Resistance to the Barley Yellow Mosaic Virus Complex — Differential Genotypic Reactions and Genetics of BaMMV-Resistance of Barley (Hordeum vulgare L.)

Barley yellow mosaic disease is caused by several viruses, i.e. barley yellow mosaic virus (BaYMV), barley mild mosaic virus (BaMMV) and BaYMV-2. The reaction of different barley germplasms to the barley mosaic viruses was studied in field and greenhouse experiments. The results show a complex situation; some varieties are resistant to all the viruses, while others are resistant to one or two of them only. Crosses between different barley germplasms were earned out in order to test whether genetic diversity of resistance against mosaic viruses does exist, particularly, BaMMV. A total of 45 foreign barley varieties were crossed to German cultivars carrying the resistance gene ym4. In F₂ of 27 crosses, no segregation could be detected, leading to the conclusion that the resistance genes of the foreign parents are allelic with ym4 e.g. Ym1 (‘Mokusekko 3’) and Ym2 (‘Mihori Hadaka 3’). A total of 18 crosses segregated in F₂ indicating that foreign parents, like ‘Chikurin Ibaraki 1’, ‘Iwate Omugi 1’, and “Anson Barley”, carry resistance genes different from the gene of German cultivars, e.g. ‘Asorbia’ or ‘Franka’. By means of statistical evaluation (Chi²-test), the observed segregation ratios were analyzed in order to obtain significant information on the heredity of resistance. All the resistance genes described here as being different from the gene ym4, act recessively. Most of the exotic varieties seem to carry only one resistance gene. In a few cases, more than one gene may be present.     

Localisation and combination of resistance genes against soil-borne viruses of barley (BaMMV,BaYMV) using doubled haploids and molecular markers

Barley yellow mosaic virus disease caused by different strains of BaYMV and BaMMV is a major threat to winter barley cultivation in Europe. Different resistance genes against these viruses have been mapped and suitable PCR-based markers have been developed. In this respect doubled haploid (DH) populations proved to be advantageous as they facilitate a repeated test for resistance against all agents of the barley yellow mosaic virus complex and besides this, dominant marker systems are as informative as co-dominant ones in DHs due to the lack of heterozygous genotypes. Using DH populations resistance genes rym4, rym5, rym11, rym13, rym15 and the BaYMV/BaYMV-2 resistance of the barley cultivar ‘Chikurin Ibaraki 1’ have been mapped. DHs are also well suited to pyramiding resistance genes against BaMMV and BaYMV. Since homozygous recessive genotypes are more frequent in DHs than in segregating F₂ populations, DHs can be efficiently used to create broad-spectrum resistance and to extend the usability of partly overcome resistance genes. Results from employing two different strategies for pyramiding, based on one and two DH-steps, respectively, combining three recessive resistance genes, i.e. rym4/rym5, rym9 and rym11, are presented. The faster strategy based on one haploidy step resulted in the identification of all three and two-way combinations of the respective resistance genes.     

Molecular assessment of genetic diversity in winter barley and its use in breeding

Summary During the last decades extensive progress has been achieved in winter barley breeding with respect to both, yield and resistance to fungal and viral diseases. This progress is mainly due to the efficient use of the genetic diversity present within high yielding adapted cultivars and – with respect to resistance – to the extensive evaluation of genetic resources followed by genetic analyses and introgression of respective genes by sexual recombination. Detailed knowledge on genetic diversity present on the molecular level regarding specific traits as well as on the whole genome level may enhance barley breeding today by facilitating efficient selection of parental lines and marker assisted selection procedures. In the present paper the state of the art with respect to virus diseases, i.e. Barley mild mosaic virus, Barley yellow mosaic virus, and Barley yellow dwarf virus is briefly reviewed and first results on a project aiming on a genome wide estimation of genetic diversity which in combination with data on yield and additional agronomic traits may facilitate the detection of marker trait associations and a more efficient selection of parental genotypes are presented. By field tests of 49 two-rowed and 64 six-rowed winter barley cultivars the genetic gain in yield for the period 1970–2003 was estimated at 54.6 kg ha⁻¹ year⁻¹ (r² = 0.567) for the six-rowed cultivars and at 37.5 kg ha⁻¹ year⁻¹ (r² = 0.621) for the two-rowed cultivars. Analysis of 30 SSRs revealed a non-homogenous allele distribution between two and six-rowed cultivars and changes of allele frequencies in relation to the time of release. By PCoA a separation between two and six-rowed cultivars was observed but no clear cut differentiation in relation to the time of release. In the two-rowed cultivars an increase in genetic diversity (DI) from older to newly released cultivars was detected.     

Gene for Resistance to Barley Mild Mosaic Virus in German Winter Barley Located on Chromosome 3L

In order to identify the chromosome arm carrying a gene for resistance to barley mild mosaic virus (BaMMV) in German winter barley cultivars, a line trisomic for the long arm of chromosome 3 (telo-trisomic 3L) was crossed to the resistant cvs. ‘Sonate’ and ‘Ogra’. Results of tests for BaMMV reaction in the F2 indicate that the gene for resistance in German cultivars is located on the long arm of chromosome 3.     

Potential and limitations of isozymes for chromosomal localization of resistance genes against barley mild mosaic virus (BaMMV)

Summary To assess the possibilities offered by isozymes to locate resistance genes against barley mild mosaic virus (BaMMV), the isozyme patterns of 19 barley (Hordeum vulgare L.) genotypes carrying genes different from ym4 were determined. Of the 15 isozyme systems tested, only three were polymorphic, namely aconitate hydratase, esterase, phosphogluconate dehydrogenase, providing markers on four of the seven barley chromosomes. Studies of F₂ progenies derived from three crosses between resistant genotypes and susceptible varieties failed to reveal linkage between resistance genes and isozymes. Another goal of the experiment was to study the linkage relationships between ym4 and the esterase locus (Est1-Est2-Est4). Our estimates of the recombination rate between these two loci (3.41 and 8.32%) were in the range of those reported between these esterases and one of the resistance genes of the Chinese variety Mokusekko 3.     

Utilization of exotic germplasm in Nordic barley breeding and its consequences for adaptation

Summary Utilization of exotic germplasm is one way to broaden genetic variation in breeding populations. This approach has recently been adopted in Sweden and Finland, where experimental barley populations has been established for research and pre-breeding purposes. The aim of the project is threefold: (1) to increase overall genetic diversity of Nordic barley breeding material; (2) to develop breeding material which possesses a high level of resistance for various barley diseases; and (3) to study effects of exotic germplasm on adaptation and agronomic performance. Both the Finnish and the Swedish barley populations include the same exotic material i.e. unadapted landraces from different parts of Asia and wild barley (Hordeum vulgare ssp. spontaneum) accessions. Locally adapted high-yielding barley lines were included in the populations. The establishment of these populations involved six crossing generations in order to promote recombination and enhance the break-up of linkage blocks. The paper discusses the third aim of the project. Studies on agronomic performance and adaptation showed that (1) agronomically valuable genotypes can be constructed through recombination using exotic germplasm for Nordic conditions, (2) that incorporation of exotic material is most successful when made in a local genetic base and (3) that exotic germplasm has an effect on adaptation.     

Development of an STS marker and SSRs suitable for marker-assisted selection for the BaMMV resistance gene rym9 in barley

Based on the RAPD marker OP‐C04H910 which is closely linked to the barley mild mosaic virus (BaMMV) resistance gene rym9 derived from the variety ‘Bulgarian 347’ the marker STS‐C04H910 cosegregating with OP‐C04H910 and generating a single additional band on plants carrying the recessive resistance encoding allele has been developed. Furthermore, the simple sequence repeats (SSRs) WMS6 and HVM67 have been integrated into the genetic map of the rym9 region on chromosome 4HL. Because of their close linkage to rym9 and distinct banding pattern STS‐C04H910 and HVM67 are well‐suited for marker‐ assisted selection, enhanced backcrossing procedures and pyramiding of resistance genes.     

The reaction of x Tritordeum and its Triticum spp. and Hordeum chilense parents to rust diseases

Summary Hexaploid and octoploid tritordeums and their parents Hordeum chilense and Triticum spp. were screened for resistance to isolates of wheat and barley yellow and brown rusts. All H. chilense lines were highly resistant to both wheat and barley brown rust, few lines were susceptible to wheat yellow rust while susceptibility to barley yellow rust was common. In general the resistance of tritordeum is predominantly contributed by the wheat parent and apparently the genes for resistance in H. chilense are inhibited in their expression by the presence of the wheat genome.Abbreviations WYR wheat yellow rust - WBR wheat brown rust - BYR barley yellow rust - BBR barley brown rust     

Characterization of wheat-Thinopyrum partial amphiploids for resistance to barley yellow dwarf virus

Resistance to viruses such as wheat streak mosaic virus (WSMV) and barley yellow dwarf virus(BYDV) is lacking in the primary gene pool of wheat, and therefore resistance is being introgressed from wild relatives such as Thinopyrum species. Resistance to BYDV was found in partial amphiploids (2n = 8x = 56, consisting of 42 wheat and14 alien chromosomes) obtained in hybrids between wheat and both Th. intermedium and Th.ponticum. GISH analysis revealed that the alien genomes of all but one resistant partial amphiploid were heterogeneous consisting of different ratios of St, J^s and J genome chromosomes obtained from theThinopyrum parent. Translocated chromosomes consisting of Robertsonian, interstitial and terminal translocations between the different genomes were also detected. The tissue blot immunoassay showed that partial amphiploids having resistance could be inoculated with the virus but both virus multiplication and spread were completely blocked. This revised version was published online in August 2006 with corrections to the Cover Date.     

Hordeum bulbosum-a new source of disease resistance-transfer of resistance to leaf rust and mosaic viruses from H. bulbosum into winter barley

An outline is given of results for the transfer of new resistances against leaf rust and barley mosaic viruses from Hordeum bulbosum into winter barley. Progenies from backcrosses of barley cultivars with H. bulbosum hybrids were tested both in conventional breeding trials and in additional tests under controlled conditions. Resistance to both pathogens proved to be stable and of good heritability, with differences occurring which depended on the combinations used. Lines with resistance to all leaf rust and mosaic viruses tested, or to either one, were selected. Both resistances segregated independently.     

Genetic analysis of resistance in barley to barley yellow dwarf virus

The inheritance of resistance to barley yellow dwarf virus (BYDV) was studied in the selected 24 spring and winter barley cultivars that showed a high or intermediate resistance level in 1994‐97 field infection tests. The polymerase chain reaction diagnostic markers YLM and Ylp were used to identify the resistance gene Yd2. The presence of the Yd2 gene was detected with both markers in all the resistant spring barley cultivars and lines from the CIMMYT/ICARDA BYDV nurseries. The results of field tests and genetic analyses in winter barley corresponded with marker analyses only when the Ylp marker was used. Genes non‐allelic with Yd2 were detected by genetic analyses and the Ylp marker in moderately resistant spring barley cultivars ‘Malvaz’, ‘Atribut’ and ‘Madras’, and in the winter barley cultivars ‘Perry’ and ‘Sigra’. Significant levels of resistance to BYDV were obtained by combining the resistance gene Yd2 with genes detected in moderately resistant cultivars. The utilization of analysed resistance sources in barley breeding is discussed.     

Screening the Spanish Barley Core Collection for disease resistance

The Spanish Barley Core Collection comprises 159 landrace-derived inbred lines and 16 cultivars adapted to Southern European conditions. The collection was screened for resistance to powdery mildew ( Blumeria graminis ), scald ( Rhynchosporium secalis ), leaf rust ( Puccinia hordei ), net blotch ( Pyrenophora teres f. teres ), Barley yellow dwarf virus (BYDV) and Barley mild mosaic virus (BaMMV). Resistance to powdery mildew was outstanding, with 58 lines presenting mean overall resistance, among them seven landrace-derived lines resistant to all seven isolates tested. About 26% of the Spanish lines were resistant to scald. Resistance to leaf rust and to net blotch was scarce, though a few accessions showed resistance levels as good as the checks. Thirteen accessions (12 Spanish) were totally resistant to BaMMV and c. 20% of accessions showed moderate tolerance to BYDV. Landrace-derived lines from the Mediterranean Coast and Southern regions of Spain were the most resistant to powdery mildew and leaf rust, but the most susceptible to viruses. Potential sources of resistance might be preserved in some accessions subjected to selective pressure in the region of origin.     

Utility of a GFP-expressing Barley yellow mosaic virus for analyzing disease resistance genes

The soil-borne plasmodiophorid Polymyxa graminis is a vector for Barley yellow mosaic virus (BaYMV), which can severely damage barley plants. Although 22 disease resistance genes have been identified, only a few have been used for breeding virus-resistant cultivars. Recently, BaYMV strains capable of overcoming the effects of some of these genes have been detected. In this study, green fluorescent protein (GFP)-expressing BaYMV was constructed and used to examine viral dynamics in inoculated barley plants. Leaf inoculations resulted in higher infection rates than root or crown inoculations. Additionally, inoculations of some resistant cultivars produced infections that were similar to those observed in a field test. The results of this study indicate that the GFP-expressing virus is a useful tool for visualizing virus replication and dynamics, and for understanding resistance mechanisms.     

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.     

Resistance of Cucurbita moschata to watermelon mosaic virus type 2 and its genetic relation to resistance to zucchini yellow mosaic virus

Summary Cucurbita moschata Menina originating from Portugal is known to carry a single dominant gene, Zym, for zucchini yellow mosaic virus (ZYMV) resistance. Resistance to watermelon mosaic virus type 2 (WMV2) was found in the same cultivar. In resistant plants, WMV2 migration from inoculated leaves or cotyledons seems to be limited and blocked. Resistance to WMV2 is conferred by one dominant gene and is effective against eight strains from different geographic origins. Results of linkage studies, based on co-inoculation of plants with WMV2 and ZYMV, indicate that resistance to these two viruses is conferred by the same gene, probably Zym, or perhaps by two closely linked genes.     

The resistance of Hordeum bulbosum and its hybrids with H. vulgare to common fungal pathogens

Summary Two tetraploid and two diploid clones of Hordeum bulbosum were screened for resistance to five isolates of powdery mildew which are virulent on cultivated barley. All were resistant and this resistance was also expressed in hybrids with H. vulgare. The tetraploid genotypes were also resistant to isolates of yellow rust and brown rust. These results show that H. bulbosum contains useful genes for resistance to these diseases and that there is a potential to transfer these into cultivated barley.     

RAPD-mapping of the distal portion of chromosome 3 of barley, including the BaMMV/BaYMV resistance gene ym4

Experiments were carried out in order to identify RAPD-markers for the BaMMV/BaYMV resistance gene ym4 to facilitate efficient marker-based selection in practical breeding programmes. Linkage analysis was carried out with 283 doubled haploid (DH) barley lines out of a cross between the BaMMV/BaYMV susceptible cv. ‘Igri’ and the resistant cv. ‘Franka’. Using bulked segregant analysis, 310 RAPD-primers were screened for polymorphism between the parents. Four of these primers gave rise to specific bands linked to the resistance gene ym4.     

Analysis of durable resistance to stem rust in barley

Summary Since the mid-1940's, barley cultivars grown in the northern Great Plains of the USA and Canada have been resistant to stem rust caused byPuccinia graminis f. sp.tritici. This durable resistance is largely conferred by a single gene,Rpg1, derived from a single plant selection of the cultivar Wisconsin 37 and an unimproved Swiss cultivar. At the seedling stage, barley genotypes withRpg1 generally exhibit low mesothetic reactions at 16–20° C and slightly higher mesothetic reactions at 24–28° C to many stem rust pathotypes. This resistance is manifested by a low level of rust infection and mostly incompatible type uredia on adult plants.Rpg1 reacts in a pathotype-specific manner since some genotypes ofP. g. f. sp.tritici are virulent on cultivars carrying this gene in the field. Several factors may have contributed to the longevity of stem rust resistance in barley, a) since barley is planted early and matures early, it can sometimes escape damage from stem rust inoculum carried from the south; b) one or more minor genes may augment the level of resistance already provided byRpg1; c) the cultivation of resistant wheat cultivars and eradication of barberry have reduced the effective population size and number of potential new pathotypes ofP. g. f. sp.tritici, respectively; and d) virulent pathotypes ofP. g. f. sp.tritici andP. g. f. sp.secalis have not become established. This situation changed in 1989 when a virulent pathotype (Pgt-QCC) ofP. g. f. sp.tritici became widely distributed over the Great Plains. However,Rpg1 may still confer some degree of resistance to pathotype QCC because stem rust severities have been low to moderate and yield losses light on barley cultivars carrying the gene during the last four seasons (1989–1992). Several sources of incomplete resistance to pathotype QCC have been identified in barley. To facilitate the transfer of resistance genes from these sources into advanced breeding lines, molecular marker assisted selection is being employed.     

Disease resistance in protected crops and mushrooms

Summary Cultivars of tomatoes, cucumbers, lettuce and peppers have been bred for resistance to one or more pathogens. Some tomato and cucumber cultivars have resistance to a wide range of diseases. Resistance has been transient in many cases and a succession of cultivars with new genes or new combinations of resistance genes has been necessary to maintain control. There has been a number of notable exceptions and these have included durable resistance to such pathogens asFulvia fulva and tomato mosaic virus. With lettuce the resistance situation is complicated by the occurrence of fungicide resistant pathotypes. There are no strains ofAgaricus bisporus purposely bred for disease resistance.In protected flower crops only resistance to Fusarium wilt in carnations has been purposely bred but differences in disease resistance are apparent in cultivars of many ornamental crops. This is particularly so in chrysanthemums where there are cultivars with resistance to many of the major pathogens. Similar situations occur with other flower crops and pot plants. Cultivars of some species have not been systematically investigated for resistance.The need for genetic resistance will increase with the further reduction, in the limits on pesticide use and an increasing public awareness and importance of pesticide pollution.ADAS is an executive agency of the Ministry of Agiculture, Fisheries and Food and the Welsh Office.     

Inheritance and linkage analysis of resistance to zucchini yellow mosaic virus, watermelon mosaic virus, papaya ringspot virus and powdery mildew in melon

A melon (Cucumis melo L.) breeding line derived from PI 414723 is resistant to three potyviruses,watermelon mosaic virus (WMV), zucchini yellow mosaic virus (ZYMV), papaya ringspot virus (PRSV), and to powdery mildew (PM). The inheritance and linkage relationships of these four resistances were studied in a segregating F2 population and derived F3 families from a cross between cultivar Top Mark and the resistant breeding line. Dominant monogenic inheritance of all four resistances was observed. We report that line 414723-4S3, which was initially selected as a source of ZYMV and WMV resistance, is also a source of dominant monogenic resistances to PRSV and PM race 1. We also report on genetic linkage (significant departure from independent segregation, χ² = 58.1, p≪ 0.0001) between resistance to WMV and ZYMV. The map distance between these loci was estimated to be 7.5 cm. The genes for resistance to PM and PRSV segregated independently from each other, and from ZYMV and WMV resistance. This revised version was published online in July 2006 with corrections to the Cover Date.