation in detached leaf assays during both years were treated as resistant, while those that produced symptoms were classified as moderately resistant, susceptible or highly susceptible depending upon incubation period, size of lesions and leaf area covered by disease symptoms. Eight species (Brassica desnottesii, Camelina sativa, Coincya pseuderucastrum, Diplotaxis berthautii, D. catholica, D. cretacea, D. erucoides, and Erucastrum gallicum) were found completely resistant, whereas others were classified as moderately resistant (12), susceptible (11) or highly susceptible (9). Since resistance is unavailable within the cultivated species, these 8 resistant wild species could be used as donor parents for introgressing resistance to leaf spot disease in Indian mustard. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of partial resistance to spot blotch in barley

Diallel crosses (without reciprocals) were made among 10 different barley genotypes with genetic variability for spot blotch resistance. Forty‐five F₁ hybrids and their parents were assessed for their combining abilities for the disease resistance. Three experiments, two in a growth chamber on detached leaf and seedlings tests and one in the field on adult plant stages, were undertaken using a randomized complete block design with five replicates. A mixed conidial suspension of nine virulent isolates of the pathogen was used for inoculation. Statistical analysis showed genetic variability for spot blotch resistance. Results showed that the cultivar Banteng, the Ethiopian line CI‐5791 and the Syrian line 79‐SIO‐9 had partial resistance in all experiments. General combining ability was significant, with either positive or negative values. Resistant genotypes show favourable GCA‐effects, and they could therefore be successfully used for breeding purposes.     

Phenotyping early blight resistance in potato cultivars and breeding clones

Thirty-two potato cultivars/clones were evaluated for resistance to early blight using different methods. The evaluations were performed under field and greenhouse conditions. In the field experiments, plants were evaluated for disease symptoms, and the relative area under the disease progress curve and percent defoliation were determined. In the greenhouse experiments, leaf lesion sizes were determined on either intact plants or detached leaves after drop inoculation with Alternaria solani. The effect of leaf position (lower, middle or upper part of the plant) on lesion size was investigated. There was no correlation between lesion sizes on lower leaves and upper leaves after inoculation. However, significant correlations between lesion sizes on lower and middle leaves and also between middle and upper leaves were found. Furthermore, we found significant correlation between the results of resistance studies in the field and in intact plant inoculation experiments in the greenhouse. In contrast, results from the detached-leaf experiment were not correlated with either greenhouse intact plant tests or field results. The results indicate that using detached-leaf assays for screening potato for early blight resistance is not accurate. We found significant differences in resistance to A. solani among cultivars/clones in both the field and in greenhouse experiments.     

Resistance toPhytophthora porri in leek and some of its wild relatives

Summary Winter leek and related plant material was evaluated for resistance to white tip disease (Phytophthora porri) in the Netherlands. Significant differences in partial resistance were found between winter leek accessions in field tests with zoospore and natural inoculation. These differences were consistent over three years, and indicate that resistance can be improved through mass selection in winter leek cultivars. Some accessions of wild or cultivated relatives of winter leek with higher levels of resistance were found in a glasshouse test with zoospore inoculation. These sources of resistance were also highly resistant in field experiments over a two year period.     

Influence of plant age,disease incubation time,and host plant resistance on Verticillium wilt symptom expression in lucerne (Medicago sativa)

Summary The most economical means to control Verticillium wilt in lucerne, caused by Verticillium albo-atrum, is through resistant cultivars. Most Verticillium wilt resistance evaluations use some form of root-cut-soak inoculation procedure. In Trial 1, three lucerne cultivars were inoculated at eight plant ages and plant symptoms were scored at ten successive weekly intervals. Trial 2 consisted of two cultivars inoculated at ten plant ages and symptoms were evaluated at eight weekly intervals. In an experiment to study the genetics of resistance to this pathogen, four plants showing no visual symptoms were selected from each of four germplasms. The selected populations were crossed in diallel fashion. The population crosses plus the parents were evaluated for resistance to V. albo-atrum. Results from Trial 1 and Trial 2 showed disease severity increased with increasing inoculation age and with increasing scoring age. Cultivar symptom expression interacted with inoculation age, but not with scoring age. The diallel analysis supported resistance to Verticillium wilt being conferred by additive gene action, as heterosis was not detected and general combining ability mean square was 3.75 times larger than the specific combining ability mean square.     

Inheritance and general combining ability studies of detached pod, leaf disc and natural field resistance to Phytophthora palmivora and Phytophthora megakarya in cacao (Theobroma cacao L.)

Black pod is a major disease of cocoa. Knowledge of the inheritance of resistance to black pod would be important in devising strategies to breed resistant varieties. The mode of inheritance of resistance to black pod disease was investigated using leaf disc, detached pod tests and natural field observation in a 6?×?4 factorial and 6?×?6 diallel mating designs analyzed using North Carolina design-II approach and Griffing??s method II model I approach, respectively. The dominant component was smaller than the additive component for pod lesion size, leaf disc scores and natural field infection. Both general combining ability and specific combining ability (SCA) effects influenced the inheritance of pod lesion number and leaf disc scores suggesting that both additive and non-additive (dominance or epistatic) effects influence the inheritance of resistance to black pod disease. However, SCA effect was not significant for pod lesion size and natural field infection suggesting the importance of additive genes in the inheritance of these traits. The higher heritability of leaf disc scores, pod lesion numbers and pod lesion sizes than the heritability of natural field infection which was almost zero, shows the influence of environment on natural field infection and suggests that ranking of resistance of cocoa genotypes need to be done under controlled environmental conditions to observe true resistance levels. Good general combiners, in combination with each other, did not necessarily yield good specific combinations suggesting that parents?? performance cannot be used for selecting superior progenies. The non-significance of reciprocal effects indicates absence of maternal effects or cytoplasmic inheritance in resistance to black pod disease. Recurrent selection procedures and inter-crossing aimed at general combining ability would be more suitable for improvement of black pod disease resistance. Cocoa genotypes with good combining ability for resistance to black pod disease such as T60/887, Pa 150, Sca 6, and Pa7/808, identified in this study could be important parents to form base population for such recurrent selection procedures.     

Expected selection efficiency for resistance to cacao pod rot (Phytophthora palmivora) comparing leaf disc inoculations with field observations

Summary A genetic analysis of resistance of cacao to Phytophthora palmivora was carried out in a 5 × 5 diallel and in a 4 × 2 NC II factorial design, involving cross-progenies and parental clones planted in the nursery and field, respectively. Resistance was scored in the laboratory by inoculation of leaf discs with P. palmivora spores with four replicates and, for the factorial design, results were compared with the percentage of rotten pods in the field assessed over a seven-year period. Significant differences between genotypes were observed for both laboratory and field evaluation methods. For the laboratory test, no reciprocal effect was observed and plant effects within seedling progenies were more significant than plant effects within clones. General combining ability was the main source of variation but specific combining ability was also significant for the leaf disc test applied to the diallel. Heritabilities increased with the number of replicates, reaching values of 0.34 and 0.67 for narrow sense heritability, and 0.60 and 0.67 for broad sense heritability, for the diallel and the factorial design, respectively. These values were higher than observed for the percentage of rotten pods in the field in the factorial design (0.42 and 0.47, respectively). For the leaf test, the expected genetic gains were around 30% with a selection intensity of 5%. The existence of significant genetic (0.71) and phenotypic (0.39) coefficients of correlation between resistance on leaves and percentage of rotten pods in the field in the factorial design confirms the feasibility of using the leaf disc test for early selection of resistance to Phytophthora pod rot of cacao.     

Inheritance of Partial Resistance to Net Blotch in Barley

Reciprocal crosses were made between 9 different barley genotypes with high genetic variability for net blotch resistance. Parents and 72 F₁ plants were used to determine the inheritance of partial resistance to net blotch. Four experiments, one in a growth chamber on seedlings and 3 others in the field on adult plants, were undertaken using a randomised complete block design. An isolate of net blotch from local cultivars was used for inoculation. Non-inoculated plants of one of the field experiments were used for the detached leaf test in petri dishes. Results show that the cultivars ‘Banteng’, and ‘Arrivate’, the Ethiopian line CI 5791, and the Syrian line 79 SIO-10, had a high partial resistance in all experiments. Diallel analysis showed high significant general and specific combining ability when maternal and reciprocal effects were not significant. As the resistance genotypes have a high additive genetic effect, they could be successfully used for breeding purposes.     

Diallel analysis of sooty stripe resistance in sorghum

Sooty stripe [Ramulispora sorghi (Ellis and Everhart) Olive and Lefebre] is a widespread foliar disease of sorghum [Sorghum bicolor (L.) Moench] in West Africa, responsible for grain yield losses up to 46%. We studied the inheritance of sooty stripe resistance in a 9 × 9 sorghum F₂-population diallel grown together with parent lines and checks in1996 under natural disease pressure at two locations in Mali. The percentage of infected leaf area was determined twice over a two-week interval during the season. At the second evaluation, the mean sooty stripe severity amounted to 13% infected leaf area at Samanko and 12% at Cinzana. The frequency distribution of the entries was approximately normal for the mean disease severity, averaged across assessment dates and locations, pointing to the involvement of multiple genes. With the data combined across the two locations, genetic differences among lines and among F₂ populations were highly significant. Genotype × location interaction variances were also significant but much smaller than the genetic variances. Broad-sense heritability estimates were 0.92 for lines and 0.94 for the F₂ populations, for the mean percentage infected leaf area across the two assessment dates. General combining ability effects (GCA) determined most of the differences among the F₂ populations. Specific combining ability effects (SCA), and the interactions of GCA or SCA with locations were also significant but less important. Line performance per se was highly correlated with GCA. Because of the high heritability and predominance of additive effects, prospects are good for the genetic improvement of resistance to sooty stripe in sorghum in Mali, using simple pedigree or recurrent selection procedures. This revised version was published online in August 2006 with corrections to the Cover Date.     

Diallel analysis of barley for resistance to leaf stripe and impact of the disease on genetic variability for yield components

Barley breeders in Syria attempting to develop barley (Hordeum vulgare L.) cultivars resistant to barley leaf stripe (BLS) disease caused by Pyrenophora graminea Ito & Kuribayashi [anamorph Drechslera graminea (Rabenh. Ex. Schlech. Shoem.)]. Information on the combining ability for BLS resistance in Syria is not available. This study was conducted to evaluate, in 10 genetically diverse barley parents, general combining ability (GCA) and specific combining ability (SCA) effects towards the determination of the genetic basis of disease resistance and to estimate genetic variability for yield components and its modification by BLS. Ten parental genotypes varying in their reactions to BLS were crossed in a half-diallel mating design to generate 45 full-sib families. The families and the parents were inoculated with P. graminea and evaluated for resistance in replicated field tests (three inoculated and three non-inoculated plots). The parents chosen showed wide variations for resistance to BLS. Genetic component analysis showed significant effects for both GCA and SCA for resistance to BLS, suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. GCA effects were more important than SCA effects. Resistant parents exhibited high negative GCA indicating that additive gene effects were more predominant, and suggesting their prime suitability for use in barley breeding programs to improve resistance to BLS. Narrow-sense heritability was 58% and broad-sense heritability was 99% indicating that selection for BLS resistance should be effective in these crosses. A high genetic variability for the agronomic traits studied was observed. Yield components decreased significantly in inoculated plants and more pronounced in diseased plants. Significant GCA was observed for all traits. Values for GCA were, in some cases, significantly modified by BLS. This indicates that attention must be paid to the danger of drawing conclusion in quantitative genetics studies dealing with both diseased and healthy plants. Two genotypes, Banteng and Igri, had high negative GCA effects and are promising parents for enhancement of BLS resistance.     

Combining ability analysis of resistance to Helminthosporium leaf blight in spring wheat

Wheat breeders in South Asia are attempting to develop wheat (Triticum aestivum L.) cultivars resistant to Helminthosporium leaf blight (HLB), which occurs mainly as a complex of spot blotch caused by Cochliobolus sativus (Ito & Kuribayashi) Drechs. ex Dastur, and tan spot caused by Pyrenophora tritici-repentis (Died.) Drechs. Information on the combining ability for HLB resistance in wheat cultivars of South Asia is not available. This study was undertaken to examine the resistance to HLB in nine genetically diverse wheat parents, and to evaluate their general combining ability (GCA) and specific combining ability (SCA) effects toward determining the genetic basis of disease resistance. Nine parents were crossed in a half-diallel mating design to produce 36 populations. The F₁ and F₂ progenies, and the parents were evaluated in replicated field tests at Rampur, Nepal. Multiple disease scores were recorded, and area under the disease progress curve (AUDPC) was calculated to measure disease severity over time. The combining ability analysis was performed using Griffing's Method 2, Model 1. The parents chosen showed wide variation for resistance to HLB. They and the F₁ and F₂ progenies differed significantly for AUDPC. GCA and SCA effects were significant in both generations suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. Wheat genotypes 'SW89-5422', 'G 162', 'NL 781'and 'Chirya 7' had significantly negative GCA effects for AUDPC in both F₁ and F₂ generations, suggesting their prime suitability for use in wheat breeding programs to improve resistance to HLB. The estimate of narrow-sense heritability was 0.77 in both generations suggesting that selection for HLB resistance should be effective in these crosses. The results indicate a predominance of additive gene action in the inheritance of HLB resistance in spring wheat.     

The influence of disease complexes involving Leptosphaeria (Septoria) nodorum on detection of resistance to three leaf spot diseases in wheat

Summary In controlled inoculation studies with Septoria nodorum and Pyrenophora tritici-repentis, estimates of the relative proportion of each pathogen demonstrated differences in the responses of cultivars to pathogen mixtures that were not apparent from measurements of diseased leaf areas. Under field conditions estimates of the relative proportion of S. nodorum, P. tritici-repentis and S. tritici varied between field screening locations in Western Australian but also between lines within locations. Lines with known resistance to P. tritici-repentis and S. tritici, but susceptible to S. nodorum, could not be distinguished from susceptible lines on the basis of leaf area diseased or grain weight depression when S. nodorum was present in the disease complex. Such conditions, while suitable for the selection of combined resistance to these pathogens, were unsuitable for identifying resistance to individual pathogens. As symptoms were similar, the proportion of diseased leaf area sporulating with each pathogen provided a means of measuring the variation in disease development induced on lines varying in resistance. Knowledge of the components of disease and their relative importance were essential in understanding varietal response information under mixed infections of these leaf spot pathogens.     

Application of multi-phase experiments in plant pathology to identify genetic resistance to <Emphasis Type="Italic">Diaporthe toxica</Emphasis> in <Emphasis Type="Italic">Lupinus albus</Emphasis>

Phenotyping assays in plant pathology using detached plant parts are multi-phase experimental processes. Such assays involve growing plants in field or controlled-environment trials (Phase 1) and then subjecting a sample removed from each plant to disease assessment, usually under laboratory conditions (Phase 2). Each phase may be subject to non-genetic sources of variation. To be able to separate these sources of variation in both phases from genetic sources of variation requires a multi-phase experiment with an appropriate experimental design and statistical analysis. To achieve this, a separate randomization is required for each phase, with additional replication in Phase 2. In this article, Phomopsis leaf and pod blight (caused by Diaporthe toxica) of Lupinus albus was used as a case study to apply a multi-phase experimental approach to identify genetic resistance to this pathogen, and demonstrate the principles of sound experimental design and analysis in detached plant part assays. In seven experiments, 250 breeding lines, cultivars, landraces, and recombinant in-bred lines from a mapping population of L. albus were screened using detached, inoculated leaves, and/or pods. The experimental, non-genetic variance in Phase 2 varied in magnitude compared to the Phase 1 experimental, non-genetic variance. The reliability of prediction for resistance to Phomopsis pod blight was high (mean of 0.70 in seven experiments), while reliability of prediction for leaf assays was lower (mean 0.35–0.51 depending on the scoring method used).     

Comparison of evaluation methods for selection of resistance to Fusarium head blight in a recurrent selection programme in wheat (Triticum aestivum L.)

The objectives of this study were to compare efficiency of evaluation for resistance to Fusarium head blight (FHB) under two inoculation methods in a recurrent selection programme. Fifty selected homozygous F₅ fertile lines, from each of five cycles (C0, C1, C2, C3 and C4) of recurrent selection, and two control cultivars were evaluated in a split-plot design in 1995 and 1996 under the soil-surface inoculation with Fusarium graminearum-colonized kernels and the single-floret inoculation with ascospore suspension. Comparison of the two inoculation methods using means, ranges, coefficients of variation, heritabilities and correlations among infected spikelet rate (ISR), reaction index (RI) and disease index (DI) indicated that FHB resistance could be evaluated with similar accuracy and precision using either of the two inoculation methods. Regressions of disease scores in the soil-surface inoculation on disease scores in the single-surface inoculation were positive and highly significant, showing a strong relationship between both inoculation methods for FHB resistance. The percentage of lines with similar performance for FHB disease scores in both inoculation methods was high. The soil-surface inoculation and single-floret inoculation appear to be useful techniques for evaluating numerous individuals of segregating population and screening advanced homozygous lines for FHB resistance in a recurrent selection programme in wheat, respectively.     

Assessment of quantitative resistance of rice cultivars to Xanthomonas campestris pv. Oryzae: a comparison of inoculation methods

Summary Lesion size and lesion number were measured on cultivars of rice inoculated by clipping or spraying with virulent isolates of Xanthomonas campestris pv. oryzae. Mean percentage diseased leaf area (%DLA) gave a similar ranking for the two inoculation methods but differences in lesion size among cultivars were much more evident after clip than after spray inoculation. Correlation between the methods was high (r=0.82^**), but some cultivars responded differently with the two techniques. Cultivars which had low scores following spray inoculation showed low disease progress during the first nine weeks after transplanting into a screen-house experiment. Assessment after clip inoculation measures resistance due to spread of bacteria within the leaf xylem system, an important component of quantitative resistance. Assessment after spray inoculation measures all resistance, including resistance to entrance of bacteria into the leaf. In order to select rice entries with improved quantitative resistance to X. c. oryzae based on both components, a screening based on lesion length after clip inoculation, followed by a test for lesion number after spray inoculation, is advised.     

Genetic Analysis for Partial Resistance to an Iranian Strain of Bacterial Leaf Streak (X. campestris pv. hordei) in Barley

Seven barley genotypes with high genetic variability for resistance to bacterial leaf streak (Xanthomonas campestris pv. hordei) were crossed in diallel fashion to determine the inheritance of resistance to this disease. Two experiments were undertaken in a controlled growth chamber using a complete-block design with four replicates. Each replicate consisted of a row of 20 seedlings per parent or F₁ hybrid. An Iranian strain of bacterial leaf streak was used for inoculation of 12-day-old seedlings. Results showed that the cultivars ‘Express’ and ‘Morex’ and the Iranian pure line Iran-3a had a high partial resistance in booth experiments. Diallel analysis showed highly significant general and specific combining abilities. ‘Morex’ and Iran-3a were the best combiners for partial resistance to bacterial leaf streak in barley and could be successfully used for breeding purposes.     

Diallel analysis of resistance to head blight caused by Fusarium culmorum in winter wheat

Summary Ten homozygous winter wheat genotypes representing different levels of resistance to Fusarium head blight were crossed in all possible combinations excluding reciprocals. Parents, F₁ and F₂ were inoculated with one pathogenic strain of Fusarium culmorum. Data for head blight, observed 21 days after first inoculation (OBS-2), and for the area under the disease progress curve, based on observations 14, 21 and 28 days after first inoculation (AUDPC), were analyzed. The contrast between parents and F₁ crosses indicated dommance effects of the resistance genes. Diallel analysis according to Griffing's Method 4, Model 1 showed significant general combining ability (GCA) effects for both F₁ and F₂; specific combining ability effects were not significant. With the exception of one genotype for which general performance for Fusarium resistance was not in agreement with its GCA, the resistance to F. culmorum was uniformly transmitted to all offspring, and the parents can be described in terms of GCA. It is suggested that in the progenies with one of the awned lines as parent, one resistance gene was linked with the gene coding for presence of awns, located on chromosome 4B. A single observation date, taken at the right time, was as effective in assessing resistance as the AUDPC.     

Components of resistance to late leaf spot and rust among interspecific derivatives and their significance in a foliar disease resistance breeding in groundnut (Arachis hypogaea L.)

Late leaf spot (LLS) and rust cause substantial yield losses and reduce the fodder and seed quality in groundnut (Arachis hypogaea L.). Adoption of resistant cultivars by the semi-arid tropic farmers is the best option to overcome yield losses. Knowledge on components of resistance to these diseases should facilitate the development of groundnut cultivars with enhanced resistance to LLS and rust. The objectives of the experiments were to study the genetic variability and relationships among components of resistance to LLS and rust, and assess their significance in disease resistance breeding. Fifteen interspecific derivatives for LLS and 14 for rust and a susceptible control, TMV 2, were evaluated in a randomised complete block design with two or three replications under greenhouse conditions. The experiments were repeated twice. Genotypic differences were highly significant for all the traits studied. Resistance to LLS is due to longer incubation and latent periods, lesser lesions per leaf, smaller lesion diameter, lower sporulation index, and lesser leaf area damage and disease score. Selection based on components of resistance to LLS may not lead to plants with higher retained green leaf area. The remaining green leaf area on the plant should, therefore, be the major selection criteria for resistance to LLS in breeding programs. Resistance to rust is due to longer incubation and latent periods, fewer pustules per leaf, smaller pustule diameter, lower sporulation index, and lesser leaf area damage and disease score. Rust resistant components appear to work additively, therefore, selection based on resistance components together with green leaf area retained on the plant should be the basis of selecting for resistance to rust in breeding programs. ICGV 99005, 99003, 99012, and 99015 for rust and ICGV 99006, 99013, 99004, 99003, and 99001 for LLS are the better parents for use in resistance breeding programs. This revised version was published online in July 2006 with corrections to the Cover Date.