Inheritance of resistance to lettuce root aphid in the lettuce cultivars 'Avoncrisp' and 'Lakeland"

The inheritance of resistance in two lettuce cultivars to lettuce root aphid, Pemphigus bursarius, was studied in a series of laboratory and field experiments at Wellesbourne between 1989 and 1992. A source of total resistance in the cv. ‘Avoncrisp’ which is linked to the downy mildew resistance gene Dm6, was shown to be governed by a single dominant gene. There were no maternal effects evident in the inheritance of this resistance. The basis of the high level of resistance which exists in the cv. ‘Lakeland’ (formerly known as ‘Jubilee’) was also shown to be controlled by the same dominant gene. The linkage between Dm6 and root aphid resistance was broken in ‘Lakeland’ as this cultivar does not possess the Dm6 gene. The linkage was presumably broken when the original cross between the parents of cv. ‘Lakeland’, ‘Calmar’ and ‘Avoncrisp’ was made. Under laboratory conditions small numbers of aphids commence development on cv. ‘Lakeland’ but colonies fail to develop and under field conditions the resistance provides adequate Protectión against the pest. The resistance in both ‘Avoncrisp’ and ‘Lakeland’ was effective against a population of lettuce root aphid collected from an endive crop in southern France as well as being effective against the Wellesbourne population of this aphid.     

Inheritance of partial resistance to the leaf aphids Macrosiphum euphorbiae and Uroleucon sonchi in lettuce

The genetics of partial resistance (PR) of lettuce to the leaf aphid species Macrosiphum euphorbiae and Uroleucon sonchi was studied in F₁ and F₂ generations of six crosses between four cultivars. Partial resistance inherited mainly additively. With M. euphorbiae the estimates of broad-sense F₂ heritability were low and non-significant in crosses between parents of the same resistance level (S x S and PR x PR), and high (0.69-0.76) in crosses between parents of different resistance level (PR x S). The level of resistance in lettuce cultivars ‘Charan’ and ‘Marbello’ led to a marked reduction in percentage of plants with colonies of M. euphorbiae in a field experiment. With U. sonchi the highest estimates of F₂ heritability were found in ‘Charan x ‘Avoncrisp’ (0.51) and ‘Charan’ x ‘Chou de Naples' (0.48). In the cross ‘Charan’ x ‘Marbello’ (PR x PR) no significant segregation for resistance to M. euphorbiae was found, indicating that these cultivars may have the same genetic constitution for resistance to M. euphorbiae.‘Charan’ has additional genes for resistance to U. sonchi, that are not, or only weakly linked to the gene(s) for resistance to M. euphorbiae. The heritability estimates for the partial resistance to M. euphorbiae and U. sonchi indicate that it is possible to effectively select for these traits in plant breeding programmes.     

Intensity of radicle fluorescence as related to the resistance of seedlings of lettuce to the lettuce root aphid and carrot to the carrot fly

A technique is described which indicates by u.v. fluorescence the concentration of caffeoylquinic, chlorogenic and isochlorogenic acids in roots of germinating lettuce and carrot. The surface fluorescence of radicles from lettuce root aphid (Pemphigus bursarius) resistant seedlings was more intense than for susceptible seedlings, attributable to the higher concentration of isochlorogenic acid. In contrast, radicles of carrot seedlings resistant to carrot fly (Psila rosae) were less fluorescent than susceptible seedlings, corresponding to the lower concentration of chlorogenic acid in resistant seedlings. The technique is non-destructive and has been developed to distinguish between cultivars or breeding lines potentially resistant to these insect pests.     

Inheritance of downy mildew resistance, β-1,3-glucanases and peroxidases in pearl millet [Pennisetum glaucum (L.) R. Br.] crosses

The inheritance of resistance to downy mildew disease and the defense-related enzymes β-1,3-glucanase and peroxidase was studied in crosses of pearl millet using a generation-mean analysis. The study material comprised six generations (susceptible and resistant parents, F₁, F₂, BC₁ and BC₂) in three crosses. Seedlings from these generations were inoculated with the downy mildew pathogen Sclerospora graminicola and disease incidence was recorded. Analysis of constitutive levels of β-1,3-glucanase and peroxidase in the seedlings of different generations indicated that the resistant populations showed higher enzyme activities, while lower activities of the enzymes were recorded in the susceptible populations. In the generation-mean analysis, the significance of scaling tests revealed the existence of non-allelic interactions in the inheritance of resistance to downy mildew as well as with the enzymes. Among the gene effects, both additive and dominant effects were significant. All the non-allelic interaction effects were significant in the crosses. Studies on the isozyme patterns of the enzymes substantiated the results of the disease-incidence experiments in most of the generations. The results indicated that the inheritance of downy mildew disease resistance and the expression of β-1,3-glucanase and peroxidase in pearl millet is not only under the control of additive and dominant genes but are also governed by complex non-allelic interactions. Received: 30 April 2000 / Accepted: 17 October 2000     

Studies on inheritance of resistance and allelic relationships for strain 2 of pigeonpea sterility mosaic pathogen

Inheritance of resistance and allelic relationships were studied in three resistant pigeonpea sources for strain 2 of sterility mosaic pathogen. The resistant genotypes (ICP 7035, ICP 7349 and ICP 8850) were crossed with susceptible genotypes (BDN1 and LRG30) to determine the inheritance of resistance. The resistant and susceptible genotypes were also crossed among themselves to obtain information on their allelic relationships. Parents, F₁ and F₂ generations were sown in pots and screened using infector-hedge technique. Observations in parents, F₁ and F₂ generations, indicated dominance of resistance in certain crosses and the dominance of susceptibility in others. Disease reaction appeared to be governed by two independent non-allelic genes, with at least three multiple alleles, at one of the loci.     

Inheritance of resistance to leaf and glume blotch caused by Septoria nodorum Berk. in winter wheat

Sixteen crosses between eight winter wheat cultivars were screened for resistance to Septoria nodorum leaf and glume blotch in the F₁ and F₄ generations using artificial inoculation in the field. The F₁ of most crosses showed dominance for susceptibility on both ear and leaf. The effects of general combining ability were of similar magnitude as the effects for specific combining ability. On the basis of the phenotypic difference of the parents, no prediction was possible about the amount and the direction of genetic variance in the segregating populations. The variation observed in this study both within and among the segregating populations suggests a quantitative inheritance pattern influencing the expression of the two traits. The components of variance between F₂ families within a population were as high as (for S. nodorum blotch on the ear) or higher (for S. nodorum blotch on the leaf) than those between populations. Therefore, strong selection within a few populations may be as effective to obtain new resistant genotypes as selection in a large number of populations. In almost all crosses, progenies were found that were more resistant than the better parent. Thus transgression breeding may be a tool to breed for higher levels of resistance to S. nodorum blotch. Highly resistant genotypes were found even in combination with two susceptible parents. The genetic source for Septoria resistance is probably broader than is generally assumed and could be used to improve S. nodorum resistance by combination breeding followed by strong selection in large populations. Received: 18 January / Accepted: 30 April 1999     

Complementarity of genes for resistance to greenbug [Schizaphis graminum (Rondani)], biotype E,in sorghum [Sorghum bicolor (L.) Moench]

Summary Gene complementarity among various sources of resistance to greenbug biotype E was assessed. Analysis of the F₂ generation of crosses between susceptible and resistant parents (mating 1) and among sources of resistance (mating 2) suggested that resistance in sorghum to greenbug biotype E was complexly inherited and, to some extent, dependent on the nature of both the resistant and susceptible parents. Positive transgressive segregation in the F₂ generations of both matings was found to be due to effective plus factors, contributed by both parents in a cross, which complemented each other. The number of plus factors ranged from one to two in the susceptible parents and from two to five in the resistant parents of mating 1, and from one to five in the parents of mating 2. The consistently significant reciprocal effects shown by Sarvasi and PI264453 indicated that these sources had major factors for resistance in their cytoplasms, which were expressed in all their crosses. The results from this study indicated that the sources of resistance complemented each other to give increased number of F₂ segregates with increased resistance. Thus, it should be possible to increase and diversify resistance of sorghum to greenbug biotype E by accumulating different, effective plus factors from various sources through recurrent selection.Contribution No. 90-106-J from the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, KS 66506, USA     

Inheritance of resistance to the cowpea aphid in cowpea

Summary Inheritance of resistance to cowpea aphid, Aphis craccivora Koch, in three resistant cultivars of cowpea, Vigna unguiculata (L.) Walp, was studied. The parents, F₁ and F₂ population were grown in an insect-proof screenhouse. Each 3-day-old seedling was infested with 10 apterous adult aphids. Seedling reaction was recorded when the susceptible check was killed. The segregation data revealed that the resistance of ICV11 and TVU310 is governed by single dominant genes. All the F₂ seedlings of the cross ICV10xTVU310 were resistant, indicating that they have the same gene for resistance. However, the F₂ populations from the crosses ICV10xICV11 and ICV11xTVU310 segregated in a ratio of 151, indicating that the dominant genes in ICV11 and TVU310 are non-allelic and independent of each other. The resistance gene of ICV10 and TVU310 is designated as Ac₁ and that of ICV11 as Ac₂.     

Foraging ofRhagoletis pomonella flies in relation to interactive food and fruit resources

Feeding behaviour of the lettuce root aphidPemphigus bursarius was monitored electronically on six lettuce varieties. Aphids, monitored for a minimum of 5 h on lettuce roots, performed recognizable feeding behaviour on the susceptible varieties, Webbs Wonderful and Borough Wonder including non probing, pattern A start of new penetration, C (stylet pathway activities) sometimes including clear B-waves (salivary sheath secretions), potential drops (intracellular penetrations), intracellular E (phloem ingestion) distinguishing E₁ and E₂ (Tjallingii, 1990) and occasionally xylem ingestion. Although aphid probing was recorded on the resistant varieties Lakeland and Grand Rapids continued penetration to the phloem elements was deterred and the time spent ingesting phloem was short. Aphids rarely probed the resistant varieties Avoncrisp and Avondefiance and all attempted penetrations quickly terminated.     

VARIETAL RESISTANCE OF LETTUCE TO ATTACK BY THE LETTUCE ROOT APHID, PEMPHIGUS BURSARIUS (L.)

Striking varietal differences in susceptibility to attack by the lettuce root aphid, Pemphigus bursarius L., were first found in lettuces grown at Wellesbourne in 1955. Subsequent work has confirmed that White Favourite and Imperials E-4 and 19551-M are highly resistant; Imperial 45634-M, Continuity and Iceberg are markedly resistant. Midas, Salad Bowl and Imperial 4164 each appear to be a mixture of susceptible and resistant plants.
Immigrant winged forms of P. bursarius showed no preference for colonizing any particular variety of lettuce, and it seems that resistance to attack results from antibiosis.
It is suggested that varietal differences in the composition of root sap may account for the differences in susceptibility to lettuce root aphid, but analyses of water and alcohol extracts of root sap from resistant and susceptible lettuce varieties have not shown consistent differences.     

Effectiveness of resistance genes to the large raspberry aphid, Amphorophora idaei Börner, in different raspberry (Rubus idaeus L.) genotypes and under different environmental conditions

The introduction into commerce of raspberry cultivars with major gene resistance to the large raspberry aphid, Amphorophora idaei, an important pest and virus vector on red raspberry in Europe, has been very effective both in decreasing pest numbers and greatly restricting infection with the viruses it transmits. However, biotypes of the aphid able to overcome these genes have developed in the field in recent years. Additionally, in field and laboratory tests, the response to aphid biotypes and recognised aphid strains of certain raspberry cultivars, such as Glen Prosen and Delight, differ markedly despite the fact that they are reputed to contain the same A. idaei‐resistance gene, A₁. In attempts to understand the reasons for this difference in response, analysis was made of the segregation of progeny seedlings from crosses between A. idaei‐resistant and ‐susceptible cultivars to two recognised strains of the aphid. These studies showed that, as expected, cv. Autumn Bliss contained the A. idaei‐resistance gene, A₁₀, and cvs Delight and Glen Prosen each contained the A. idaei‐resistance gene, A₁. When progeny seedlings were assayed in a heated glasshouse as young plants and in an unheated Tygan house as 1 m tall plants, the segregation ratios for resistance and susceptibility to A. idaei were largely unchanged. However, when the resistance of individual progeny plants was assessed, c. 37% of the putative gene A₁‐containing progeny and 9–23% of the putative gene A₁₀‐containing progeny, behaved differently in these two environments. Experiments involving an A. idaei‐resistant and ‐susceptible parent cultivar showed that shading plants increased their susceptibility to A. idaei colonisation. Whilst this shading effect has implications for experimentally detecting A. idaei‐resistant progeny in segregating raspberry seedlings, it does not explain the difference in field resistance to A. idaei of cvs Delight and Glen Prosen. Such differences in the field seem best explained by the presence in these cultivars of ‘minor’ genes for A. idaei resistance and/or susceptibility that influences the effectiveness of gene A₁.     

Inheritance and mechanism of resistance to herbicides inhibiting acetolactate synthase in Sonchus oleraceus L.

A biotype of Sonchus oleraceus L. (Compositae) has developed resistance to herbicides inhibiting acetolactate synthase (ALS) following field selection with chlorsulfuron for 8 consecutive years. The aim of this study was to determine the inheritance and mechanism of resistance in this biotype. Determination of ALS activity and inhibition kinetics revealed that K_m and V_max did not vary greatly between the resistant and susceptible biotypes. ALS extracted from the resistant biotype was resistant to five ALS-inhibiting herbicides in an in vitro assay. ALS activity from the resistant biotype was 14 19, 2, 3 and 3 times more resistant to inhibition by chlorsulfuron, sulfometuron, imazethapyr, imazapyr and flumetsulam, respectively, than the susceptible biotype. Hybrids between the resistant and a susceptible biotype were produced, and inheritance was followed through the F₁, F₂ and F₃ generations. F₁ hybrids displayed a uniform intermediate level of resistance between resistant and susceptible parents. Three distinct phenotypes, resistant, intermediate and susceptible, were identified in the F₂ generation following chlorsulfuron application. A segregation ratio of 121 was observed, indicative of the action of a single, nuclear, incompletely dominant gene. F₃ families, derived from intermediate F₂ individuals, segregated in a similar manner. Resistance to herbicides inhibiting ALS in this biotype of S. oleraceus is due to the effect of a single gene coding for a resistant form of the target enzyme, ALS.     

Nature and type of gene action governing resistance to Helminthosporium turcicum leaf blight in maize (Zea mays L.)

Six generations, consisting of three resistant parents, three susceptible parents, their 15 possible F₁ crosses, 15 F₂'s, 15 BC₁'s (F₁ x resistant female parent) and 15 BC₂'s (F₁ x susceptible male parent) were analysed following Hayman (Heredity 12: 371–390, 1958) to evaluate the nature and type of gene action governing resistance to H. turcicum. The results showed that all types of gene effects, viz., additive, dominance and epistasis (i.e., additive x additive, additive x dominance and dominance x dominance) were operating in one cross or the other in controlling resistance. However, it was additive gene action and dominance x dominance type of epistasis with duplicate nature that were important in controlling resistance in most crosses. Depending upon the final objectives, one of the breeding methods, viz., recurrent selection, heterosis breeding, back-cross method or full-sib selection (bi-parental mating) may be followed.     

Inheritance of adult field resistance to yellow rust disease among broad-based hexaploid spring wheat germplasm

 Complete F₁ and F₂ diallel crosses were used to investigate the inheritance of yellow rust resistance among eight bread wheat lines, developed by CIMMYT for the East African Highlands, which showed a wide response to this disease. Both diallel sets were grown at a site with a high incidence of yellow rust, although for one season, during which the F₁ diallel was grown, disease incidence was unusually low. Analyses disclosed the presence of additive, dominance and epistatic effects among those genes controlling rust resistance, with the former being the most important. At normal disease levels, excluding two arrays having resistant common parents removed non-allelic interactions from the F₁ diallels. For all F₂ diallels, and the remaining F₁ diallel, omitting two arrays based on susceptible parents removed these interactions. Local selection of material from a broadly based germplasm appears to be a feasible method of developing adapted cultivars resistant to endemic diseases. Received: 1 March 1998 / Accepted: 19 March 1998     

Inheritance of bipyridyl herbicide resistance in Arctotheca calendula and Hordeum leporinum

The mode of inheritance of resistance to bipyridyl herbicides in bipyridyl-resistant biotypes of Arctotheca calendula and of Hordeum leporinum was investigated. F₁ plants from reciprocal crosses between diquat-resistant and -susceptible plants of A. calendula showed an intermediate response to diquat application that was nuclearly inherited. Treatment of F₂ plants with 100 g ai ha^-1 of diquat or 800 g ai ha^-1 of paraquat killed all homozygous-susceptible plants, caused severe injury to heterozygous plants but only slight or no injury to homozygous-resistant plants. Back crosses of F₁ to susceptible plants exhibited intermediate and susceptible phenotypes. The observed segregation ratios in F₂ and test-cross populations fitted predicted segregation ratios, 1:2:1 (R:I:S) and 1:1 (I:S) respectively, showing that bipyridyl resistance is conferred by a single incompletely-dominant gene. Biotypes of paraquat-resistant and -susceptible H. leporinum were crossed reciprocally. F₁ plants from reciprocal crosses showed an intermediate response to paraquat application. The F₂ progeny showed segregation ratios that fitted the predicted segregation ratio of 1:2:1 (R:I:S) forinheritance of resistance being governed by a single partially-dominant gene.     

The genetics of race specific resistance in lettuce (Lactuca sativa) to downy mildew (Bremia lactucae)

Data are presented on the segregation of resistance to five British races and two Dutch races of Bremia lactucae in the F₂ progenies of crosses involving seven resistant and several susceptible lettuce cultivars. These data and also those previously published by other workers are considered in relation to the systematic model proposed by Crute & Johnson (1976) to explain the genetics of race specific resistance to B. lactucae in lettuce. It is shown that, with minor modifications, the model accommodates almost all of the previously published data and correctly predicts the new data, except for one set which cannot at present be interpreted. It is concluded that genetic evidence exists for the presence, among various cultivars of lettuce, of at least four and possibly five different dominant resistance genes of major effect designated Dm₂, Dm₃, Dm₄, Dm₆ and Dm₈; and of a pair of dominant genes with complementary effect designated Dm7/1 and Dm7/2. The resistance conferred by these genes is specified in relation to five British races, five Dutch, three Israeli and one United States race of the fungus. Resistance genotypes are proposed for cultivars Avoncrisp, Avondefiance, Calmar, Great Lakes 659, Kares, Meikoningen, Mildura, Proeftuins Blackpool, Solito, Valverde, Ventura and the USDA line PI 164937.     

Paraquat resistance in a Lolium rigidum population is governed by one major nuclear gene

Paraquat resistance in an annual ryegrass (Lolium rigidum Gaud.) population (AFLR1) has been attributed to reduced paraquat translocation. Genetic inheritance of paraquat resistance in this population was investigated in the present study. The paraquat dose response of progeny from 8 F₁ families was more similar to that of the resistant than the susceptible parent, while the equivalent data for a further three families were intermediate compared to those of the parental populations. No significant differences in dose response were observed between reciprocal crosses of specific F₁ families. These results suggest that paraquat resistance in AFLR1 is inherited as a dominant or partially dominant nuclear-encoded trait. Pseudo-F₂ (ψ-F₂) generation seedlings were treated with multiple dose rates sufficient to control the susceptible parental population, and observed segregation ratios in all instances conformed to a 3:1 (resistant:susceptible) segregation ratio, and this ratio was further confirmed by individual phenotyping of cloned plant genotypes. A single major nuclear gene is hence apparently responsible for evolved paraquat resistance in AFLR1.     

The inheritance of resistance to Rhynchosporium secalis in some European spring barley cultivars

The inheritance of partial resistance to Rhynchosporium secalis, which appears to be race non-specific, was studied in a diverse range of European spring barley cultivars. Data from the F₂ generation of a 6 × 6 diallel cross and the F₃ generation of three crosses selected from this diallel set suggested that resistance was complex in inheritance, the results being incompatible with any hypothesis involving less than four genes. The F₂ studies indicated that both dominant and recessive genes were active in conferring resistance, and that there were significant additive gene effects. Transgressive segregation occurred in all cross combinations in the F₃ material. Consequently the alleles conferring resistance were not completely concentrated in the most resistant cultivar studied (cv. Proctor). Heritability estimates obtained from F₂ and F₃ material suggested that field selection was of limited reliability, even when spreader drills were incorporated amongst the segregates. Single-plant selection (in F₂) was considered to be of little value, and the results of F_a head-row tests would require confirmation by replicated tests in subsequent generations.     

Inheritance of Cry1Ac resistance and associated biological traits in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

The analysis of reciprocal genetic crosses between resistant Helicoverpa armigera strain (BH-R) (227.9-fold) with susceptible Vadodara (VA-S) strain showed dominance (h) of 0.65-0.89 and degree of dominance (D) of 0.299-0.782 suggesting Cry1Ac resistance as a semi-dominant trait. The D and h values of F₁ hybrids of female resistant parent were higher than female susceptible parent, showing maternally enhanced dominance of Cry1Ac resistance. The progeny of F₂ crosses, backcrosses of F₁ hybrid with resistant BH-R parent did not differ significantly in respect of mortality response with resistant parent except for backcross with female BH-R and male of F₁ (BH-R × VA-S) cross, suggesting dominant inheritance of Cry1Ac resistance. Evaluation of some biological attributes showed that larval and pupal periods of progenies of reciprocal F₁ crosses, backcrosses and F₂ crosses were either at par with resistant parent or lower than susceptible parent on treated diet (0.01 μg/g). The susceptible strain performed better in terms of pupation and adult formation than the resistant strain on untreated diet. In many backcrosses and F₂ crosses, Cry1Ac resistance favored emergence of more females than males on untreated diet. The normal larval period and the body weight (normal larval growth) were the dominant traits associated with susceptible strain as contrast to longer larval period and the lower body weight (slow growth) associated with resistance trait. Further, inheritance of larval period in F₂ and backcross progeny suggested existence of a major resistant gene or a set of tightly linked loci associated with Cry1Ac sensitivity.     

Inheritance of potato aphid resistance in hybrids between Lycopersicon esculentum and L. pennellii

Summary The potato aphid, Macrosiphum euphorbiae Thomas, is an important pest of tomato, Lycopersicon esculentum Mill., because it transmits tomato viruses and directly reduces crop yields by its feeding. This study was conducted to determine whether the wild tomato species, Lycopersicon pennellii (Corr.) D'Arcy, would be useful as a source of potato aphid resistance for tomato. Type IV trichome density and aphid resistance were assessed in six generations (P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂) from crosses between L. pennellii (LA 716) and two tomato cultivars, New Yorker and VF Vendor. Weighted leastsquares were used in joint scaling tests to estimate the relative importance of gene effects on type IV trichome density and potato aphid resistance of the hybrids. A simple additive-dominance model adequately explained the variation in type IV trichome density. Models which included digenic epistatic effects were required to explain the variation in aphid resistance. Standard unit heritability estimates of aphid resistance in the backcross to L. esculentum were obtained by regression of BC₁F₂ off-spring families on BC₁F₁ parents. Regression coefficients and heritability estimates varied between years with the level and uniformity of the aphid infestation. In the 1985–1986 growing seasons, when aphid infestations were uniform, aphid resistance exhibited a moderate level of heritability (29.8% ± 14.1% and 47.1% ± 11.5% in New Yorker and VF Vendor backcross populations, respectively). The non-uniform aphid infestation of 1984 resulted in lower heritability estimates in the 1984–1985 growing seasons (16.1% ± 15.7% and 21.9% ± 14.8% in the New Yorker and VF Vendor backcross populations, respectively). Selection for potato aphid resistance would probably be most efficient if it were delayed until gene combinations are fixed in later generations, because of the large epistatic effects and the low heritability of this trait in seasons with variable aphid infestations.