Congruence of conventional and molecular studies to locate genes that control flavone synthesis in maize silks

The presence of maysin, a flavone glycoside, and its analogues in the silks of corn is an important defence against invasion of the ear by corn earworm in the southeastern USA. Three dent maize inbreds with high silk‐maysin concentrations were evaluated for chromosomal location of major genes controlling synthesis of three antibiotic chemicals by crossing them to a series of waxy‐marked chromosome 9 reciprocal translocations. The data indicated that genes for maysin and its analogues are probably present on the short arms of chromosomes 1 and 10, and long arms of chromosomes 5 and 8 for inbred line GT114, the short arms of chromosome 1 and 6, and the long arms of chromosome 5 and 8 for inbred line GE37, and the short arms of chromosomes 1 and 10, and the long arm of chromosome 4 for inbred line SC102. These results are in general agreement with other translocation studies on corn earworm injury to sweet corn inbreds and gene and restriction fragment length polymorphism marker studies to locate quantitative trait loci (QTL) for maysin synthesis, with the exception that QTL on chromosome 9 have been found to be strongly associated with flavone synthesis. The most significant concordance between conventional and molecular techniques for locating chromosome regions influencing synthesis of antibiotic chemicals from silks of high silk antibiosis sources is found for the short arm of chromosome 1. This region is the most likely one on which to place emphasis during the initial stages of transferring high‐maysin silks to elite inbred lines. A chromosome 1 marker plus at least one more from any chosen high‐maysin inbred source should be sufficient to transfer high maysin silks to other lines. Other markers may be needed for transfer of specific traits when recovering recurrent parent genome types in a backcrossing procedure.     

Recurrent selection for maysin, a compound in maize silks, antibiotic to earworm

A thorough knowledge of the inheritance of maysin, a flavone glycoside with antibiosis to corn earworm, Helicoverpa zea Boddie, in the silks of maize will assist breeders in choosing the most efficient method of incorporating this trait into elite inbreds. Two breeding populations, one having exotic origin (EPM), the other from southern inbred lines (SIM) were subjected to six cycles of recurrent selection for increased silk maysin. Ten per cent of the individuals evaluated in populations were selected for recombination as S₁ progenies in each cycle. Progress was evaluated as C0 to C6 population cycles in a randomized complete‐block experiment with five replications in 1997 and 1998. Maysin fresh weights in silks of the C0 populations were 0.54% for EPM and 0.36% for SIM. Responses of 0.19% per cycle in EPM and 0.22% per cycle in SIM resulted in an EPM (C6) population with 1.76% maysin and an SIM (C6) population with 1.69% maysin. Silk maysin concentrations above 0.2% begin to substantially reduce larval growth and prevent completion of the life cycle when husk coverage is sufficient to force the insect to feed on silks while entering the ear. Chlorogenic acid and two analogues of maysin, apimaysin and 3′‐methoxymaysin, were found in such minor quantities in silks that they could not be credited with any impact on antibiotic activity against the insect. Selection has effectively increased silk maysin concentration in both EPM and SIM. Trait responses for maysin are highly heritable and will allow the plant breeder to introgress resistance to the corn earworm into elite material easily.     

Inheritance of partial rust resistance in pearl millet

Quantitative disease resistance should be exploited to complement the use of genes for qualitative or hypersensitive resistance. The expression and inheritance of partial rust resistance of pearl millet inbreds 700481-21-8 and ‘ICMP 501’ crossed to moderately susceptible Tift 383’ were evaluated in seedling assays in the greenhouse and in generation mean and single-seed descent populations in the field. Uredinium sizes on seedling leaves of hybrids were generally intermediate to those of the parental inbreds and consistent differences could be discerned in uredinium lengths. Area under the disease progress curves (AUDPCs) of individual plants of the parents, F₁, F₂, and backcross F₁S to each parent were determined from field trials. Broad-sense heritability estimates for both crosses were 43%. In generation mean analyses, additive genetic effects were significant in the cross of 700481–21–8 × Tift 383′, whereas additive, dominance, and dominance × dominance epistatic effects were significant for ‘ICMP 501’בTift 383’. The number of genes conferring partial resistance was estimated to be two for 700481–21–8 and 2.5 for ‘ICMP 501’. A hierarchical single-seed descent analysis revealed significant differences in AUDPC among F₃-derived F₄ progenies in the F₆ generation. Selection for progenies with greater resistance should be possible among F₄ families. Higher levels of resistance were observed in progeny derived from ‘ICMP 501’. Because segregation of resistance differed among progeny derived from 700481–21–8 and ‘ICMP 501’, the genetic basis for resistance probably differs between the two inbreds.     

Inheritance of tillering and flowering-time in early maturing maize

Summary A diallel cross and F₂ populations derived from eight early maturing maize inbreds were used to investigate the inheritance of tillering and flowering-time (anthesis), and the possible relationship between tillering and flowering-time. Incomplete dominance for increased tillering was observed; potence ratios, representing the overall degree of dominance, ranged from 0.26 to 0.52. Dominance for early flowering ranged from incomplete with a potence ratio of 0.55 to overdominance with a potence ratio of 1.40. Broad-sense heritabilities were low for both characters. The genetic component of variation for tillering was due to general combining ability effects; specific combining ability effects were not significant. A significant negative linear relationship between tillering and flowering-time was found. Lack of independent assortment of tillering and flowering-time in F₂ populations indicated that the two characters are genetically related.     

Genetic and statistical models for estimating genetic parameters of maize seedling resistance to Fusarium graminearum Schwabe root rot

Root lodging is an important problem in corn fields. Fungi recovered from roots include seedling blight and stalk rot pathogens. The objective of this work was to study the inheritance of maize seedling resistance to pathogens causing maize lodging. The Fusarium graminearum strain, 241 Fr1, was isolated from maize lodged plants and identified as the most pathogenic isolate for root rotting. Nine inbred lines of maize and their diallel F₁ crosses plus control genotypes were studied. Seedlings were inoculated at the stage of four-leaves. Disease severity was measured as percentage of the root rotted area. Highly significant differences between inoculated and non-inoculated genotypes were found. Four genetic models and two statistical approaches—the mixed model for the best linear unbiased prediction (BLUP) and the general linear model (GLM)—were used for the analysis. Favorable heterosis of resistance of hybrids over inbreds was the most important effect detected. The general combining ability (GCA) effects were significant for all genetic models and statistical methods studied, and a good agreement existed among the GCA estimates by the different methods. The type of gene action, either additive or dominance, showed a large variation among the parental inbreds and hybrids. Selection of additive effects based exclusively on inbred lines is not sufficient to confer resistance to hybrids, additional selection should be practiced on hybrids to look for favorable dominance effects. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic analysis of prolificacy in “Sikkim Primitive”—A prolific maize (Zea mays) landrace of North‐Eastern Himalaya

Prolificacy assumes significance for development of high‐yielding baby corn hybrids. “Sikkim Primitive” is a native landrace of North‐Eastern Himalaya, and is the highest prolific maize germplasm. So far, the genetics of prolificacy in “Sikkim Primitive” has not been deciphered. Here, a prolific inbred (MGU‐SP‐101) developed from “Sikkim Primitive” was crossed with four non‐prolific inbreds viz., LM13, BML7, HKI161 and HKI1128. Six generations (P₁, P₂, F₁, F₂, BC₁P₁ and BC₁P₂) of the crosses were evaluated at two locations during rainy season 2018. MGU‐SP‐101 possessed 2.50–3.78 ears per plant compared to 1.06–1.86 among non‐prolific inbreds. The variation for ears was the highest in F₂s (1–8), followed by BC₁P₁ (1–7) and BC₁P₂ (1–5). The quantitative inheritance pattern of prolificacy with prevalence of non‐allelic interactions of duplicate epistasis type has been observed. Dominance × dominance effect was predominant over additive × additive and additive × dominance effects. Total number of major gene blocks ranged from 0.41 to 2.86, thereby suggesting the involvement of at least one major gene/QTL governing the prolificacy. This is the first report of genetic dissection of prolificacy in “Sikkim Primitive”.     

Genetics of spike length in durum wheat

Gene effects were analyzed using mean spike length of 12 populations, viz., both parents, F₁, F₂, first back cross generation, BC₁ and BC₂, second backcross generations, BC₁₁,BC₁₂, BC₂₁ and BC₂₂ along with BC₁ self and BC₂ self derived by selfing BC₁ and BC₂populations of three crosses involving six diverse cultivars of Triticum durumto determine the nature of gene actions governing spike length through generation mean analysis under normal and late sown environments. The six-parameter model was adequate in most of the cases to explain genetic variation among the generation means under both the sowing environments. Additive (d) gene effect was significant in all the cases, whereas dominance (h) gene effect was not so frequently observed significant. Epistatic effects, particularly digenic types were predominant over additive and dominance effects in most of the cases under both normal and late sown environments except in the cross Cocorit 71 × A-9-30-1 (normal sown).Additive × dominance × dominance (y), trigenic interaction played significant role in controlling the inheritance of this trait in the cross HI 8062 × JNK-4W-128under late sown condition. Duplicate epistasis was observed in the cross HI 8062× JNK-4W-128 (normal sown). Non-fixable gene effects were of higher magnitude than fixable gene effects in almost all cases, confirmed the major role of non-additive gene effects to control the inheritance of spike length in durum wheat. Significant heterosis over better parent was not observed. Similarly, inbreeding depression was not commonly observed. Favourable and suitable environment must be considered before finalizing breeding programme for its simple inheritance to get desirable improvement for high grain yield. Hybridization systems, such as biparental mating and / or diallel selective mating, which exploit both additive and non-additive gene effects, simultaneously, could be useful in the improvement of spike length in durum wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of β-carotene concentration in durum wheat (Triticum turgidum L. ssp. durum)

Despite being one of the important characteristics in determining pasta quality in durum wheat (Triticum turgidum ssp. durum), there is no direct report on inheritance of β-carotene concentration. The objectives of this study were to determine the inheritance of β-carotene concentration and the number of genes involved in six crosses of durum. For the cross PDW-233 (P₁) × Bhalegaon-4 (P₂), F₁, F₂, BCP₁ and BCP₂ populations were developed. For all other crosses, only the F₁ and F₂ populations were developed. β-carotene concentration was determined for all populations and parents of each cross grown at Hol, Maharastra, India. The cross PDW-233 × Bhalegaon-4 was also evaluated at Dharwad, Karnataka, India. Low β-carotene concentration was partially dominant in most of the crosses. Broad sense heritability was 67 and 91% at Dharwad and Hol, respectively, for the cross PDW-233 × Bhalegaon-4 and varied from 74 to 93% for the other five crosses indicating the presence of additive gene effects. The frequency distributions of the trait in the F₂ populations were not normal and were skewed towards the lower parent. Segregation of β-carotene concentration in the six F₂ populations indicated that at least two major genes and two or three minor genes with modifying effects govern the trait. Analysis of variance indicated that environment had comparatively little influence on the trait and this should allow for easy selection. The joint scaling test revealed additive × additive, additive × dominance and dominance × dominance epistatic interactions in the cross PDW-233 × Bhalegaon-4. These authors contributed equally.     

Genetics of resistance to early blight (Alternaria solani Sorauer) in tomato (Lycopersicon esculentum L.)

The genetic nature of early blight resistance in tomato was studied in three crosses at seedling and adult plant stages. A six generation mean analysis of the cross Arka Saurabh (susceptible) × IHR1939 (resistance) and its reciprocal cross revealed that the resistance to early blight was conferred by recessive polygenes at both seedling and adult plant stages. This polygenic early blight resistance revealed the importance of additive and additive × additive gene effects at seedling stage and higher magnitude of dominance and dominance× dominance gene effects at adult plant stage. Evaluation of parents, F₁, F₂ and backcross generations of IHR1816 (resistance) × IHR1939 (resistance) revealed that the early blight resistance genes in IHR1816 (Lycopersicon esculentum NCEBR-1) and IHR1939 (Lycopersicon pimpinellifolium L4394) are independent. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of the stay-green trait in tropical maize

Stay-green maize genotypes have been associated with tolerance to biotic and abiotic stresses, including tolerance to drought, and to stalk and root lodging, but there is limited information on its inheritance. Thus, this research was conducted to study the inheritance of the stay-green trait using both conventional analysis and QTL mapping of the Design III in a tropical maize population developed from two inbred lines genetically divergent for this trait. Two-hundred and fifty F₂ plants were genotyped with 177 microsatellite markers, and their backcrossed progenies to both parental inbreds were evaluated at three locations. Ten plants per plot were assessed 120 days after sowing and the plot means scores for stay-green, adjusted for days to silking emergence, were used for analysis. The additive variance was larger than the dominance variance, the genetic by location interaction variance presented a high magnitude, and the heritability coefficient on a plant-basis a low magnitude. Seventeen QTL were mapped, most of them were clustered on four chromosomes and accounted for by 73.08 % of the genetic variance. About half of the QTL interacted with location, and the average level of dominance was partial dominance. The additive effects were larger than the dominance effects; the latter were not unidirectional, so that heterosis could not be exploited in crosses. Procedures for marker-assisted selection to increase the level of stay-green are discussed and an approach is suggested for using both stable and non-stable QTL in a marker-assisted backcross program.     

Inheritance of time to flowering in cowpea (<Emphasis Type="Italic">Vigna unguiculata</Emphasis> (L.) Walp.)

Time to flowering and maturity is an important adaptive feature in annual crops, including cowpeas (Vigna unguiculata (L.) Walp.). In West and Central Africa, photoperiod is the most important environmental variable affecting time to flowering in cowpea. The inheritance of time from sowing to flowering (f) in cowpeas was studied by crossing a photoperiod-sensitive genotype Kanannnado to a photoperiod-insensitive variety IT97D-941-1. Sufficient seed of F₁, F₂, F₃ and backcross populations were generated. The parental, F₁, F₂, F₃ and the backcross populations were screened for f under long natural days (mean daylength 13.4 h per day) in the field and the parents, F₁, F₂ and backcross populations under short day (10 h per day) conditions. The result of the screening showed that photoperiod in the field was long enough to delay flowering of photoperiod-sensitive genotypes. Photoperiod-sensitivity was found to be partially dominant to insensitivity. Frequency distribution of the trait in the various populations indicated quantitative inheritance. Additive (d) and additive × dominance (j) interactions were the most important gene actions conditioning time to flowering. A narrow sense heritability of 86% was estimated for this trait. This will result in 26 days gain in time to flowering with 5% selection intensity from the F₂ to F₃ generation. At least seven major gene pairs, with an average delay of 6 days each, were estimated to control time to flowering in this cross.     

Estimation of genetic parameters for in vitro traits in wheat immature embryo cultures involving high X low regeneration capacity genotypes

Summary Estimates of gene actions were obtained for five in vitro traits of immature wheat (Triticum aestivum L.) embryo cultures from a cross of two wheat cultivars and the resulting reciprocal, F₁, F₂ and backcross populations. The contribution of additive gene effects to in vitro traits was not as important as the dominance gene effects. Epistatic gene effects were relatively more important than either additive or dominance gene effects. Of the individual types of digenic epistatic effects, the dominance x dominance estimates were relatively larger in magnitude for all in vitro culture traits measured. The maternal effect played a minor role in the inheritance of the in vitro studied traits since the difference among the reciprocal values was not significant. It is shown from the generation mean method that epistasis played a major role in the inheritance of most of the traits under study. The negative values of additive and dominance genetic variance were estimates of zero. Heritability estimates, in broad sense, were relatively high for the in vitro studied traits. In some cases, heritability estimates in broad and narrow senses are almost equal since the estimation of dominance genetic variance led to negative values. According to the results of the gene effects, dominance and epistasis were important for the shoot formation trait. Selection would be effective among the isolated genotypes on individual basis.     

Inheritance of internode length,plant form,and annual habit in a cross of cabbage and broccoli (Brassica oleracea var. capitata L. and var. italica Plenck.)

Summary When an inbred line of cabbage, Brassica oleracea L. var. capitata L., was crossed with an inbred line of broccoli B. oleraceae var. italica, the F₁ progeny were vigorous late annuals. All F₁ × broccoli backcross plants and 92% of the 3260 F₂ plants were annuals, while 40% of the F₁ × cabbage backcross plants were biennials. Annual habit is thus dominant and controlled by more than a single gene. Number of days to bud appearance in annuals varied continuously, and was primarily additive in inheritance. F₁ data suggested partial dominance for lateness but this was not supported by the F₂. Internode length was also continuous in distribution and primarily additive in inheritance, but with some dominance for short internodes in the F₁. Cabbage head forming ability was recessive and multigenic, with 2% of the F₂ plants forming heads, of which none were of commercial type and about half bolted as annuals. There was a significant chi square association between biennial habit and tendency for cabbage head formation. Clasping habit of terminal leaves was recessive to open leaves, multigenic, and associated with both cabbage heading and biennial habit.Technical Paper 4836, Oregon Agricultural Experiment Station; from an M.S. thesis by the senior author.     

Genetic analysis of Fusarium wilt resistance in cowpea (Vigna unguiculata Walp.)

This study investigated the inheritance of resistance to Fusarium oxysporum f.sp. tracheiphilum (Fot) in cowpea lines. Resistant and susceptible cowpea lines were crossed to develop F₁, F₂ and backcross populations. Reaction to Fot was evaluated in 2015 and 2016 using seed soak and modified root‐dip inoculation methods. The expression of resistance reaction in the F₁ and segregation in F₂ generations indicated the role of dominant gene controlling Fot in cowpea. These results were further supported by the result of backcross (BC₁P₁F₁ and BC₁P₂F₁) progeny tests. The backcross of F₁ with the resistant parent produced progeny that were uniformly resistant, whereas backcross of F₁ with the susceptible parent produced progeny that segregated into 1:1 ratio. The F₂ segregation ratio in the reciprocal cross showed no evidence of maternal effect in the inheritance of the resistance. Allelism test suggests that the gene for resistance in TVu 134 was the same in TVu 410 and TVu 109‐1. We also identified an SSR marker, C13‐16, that cosegregated with the gene conferring resistance to Fot in cowpea.     

Development of Yellow Seeded Brassica napus Through Interspecific Crosses

Yellow seeded Brassica napus was developed through interspecific crosses with the two mustard species, B. juncea and B. carinata. The objective of these two interspecific crosses was the introgression of genes for yellow seed colour from the A genome of B. juncea and C genome of B. carinata into the A and C genomes of B. napus, respectively. The interspecific F₁ generations were backcrossed to B. napus in an attempt to eliminate B genome chromosomes and to improve fertility. Backcross F₂ plants of the (B. napus×B. juncea) ×B. napus cross were then crossed with backcross F₂ plants of the (B. napus×B. carinata) ×B. napus cross. The objective of this intercrossing was to combine the A and C genome yellow seeded characteristics of the two backcross populations into one genotype. The F₂ generation of the backcross F₂ intercrosses was grown in the field, plants were individually harvested and visually rated for seed colour. Ninety-one yellow seeded plants were identified among the 4858 plants inspected. This result indicated that the interspecific crossing scheme was successful in developing yellow seeded B. napus.     

Single-gene resistance to zucchini yellow mosaic virus in Cucurbita moschata

Summary The mode of inheritance for resistance to zucchini yellow mosaic virus (ZYMV) in Cucurbita moschata was determined from F₁, F₂, and backcross progenies of the cross between the susceptible cultivar Waltham Butternut from the U.S.A. and a resistant inbred line of the Menina cultivar from Portugal. Resistance to ZYMV in Menina was conferred by a single dominant gene designated Zym.     

Inheritance of shelf life and other quality traits of tomato fruit estimated from F<Subscript>1</Subscript>’s,F<Subscript>2</Subscript>’s and backcross generations derived from standard cultivar, <Emphasis Type="Italic">nor</Emphasis> homozygote and wild cherry tomato

A tomato cultivar with high quality fruit and a long shelf life is a main goal in tomato breeding and it would be achieved using wild germplasm. The objective of this work was to explore the inheritance for fruit quality traits, especially fruit shelf life, in three tomato crosses using a standard Argentinean cultivar (Ca, cv ‘Caimanta’), a ripening mutant (nor, homozygous for the nor gene) of Solanum lycopersicum, and a wild cherry type (Ce, LA1385 of S. lycopersicum var. cerasiforme). The wild parent had a shorter fruit shelf life than the mutant genotype but higher than Ca. When the Ce genotype was analyzed in hybrid combination, the F₁ (Ca×Ce) was similar to the wild genotype for shelf life whereas the F₁ (nor × Ce) had a longer shelf life. Both F₁ crosses and backcrosses to the cherry type genotype had significantly lower fruit weight than the cultivated genotypes but higher than the cherry type parent. In the F₂ analysis, it was found that the inheritance underlying quality traits is complex since non allelic interactions were detected. A significant additive genetic variance was found for fruit shelf life as well as for other fruit quality traits in each cross. The genetic parameters analyzed by mean values and variances in parental, F₁ and F₂ and backcross generations indicated that the cross between the normal ripening cultivar and LA1385 of S. lycopersicum var. cerasiforme offers the best possibility to obtain long shelf life tomato genotypes with good fruit quality.     

Inheritance of Bolting in Three Sugar Beet Crosses after Different Periods of Vernalization

The induction of bolting after different periods of vernalization was studied in the parental, F₁, F₂ and backcross generations (F₂× P₁ and F₂× P₂) of three crosses between five sugar beet lines. The parental lines represented different levels of bolting resistance. The populations were evaluated in replicated field or greenhouse trials. Generation mean analysis was used to quantify the genetic effects of bolting. The additive (d) parameter was predominant for those levels of cold treatment that stimulated the bolting to occur. The dominance (h) parameter was also shown to be important in most cases. In one cross, the additive × additive (i) type of epistasis was significant. The chances of detecting genetic effects are increased by exposing the plants to vernalization conditions which maximize the difference between two parental lines of each cross. Hence, it would be sensible to exploit both additive and non-additive genetic effects in any selection programme.     

Inheritance of coffee bean sucrose content in the interspecific cross Coffea pseudozanguebariae×Coffea liberica'dewevrei'

The sucrose content of coffee bean is an important component of the coffee flavour: the higher the sucrose content in green beans, the more intense coffee cup flavour: Beans of the anciently cultivated cultivar Coffea liberica‘dewevrei’ have low levels of sucrose compared with beans of the wild species Coffea pseudozanguebariae. In the present study, the inheritance of the sucrose accumulation in the interspecific cross C. pseudozanguebariae×C. liberica‘dewevrei’ was examined. The sucrose content was measured in mature beans of both parental species, and in F₁ and backcross hybrids using high‐pressure liquid chromatography coupled to pulsed amperometric detection. The sucrose accumulation in all but one hybrid, showed genetic additivity. A segregation distortion was identified in the offspring of the backcross to C pseudozanguebariae. There was no year effect and only a low genotype × year interaction. Consequences for breeding, in relation to the coffee cup taste improvement, are discussed.     

Identification of Genes Controlling Pod Length in Spring Rapeseed, Brassica napus L., and their Utilization for Yield Improvement

Experiments were undertaken to determine the inheritance of pod length in a cross with spring rapesecd, Brassica napus, and to assess the value of pod length as a criterion of selection for high seed yield. Analyses of patterns of variation in F₂; and backcross populations derived from a cross between a short-pod line TB42 and long-pod line CA553 indicated that much of the variation in pod length could be attributed to two major genes interacting in a complementary manner. Short-pods were produced when cither one or both genes were homozygous for the recessive allele. Analyses of F₃ progenies of selected F₂ and inbred-backcross lines derived from the same cross supported the two-gene hypothesis but also indicated that the effects of the major genes on pod length were possibly modified by genes of minor effect. Field testing of families derived from random intermating between F₂, plants of the TB42 × CA553 cross showed that number of pods per plant varied independently of pod length, but seed weight per pod tended to increase with increasing pod length. As a result, families with the longest pods generally had significantly higher yields than those with short pods. It was concluded that yield improvement in B. napus could be achieved through introgression of long-pod genes into cultivars with an appropriate genetic background to ensure that selection for the long-pod character would be accompanied by an increase in seed weight per pod with little or no reduction in number of pods per plant.