A single dominant gene for downy mildew resistance in broccoli

Downy mildew, incited by Peronospora parasitica (Pers.: Fr.) Fr., is a destructive disease of broccoli (Brassica oleraceaL., Italica Group). Resistant cultivars represent a desirable control method to provide a practical, environmentally benign, and long-term means of limiting damage from this disease. Doubled-haploid (DH) lines developed by us exhibit a high level of downy mildew resistance at the cotyledon stage. To determine the mode of inheritance for this resistance, a resistant DH line was crossed to a susceptible DH line to make an F₁, from which F₂ and backcross (BC) populations were developed. All populations were evaluated for response to artificial inoculation with P. parasitica at the cotyledon stage. All F₁ plants (including reciprocals) were as resistant as the resistant parent, indicating no maternal effect for this trait. F₂ populations segregated approximately 3resistant to 1 susceptible, BC populations using the resistant parent as the recurrent parent contained all resistant plants, and the BC to the susceptible parent segregated 1 resistant to 1 susceptible. These results indicate that resistance is controlled by a single dominant gene. This gene should be easily incorporated into F₁ hybrids and used commercially to prevent downy mildew at the cotyledon stage. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of downy mildew resistance in Indian cauliflower (group III)

Summary Inheritance of downy mildew (Peronospora parasitica) resistance in three resistant x susceptible crosses, one susceptible x susceptible and one resistant x resistant cross were studied in Indian cauliflower (Group III) over the two years (1990 and 1991). No significant difference was observed between the years for various estimates and hence pooled data are presented. Downy mildew resistance in crosses cc×HR 5-4 and 3-5-1-1×244 (R×S) is governed by single dominant gene PPA3 but in cross cc×244 (R×S), recessive epistasis was observed. The resistance level was not improved in both the cc×3-5-1-1 (R×R) and 244×267-6-9 (S×S) crosses. Exploitation of downy mildew resistance from cc and 3-5-1-1 in F₁ hybrid is explained in detail.     

Shading of plants facilitates selection for powdery mildew resistance in Squash

Summary Powdery mildew development was assessed on squash (Cucurbita pepo) plants of a susceptible cultivar, a resistant accession, their F₁, and their F₂ in an early summer planting in the field, covered or not covered with a shading net. Three reaction types were observed: susceptible, powdery mildew on stems and on both, the upper and lower leaf surfaces, as in the susceptible parent; resistant, no powdery mildew on leaves or stems, as in the resistant parent; and partially resistant, powdery mildew on upper leaf surfaces only, as in the F₁. Disease presence on the stem was associated with susceptibility. Shading hastened the appearance of powdery mildew and increased the severity of infection on partially resistant and susceptible plants, facilitating identification of resistant individuals in the F₂ population.Contribution No. 1613-E from the Agricultural Research Organization, Bet Dagan, Israel     

Monogenic resistance in tomato to Fusarium oxysporum f. sp. lycopersici race 3

Summary Resistance to fusarium wilt, incited by Fusarium oxysporum (Schlecht.) f. sp. lycopersici (Sacc.) Snyder & Hansen race 3 in tomato (Lycopersicon esculentum Mill.) was discovered in LA 716, a L. pennellii accession. A resistant BC₁F₃ breeding line, E427, was developed from LA 716. E427 was crossed with the susceptible cv. Suncoast and F₁, BCP₁, BCP₂ (to Fla 7155, a susceptible parent) F₂, F₃, and BCP₂S₁ seeds were obtained. Segregation for resistance following root dip inoculation over three experiments indicated a single dominant gene controlled resistance. Five of the 12 BCP₁S₁'s segregated more susceptible plants, whereas one of the 12 segregated more resistant plants than expected (P<0.05). Three of 23 F₃ lines segregated more susceptible plants than expected while 1 of the 23 had more resistant plants than expected (P<0.05). Segregation in all other lines fit expected ratios. Five of the 23 F₃'s were homozygous resistant which was an acceptable fit to expectations (P=0.1–0.5). The gene symbol I ₃ is proposed for resistance to race 3 of the wilt pathogen. Deviations from expected ratios in data reported here and for other breeding lines indicate an effect of modifier genes and/or incomplete penetrance. Plant age at inoculation and seed dormancy did not affect results.Florida Agricultural Experiment Station Journal Series No. 8101.     

Expression of resistance to downy mildew at cotyledon and adult plant stages in Brassica oleracea L.

This research compared the expression of resistance to downy mildew at cotyledon and adult plant stages in seven Brassica oleracea genotypes against two P. parasitica isolates of different virulence. Seven day old seedlings were dual inoculated under controlled environment by depositing two 10 μl droplets of a spore suspension of a different isolate on each cotyledon and the interaction phenotype (IP) evaluated 7 days later using a 0–5 scale of increasing susceptibility. The seedlings were transplanted to 16 cm pots and grown in the greenhouse for 110 days (15 to 28 leaves). Adult plants were tested using a single leaf inoculation method that allowed the same plant to be simultaneously inoculated with the two P. parasitica isolates. Leaves were scored 10 days after inoculation using a 0–5 scale of increasing susceptibility. The inoculation of the same plant with the two isolates produced different combinations of cotyledon (CT) and adult-plant (AP) interaction-phenotype according to the genotype. CrGC 3.1 was susceptible at CT and AP stages, ‘Algarvia’ resistant at CT and AP stages, and broccoli ‘A’ susceptible at CT and resistant at AP stage against the two isolates. ‘Murciana’ and broccoli ‘B’ were differential at CT and AP stages. Savoy and Shetland cabbages were differential at CT stage and resistant to both isolates as adult plants. Cotyledon resistance could not be used to predict adult-plant resistance since the two types of resistance were very poorly correlated. Resistance can be race specific either at cotyledon or at adult-plant stage. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparative genetic analysis of field resistance to downy mildew in the lettuce cultivars ‘Grand Rapids’ and ‘Iceberg’

Downy mildew on lettuce is currently controlled using host resistance genes (Dm genes) that confer race-specific resistance in seedlings. Field resistance (FR) that is active in adult plants but not seedlings was identified in the cvs. Grand Rapids and Iceberg. The goal of our study was to evaluate the utility of Grand Rapids as a source of novel Bremia resistance alleles, particularly in comparison with Iceberg. To measure FR, downy mildew symptoms were evaluated following natural infection in field experiments. The responses of Grand Rapids and Iceberg were similar in many respects. Although both cultivars had a small percentage of plants exhibiting disease symptoms, the average disease ratings were as low as for cultivars with effective Dm genes. We observed no evidence for race specificity. FR was effective over 3 years of our study, despite documented variation within pathogen populations. Both cultivars lacked all known seedling resistance genes except Dm13, which was not responsible for the resistance observed in field experiments. Similar segregation of FR was observed in F₂ populations for both Grand Rapids and Iceberg. The presence of highly susceptible families within Grand Rapids × Iceberg populations suggested the presence of at least one unique resistance allele in each cultivar. Preliminary genetic analysis of FR from Grand Rapids revealed a high estimate of narrow-sense heritability that suggested simple inheritance, but single gene models did not fit the observed data. Our results suggest that Grand Rapids may represent an underutilized resource for controlling downy mildew in lettuce.     

Inheritance of adult-plant resistance to Phytophthora capsici in pepper

Summary Inheritance studies were conducted to determine the genetic basis of adult-plant resistance in pepper (Capsicum annuum L.) to Phytophthora capsici. F₁, backcrosses and F₂ populations were developed using the resistant parent Criollo de Morellos 334 and susceptible parents Agronômico 10-G and Yolo Wonder. Pepper plants, at 36 days post-emergence, were inoculated near the base of the stem with an inoculum suspension of 5×10⁴ zoospores/ml. Segregation ratios in the F₂ generation of 13 resistant to 3 susceptible plants fit a 2-gene model for resistance with dominant and recessive epistasis.     

A demonstration of the inheritance of field resistance to lettuce downy mildew (Bremia lactucae Regel.) in progeny derived from cv. Grand Rapids

Summary The inheritance of field resistance to downy mildew in lettuce derived from cv. Grand Rapids was studied in F₂, F₃ and F₄ progeny from a cross with the more susceptible cv. Lobjoits Green Cos. The results for 32 randomly produced F₃ families and the F₂ population corresponded closely although the absolute level of resistance varied from season to season according to disease pressure and environmental conditions. The field resistance character showed a quantitative pattern of inheritance but the rapid response to selection at F₃ implied that only a few genes may be involved. F₄ families raised from families selected from the two extremes of the F₃ distribution fell into distinct categories with resistance levels corresponding to that of their F₃ parents. An association between resistance and morphological features of the resistant parent was present at F₂ but not in subsequent generations when individuals and families which were as resistant as cv. Grand Rapids and yet bore little morphological resemblance to this cultivar were readily recovered.     

Identification of alleles at two loci controlling resistance to Phomopsis stem blight in narrow-leafed lupin (Lupinus angustifolius L.)

The genetics of resistance to Phomopsis stem blight caused by Diaporthe toxica Will., Highet, Gams & Sivasith. in narrow-leafed lupin (Lupinus angustifolius L.) was studied in crosses between resistant cv. Merrit, very resistant breeding line 75A:258 and susceptible cv. Unicrop. A non-destructive glasshouse infection test was developed to assess resistance in the F₁, F₂, selected F₂-derived F₃ (F_2:3) families, and in selfed parent plants. The F₁ of Unicrop × 75A:258 (and reciprocal cross) was very resistant, and the F₂ segregated in a ratio of 3:1 (resistant: susceptible), which suggested the presence of a single dominant allele for resistance in 75A:258. In Merrit × Unicrop (and reciprocal), the F₁ was moderately resistant, and the F₂ segregated in a ratio of 3:1 (resistant: susceptible). Thus Merrit appeared to carry an incompletely dominant resistance allele for resistance. The F₁ of Merrit × 75A:258 (and reciprocal) was very resistant and the F₂ segregated in a ratio of 15:1 (resistant: susceptible), which supported the existence of independently segregating resistance alleles for resistance in 75A:258 and Merrit. Alleles at loci for early flowering (Ku) and speckled seeds (for which we propose the symbol Spk) segregated normally and independently of the resistance alleles. Resistant F₂ plants gave rise to uniformly resistant or segregating F_2:3 families, whereas susceptible F₂ plants gave rise only to susceptible F_2:3 families. However, the variation in resistance in the F₂ and some F_2:3 families of crosses involving 75A:258, from moderately to extremely resistant, was greater than that expected by chance or environmental variation. We propose the symbols Phr1 to describe the dominant resistance allele in 75A:258, and Phr2 for the incompletely dominant resistance allele in Merrit. Phr1 appears to be epistatic to Phr2, and expression of Phr1 may be altered by independently segregating modifier allele(s). This revised version was published online in July 2006 with corrections to the Cover Date.     

The oat line Pc54 as a source of resistance to crown rust,stem rust and powdery mildew in Europe

Summary The oat line Pc54 was found to be resistant to powdery mildew under both field and glasshouse conditions. The ratio of resistant to susceptible F₂ and F₂ progeny of a cross between a selection from the Pc54 line (Cc7422) and a susceptible cultivar (Selma) showed that, in addition to carrying the crown rust resistance gene Pc54 and the pg15 gene for stem rust resistance, the mildew resistance of the Pc54 line was conditioned by a single incompletely dominant gene along with additional factors which modified the expression of resistance. Previous results, that there was no linkage between genes Pc54 and Pg15, were confirmed. In addition, there was no evidence of linkage between the mildew resistance gene and gene Pc54. Evaluation of selections from within the Pc54 line showed that the expression of both stem rust and mildew resistance was modified by, or linked to, plant height. The effectiveness of genes Pc54 and Pg15, as measured by virulence frequencies, in central and eastern Europe is described.     

Single-gene resistance to powdery mildew in zucchini squash (Cucurbita pepo)

Cucurbita pepo (pumpkin, squash,gourd) is an economically important species that is susceptible to the cucurbit powdery mildew fungus, Podosphaera xanthii(syn. Sphaerotheca fuliginea). ‘True French’, an open-pollinated cultivar of the Zucchini Group of C. pepo, was crossed with an unnamed powdery-mildew resistant straight neck-type accession, the resistance of which was apparently derived from an interspecific cross with a resistant wild species of Cucurbita,and resistant plants were selected in the F₂ generation. This was followed by six cycles of backcross-pedigree selection for resistance, and resulted in the development of an accession true-breeding for resistance to powdery mildew and nearly isogenic to ‘True French’. The resistant and susceptible near-isogenics were crossed and seeds of the filial and backcross generations were produced. Plants of the parental accessions and their progenies were grown together in a controlled-environment chamber, exposed to the pathogenic fungus, and scored as resistant, partially resistant, or susceptible 27–33 days after sowing. The results indicated that resistance is conferred by a single incompletely dominant gene, designated Pm-0. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of resistance to two races of sunflower downy mildew (Plasmopara halstedii) in two Helianthus annuus L. lines

Sunflower downy mildew caused by Plasmopara halstedii is an important disease of sunflower capable of causing losses of more than 80% of production. Races 100, 300, 310, 330, 710, 703, 730 and770 of the fungus have been identified in Spain. Race 703, of high virulence, has been identified frequently in the northeast, while race 310 seems to occur over the south, the main sunflower growing region of the country. Oil sunflower lines RHA-274 and DM4 were studied for their resistance to races 310(RHA-274 and DM4) and 703 (DM4). In each cross, only one plant of the resistant parent was crossed to the inbred susceptible line HA-89 (or cmsHA-89).Plants from F₂ and backcross(BC₁F₁ to susceptible parent)generations were evaluated for fungal sporulation on true leaves and/or cotyledons. The resistant-to-susceptible ratios obtained in the F₂ and BC₁F₁ progenies from the crosses cmsHA-89 × RHA-274 and HA-89 × DM4suggested that one major gene in each line is responsible for resistance to race 703.The segregations of the progenies of the cross HA-89 × DM4 inoculated with race 703also fitted the ratios 1:1 and 3:1 (for BC₁F₁ and F₂, respectively)corresponding to control of resistance by a single dominant gene. In RHA-274, the gene for resistance to race 310 was designated Pl ₉, whereas Pl _v is tentatively proposed to designate the gene in DM4 responsible for resistance to races310 and 703. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic analysis of adult plant resistance to powdery mildew in pea (Pisum sativum L.)

Summary An analysis of adult plant resistance of powdery mildew in 15 F₁, F₂ and F₃ populations of pea derived from crossing 15 diverse and susceptible lines with one resistant line revealed that resistance to powdery mildew is controlled by duplicate recessive genes. The genes were designated as er₁ and er₂.Disease reaction showed independent segregation with three known markers in the resistant parent, namely, af (afila, chromosome 1), st (stipule reduced, chromosome 3) and tl (clavicula, chromosome 7).Contribution form the Department of Genetics and Plant Breeding Banaras Hindu University, Varanasi-221005, India.     

Inheritance of the Powdery Mildew Resistance Mlk in Wheat

The inheritance of the Mlk powdery mildew resistance originating from ‘Heine 2174.50’ was analyzed by crossing the Mlk resistant cultivar ‘Ralle’× cv. ‘Amor’ (highly susceptible) and vice versa and by observing the reactions of F₁- and F₂-plants after inoculation with two different Mlk avirulent powdery mildew isolates. In all cases, a 3 (resistant): I (susceptible) segregation was found in F₂. The reactions of the F₂plants against the two powdery mildew isolates were identical in each case. Therefore, it is supposed that one dominant resistant gene is responsible for the resistant reactions against these two isolates. These results support the earlier assumption of Heun and Fischbeck (1987b) that the whole Mlk resistance pattern is controlled by a single gene.     

Inheritance of resistance to angular leaf spot in Nicotiana tabacum L. cultivars burley 21 and kentucky 14

Summary The genetics of resistance to angular leaf spot caused by Pseudomonas syringae pv. tabaci in Nicotiana tabacum cultivars Burley 21 and Kentucky 14 was investigated by studying disease reactions to three isolates of parental, F₁, F₂ and backcross generations derived from crosses between the resistant cultivars and the susceptible cultivar Judy's Pride. Studies were conducted in the greenhouse and in field plant beds. Chi-square values were computed to determine whether the observed ratios for disease reactions deviated from expected Mendelian ratios for a single, dominant gene controlling resistance to angular leaf spot in tobacco. Based on the resistance of the F₁ and the backcross generation to the resistant parent (BC-R), a 3 resistant: 1 susceptible segregation ratio in the F₂, and a 1 resistant: 1 susceptible segregation ratio in the backcross generation to the susceptible parent (BC-S), it was concluded that resistance to three isolates of Pseudomonas syringae pv. tabaci is governed by a single, dominant gene.     

Screening of melon (Cucumis melo L.) germplasm for multiple disease resistance

Summary Melon germplasm was screened for cucumber green mottle mosaic virus (CGMMV), powdery mildew (Sphaerotheca fuliginea), downy mildew (Pseudoperonospora cubensis) and Fusarium wilt (Fusarium oxysporum f. sp. melonis) resistance under artificial conditions except downy mildew for which screening was done under natural epiphytotic conditions. High level resistance to all the four diseases was not recorded in any of the collections tested. Nevertheless, ertheless, resistance to three diseases was located in three germplasm. Wild Cucumis species C. figarei exhibited absolute resistance to CGMMV and Fusarium wilt and high level resistance to downy mildew. Phoot or snapmelon (Cucumis melo var. momordica) — a non-dessert from of Indian origin—was highly resistant to downy mildew and resistant to CGMMV and medium resistant to Fusarium wilt. Iroquois was resistant to powdery mildew and medium resistant to downy mildew and CGMMV.     

Genetic variability for resistance to sorghum downy mildew (Peronosclerospora sorghi) and Rajasthan downy mildew (P. heteropogoni) in the tropical/sub-tropical Asian maize germplasm

Downy mildews cause considerable damage to maize (Zea mays L.) worldwide, particularly in the tropical Asia. We have evaluated a set of 42 tropical/sub-tropical maize inbred lines developed in different countries in Asia (India, Thailand and Philippines), and Mexico, for analysing the genetic variability for resistance to sorghum downy mildew [Peronosclerospora sorghi; SDM] and Rajasthan downy mildew [P. heteropogoni; RDM]. Experiments were carried out in replicated trials under artificial infection in field conditions against SDM and RDM at Mandya in Karnataka, India, and Udaipur in Rajasthan, India, respectively, during 1999 and 2000. The study resulted in identification of five inbred lines offering consistent and strong resistance to both SDM and RDM, while several inbred lines revealed resistance only to RDM. It was also revealed that the SDM-resistant inbreds are invariably resistant to RDM, while the RDM-resistant inbreds might show differential responses to the SDM. The maize inbred lines identified in this study with broad-spectrum resistance to downy mildews could be potentially useful for basic and applied research work on downy mildews in tropical Asia. This revised version was published online in August 2006 with corrections to the Cover Date.     

Colocation of downy mildew (Plasmopara halstedii) resistance genes in sunflower (Helianthus annuus L.)

Summary The Pl6 locus in the inbred sunflower (Helianthus annuus L.) line HA335 giving resistance to French races of downy mildew (Plasmopara halstedii (Farl.) Berl. & de Toni. was localized by molecular techniques. A bulked segregant analysis was made on the F2 progeny from a cross between this line and H52, a downy mildew susceptible line. The resistance gene in HA335 was found to have the same linked RFLP marker loci as those determined for Pl1 (resistance to race 1 in the line RHA266) on linkage group 1 of the consensus RFLP map of the cultivated sunflower. Pl1 and Pl6 thus appear either to be allelic or closely linked. The implications for sunflower breeding are discussed.     

Inheritance of resistance to race 330 of Plasmopara halstedii in three sunflower lines

Sunflower lines RHA‐274, HA‐61 and RHA‐325 were studied for their resistance to race 330 of downy mildew (Plasmopara halstedii). The same inbred line, with normal (HA‐89) or sterile cytoplasm (cmsHA‐89) was used in all the crosses as susceptible parent, and, in each cross, only one genotype of the resistant parent was studied. The resistant‐to‐susceptible ratios obtained in the BC₁ and F₂ progenies from the crosses of the lines RHA‐274 and HA‐61 to cmsHA‐89 and HA‐89, respectively, suggested that, in each resistant line, two dominant genes are responsible for resistance to this downy mildew race. One of the genes (A) is epistatic to the other (B), and the recessive allele b in homozygosity is also epistatic to aa, with plants carrying aabb genotypes being resistant. Resistance to race 330 seemed to be controlled by two complementary genes in the sunflower inbred line RHA‐325, the dominant allele of one of them being present in cmsHA‐89. In the genotypes HA‐89 or cmsHA‐89, the existence of genes that modify the expected segregations following the crosses with resistant parents is proposed. It is concluded that, although major genes have been described as responsible for monogenic resistance to downy mildew, other types of regulation of this character, such as complementarity and epistatic relationships, do occur.     

RAPD markers linked to resistance to downy mildew disease in soybean

To determine and utilize RAPD markers linked to resistance to downymildew incited by Peronospora manshurica in soybean, a resistantcultivar `AGS129' was crossed to a susceptible cultivar `Nakhon Sawan 1'(NS1). F₂ and BC₁ populations were advanced from the F₁ and evaluatedfor resistance to the disease. ²-test demonstrated that the resistancewas controlled by a single dominant gene (Rpmx). Near-isogenic lines(NILs) and bulked segregant analysis (BSA) were used to identify RAPDmarkers linked to the gene. Six DNA bulks namely F₅(R), F₅(S),BC₆F₃(R), BC₆F₃(S), F₂(R) and F₂(S) were set up by pooling equalamount of DNA from 8 randomly selected plants of each disease responsetype. A total of 180 random sequence decamer oligonucleotide primerswere used for RAPD analysis. Primer OPH-02 (5 TCGGACGTGA 3 andOPP-10 (5 TCCCGCCTAC 3) generated OPH-02₁₂₅₀ and OPP-10₈₃₁fragments in donor parent and resistant bulks, but not in the recurrentparent and susceptible ones. Co-segregation analysis using 102 segregatingF₂ progenies confirmed that both markers were linked to the Rpmxgene controlling downy mildew disease resistance with a genetic distance of4.9 cm and 23.1 cm, respectively. Marker OPH-02₁₂₅₀ was presentin 13 of 16 resistant soybean cultivars and absent in susceptible cultivars,thus confirming a potential for MAS outside the mapping population.