Comparison of bread wheat lines selected by doubled haploid, single-seed descent and pedigree selection methods

 Yield performance of each group of ten spring bread wheat lines selected by doubled haploid (DH), single-seed descent (SSD) and pedigree selection (PS) methods from three F₁ crosses was compared with the aim of evaluating the DH method in breeding programs. Populations of 65–97 DH lines and 110 SSD lines per cross were used for selection. PS lines were developed by repeated selections from 1500 F₂ plants. Yield evaluation was performed at the F₆ generation of SSD and PS lines along with DH lines in a 2-year field experiment. It took only 2 years from the planting of wheat materials for DH production to the planting of selected DH lines for yield evaluation. There was no significant difference in grain yield between DH lines and PS lines selected from an F₁ cross whose parental varieties were closely related in their pedigrees. In two crosses with low coefficients of parentage and a large variation in their progenies, grain yield of selected DH lines was significantly lower than those of selected SSD and PS lines. These results confirm that the DH method can save time in obtaining recombinant inbred lines ready for yield evaluation. However, a larger DH population is required to achieve the same level of genetic advance with the PS method in crosses containing greater genetic variation. Received: 23 December 1997 / Accepted: 12 March 1998     

Comparison of frequency distributions of doubled haploid and single seed descent lines in barley

Summary Both doubled haploid (DH) and single seed descent (SSD) methods were used to derive homozygous lines from two crosses of barley. The frequency distributions of grain yield, heading date, and plant height of the DH and SSD lines were compared by the Mann-Whitney U test, Kolmogorov-Smirnov twosample test and Wald-Wolfowitz runs test. It was found that the DH lines distributed in the same manner as the SSD lines with respect to the three characters. The results indicated that although the SSD method had more opportunity for recombination than the DH method, it did not produce a sample of recombinants which differed significantly from the DH sample; thus both methods were equally efficient for use in deriving homozygous lines from F₁ hybrids in a relatively short time.Contribution no. 455 Charlottetown Research Station, Agriculture Canada, P.E.I. (Canada)     

The feasibility of producing inbred rather than F1 hybrid cultivars in Brussels sprouts: a preliminary genetical analysis and choice of material for inbreeding

The feasibility of producing commercially acceptable inbred lines of Brussels sprouts is investigated. Two experiments are described which examined the nature and extent of heterosis in this crop using the techniques of biometrical genetics. They indicate incomplete dominance for several production traits, lower F₁ uniformity and the presence of inbreds that outyield some F₁'s. These results strongly suggest that recombinant inbred lines, possibly outyielding the better F₁'s could be produced. Five crosses were identified for inbreeding by SSD and 1000 lines are being produced from each. The principles upon which these crosses were identified are described.     

A comparison of androgenetic doubled-haploid,and single seed descent lines in Triticale

Summary Sixty single seed descent (SSD) lines and about 25 anther-derived doubled-haploid (DH) lines were obtained from two triticale crosses. The frequency distributions of 10 quantitative agronomic traits were compared using parametric and non-parametric tests. A multivariate discriminant analysis was subsequently carried out. Gliadin patterns obtained from each line by polyacrylamide gel electrophoresis were used to calculate intra- and inter-population diversities from relative dissimilarity indices. It was found that DH and SSD lines show significant differences in frequency distributions of 1000 grain weight in both crosses, of heading date for one cross, and of lodging susceptibility for the other cross. The results of intra- and inter-population gliadin diversity indicate that although the SSD method theoretically provides more opportunity for recombination to occur than the DH method, it did not produce a greater range of recombinants. Since there is no significant difference between SSD- and DH-line distributions for grain yield, anther culture appears to be an efficient method for producing high yielding homozygous lines from F₁ hybrids of triticale in a relatively short time.     

The feasibility of producing inbred rather than F1 hybrid cultivars in Brussels sprouts: predictions from early generations

Predicting the performance of recombinant inbred lines derived by single seed descent (SSD) following crosses between inbred lines is most reliable for traits in which epistasis and genotype environment interaction are absent. The analysis of two field trials of Brussels sprouts indicated the presence of such effects in seven quantitative characters. These effects could be minimised by transforming the scale on which each character was measured. Predictions were made of the likely performance of the recombinant inbred lines currently being produced by SSD at the Institute of Horticultural Research, Wellesbourne, compared with that of current F₁ hybrid varieties. The predictions from the first trial of F₂ material were encouraging for harvest date, stem length and waste weight, but low for sprout number, quality, marketable yield and total yield. Predictions from F₃ material in the second trial were good for sprout number and total yield (in addition to harvest date, stem length and waste weight as before), but still low for marketable yield and quality, with less than 0 01% of lines expected to perform better than the F₁. Model fitting to variances suggested the presence of linkage disequilibrium in dispersion for waste weight, but no information could be obtained about linkage disequilibrium for the two most important characters, marketable yield and quality. The broad heritability of each character except stem length was lower than 0 5 (0 3 or lower for total yield, waste weight and quality), suggesting that advanced generation selection, as used in SSD, might be more successful than selection in the early generations.     

Comparative efficiency of four selection methods for deriving high-yielding lines in mungbean [Vigna radiata (L.) Wilczek]

The comparative efficiency of four selection methods, viz., honeycomb (HC), pedigree selection (PS), single-seed descent (SSD) and the bulk method (BM), was assessed in three crosses of mungbean. The lines derived by each method, along with check varieties, were yield-tested in a compact family block design in F₅ and F₆ generations during summer and kharif of 1990. On the basis of the mean of the lines, the range, the number of superior lines over the best check, and the proportion of the top 10% lines in all the crosses and generations, the honeycomb method exhibited superiority over PS, SSD and BM for yield per plant and its component traits. PS, SSD and BM did not differ from each other. The honeycomb and SSD methods were found suitable for deriving superior lines for seed yield and pods per plant in mungbean.     

Differentiation and classification of parental lines and favorable genic interactions affecting F1 fertility in distant crosses of rice (Oryza sativa L.)

 This study was intended to investigate the extent of genetic differentiation in parental lines of rice hybrids and to analyze the genetic basis underlying the fertility phenomenon in distant crosses. Two subsets of rice material (111 entries in total) were used, including 81 doubled-haploid (DH) lines and 30 Indica and Japonica rice varieties or lines (as a control). The DH lines was derived from a heterotic Indica/Japonica cross (Gui630/02428) by anther culture. The materials in the control represent a broad spectrum of the Asian cultivated rice gene pool including landraces, primitive cultivars, historically important cultivars, modern elite cultivars, super rice and parents of superior hybrids. In accordance with the NC II design, 57 out of the DH lines were test-crossed to two important wide compatibility lines: photoperiod-sensitive genetic male sterile (PGMS) line N422s and thermo-sensitive genetic male sterile (TGMS) line Peiai64s. The F₁s and their parents, 182 entries in total, were examined for the performance of seven traits in a replicated field trial. All the rice materials was surveyed for polymorphisms using 92 RFLP markers selected from two published molecular marker linkage maps. Genotypes of the F₁ hybrids at the molecular-marker loci were deduced from the parental genotypes. The analysis showed that there were two types of genetic differentiation in the two subsets of rice material; that is, qualitative differentiation in the control and quantitative differentiation in the DH lines. In addition, favorable genic interactions (both intra- or inter-locus) contributed to better increase the fertility in hybrids of distant crosses through incorporation of a wide-compatibility line as the female parent. Favorable genic interactions can be applied in hybrid rice breeding programs by selecting parents with an appropriate extent of genetic differentiation. Received: 5 June 1997 / Accepted: 10 September 1997     

Should maize doubled haploids be induced among F<Subscript>1</Subscript> or F<Subscript>2</Subscript> plants?

Maize (Zea mays L.) doubled haploid lines are typically produced from F₁ plants. Studies have suggested that the low frequency of recombinants in doubled haploids may reduce the response to selection. My objective was to determine if, for sustaining long-term response, doubled haploids should be induced in F₁ or F₂ plants during maize inbred development. In simulation experiments, I examined the response to multiple cycles of testcross selection among doubled haploid lines derived from F₁ plants (denoted by DH), doubled haploid lines derived from F₂ plants (DH_F2), and recombinant inbred (RI) lines derived by single-seed descent. For a trait controlled by 100 or more quantitative trait loci (QTL), the cumulative responses to selection were up to 4–6% larger among DH_F2 lines than among DH lines. The cumulative responses were up to 5–8% larger among RI lines than among DH lines. The QTL become unlinked as the number of QTL in a finite genome decreases, and the responses among RI, DH, and DH_F2 lines were equal or nearly equal when only 20 QTL controlled the trait. Metabolic-flux epistasis reduced the differences in the response among RI, DH, and DH_F2 lines. Overall, the results indicated that doubled haploids should be induced from F₂ plants rather than from F₁ plants. If year-round nurseries are used and new F₁ crosses for inbred development are initially created on a speculative basis, the development of doubled haploids from F₂ rather than F₁ plants should not cause a delay in inbred development.     

The agronomic performance of anther culture derived plants of barley produced via pollen embryogenesis

Anther culture was used to generate microspore-derived doubled haploid (DH) plants from four spring barley crosses. The culture medium used contained maltose as the sole carbohydrate source and the mode of plantlet regeneration was mainly via pollen embryogenesis. Both haploid and spontaneously doubled regenerants were produced and the doubled haploids were compared to recom-binant inbred lines generated by several rounds of selfing (single seed descent). Parental, DH and single seed descent (SSD) lines were grown in randomised, replicated field trials and the samples were scored for a range of agronomic traits. The mean performance and phenotypic distribution of the DH and SSD samples were similar and there was little evidence to support the conclusion that anther culture derived lines exhibit a reduction in vigour. Where significant differences were detected between groups these were mainly confined to crosses which were segregating for the denso dwarfing gene. The differential transmission of particular regions of the barley genome may therefore influence and confound the expression of agronomic traits in DH populations. This is the first report of the agronomic performance of anther culture lines produced via pollen embryogenesis and the results are discussed in relation to the exploitation of anther culture technology in barley breeding.     

A simple method to estimate the percentage of hybridity in canola (Brassica napus) F1 hybrids

We have developed an efficient PCR-based system that uses RAPD markers for the certification of F₁ hybrids of canola. These markers were selected by screening five parental lines used in three crosses X, Y and Z with 131, 131 and 322 primers respectively. Stable DNA fragments that were homozygous and specific to the male inbreds were used to certify F₁ hybrid populations. The hybrid production system was based on self-incompatibility (SI) alleles that prevent self-pollination of the female parent. The efficiency of two S-alleles was compared under both field and greenhouse conditions. The percentage of hybridity was estimated in different F₁ populations. We found a significant difference between the two alleles for their efficiency in controlling selfing; both alleles were stable under greenhouse conditions, one allele appeared less reliable under field conditions.     

Prediction of progeny variation in oat from parental genetic relationships

Summary The ability to predict agronomic performance of progeny from a cross would be a great benefit to plant breeders in selecting parents. The predictive value of parental genetic relationships estimating F₁ progeny means and F₄ family variances of nine argronomic traits was tested in 76 oat crosses, using genetic distance measures based on coefficients-of-parentage, quantitatively inherited morphological characters, and discretely inherited biochemical and morphological characters. Coefficients-of-parentage were better predictors of F₁ performance than similarity measures derived from plant morphology or discretely inherited characters. Combined distance measures were better estimators of F₁ specific combining ability (SCA) effects than any single measure. Among cultivars of similar adaptation and quantitative morphology, crosses between parents with high coefficients-of-parentage gave higher SCA effect values than crosses of distantly related parents for grain yield and total biomass. The opposite was found for crosses among cultivars of different adaptation or quantitative morphology. The best predictor of trait variances among F₄ families was coefficients-of-parentage. Crosses between more distantly related parents produced larger variances among families than crosses between closely related parents for plant biomass. For grain yield, test weight, heading date, grain filling period, and maturity date, crosses between more closely related parents produced larger among-family variances than crosses of distantly related parents. Crosses between more distantly related parents involved at least one parent unadapted to central New York, and resulted in most of the progeny being generally unadapted. This, in part, may account for the low genetic variances for heading date, test weight, and grain yield in crosses of distantly related parents.     

Prediction of testcross means and variances among F3 progenies of F1 crosses from testcross means and genetic distances of their parents in maize

 Prediction of the means and genetic variances in segregating generations could help to assess the breeding potential of base populations. In this study, we investigated whether the testcross (TC) means and variances of F₃ progenies from F₁ crosses in European maize can be predicted from the TC means of their parents and F₁ crosses and four measures of parental genetic divergence: genetic distance (GD) determined by 194 RFLP or 691 AFLP^{TM 1} markers, mid-parent heterosis (MPH), and absolute difference between the TC means of parents (∣P₁−P₂∣). The experimental materials comprised six sets of crosses; each set consisted of four elite inbreds from the flint or dent germplasm and the six possible F₁ crosses between them, which were evaluated for mid-parent heterosis. Testcross progenies of these materials and 20 random F₃ plants per F₁ cross were produced with a single-cross tester from the opposite heterotic group and evaluated in two environments. The characters studied were plant height, dry matter content and grain yield. The genetic distance between parent lines ranged between 0.17 and 0.70 for RFLPs and between 0.14 and 0.57 for AFLPs in the six sets. Testcross-means of parents, F₁ crosses, and F₃ populations averaged across the six crosses in a particular set generally agreed well for all three traits. Bartlett’s test revealed heterogeneous TC variances among the six crosses in all sets for plant height, in four sets for grain yield and in five sets for dry matter content. Correlations among the TC means of the parents, F₁ crosses, and F₃ populations were highly significant and positive for all traits. Estimates of the TC variance among F₃ progenies for the 36 crosses showed only low correlations with the four measures of parental genetic divergence for all traits. The results demonstrated that for our material, the TC means of the parents or the parental F₁ cross can be used as predictors for the TC means of F₃ populations. However, the prediction of the TC variance remains an unsolved problem. Received: 4 August 1997 / Accepted: 17 November 1997     

The agronomic performance of wheat doubled haploid lines derived from wheat x maize crosses

Summary The agronomic performance of 9 doubled haploid (DH) lines of Chinese Spring, 6 DH lines of Hope, 14 DH lines of the single chromosome substitution line Chinese Spring (Hope 5 A) and their respective parents was analyzed under field conditions. Seventeen Chinese Spring DH lines derived from wheat x Hordeum bulbosum crosses were also included for comparison. No significant variation was detected in either population of Chinese Spring DH lines and neither DH population differed from its parent. The Hope DH lines differed significantly for tiller biomass, spikelet number per ear, ear grain weight and 50-grain weight. However, all the variation could be attributed to the poor performance of only one line. Chinese Spring (Hope 5 A) DH lines showed significant variation for ear emergence time, but this was probably due to genetic heterogeneity in the parental stock. Overall, the results suggest that most DH lines produced by the wheat x maize method resemble their wheat parent, and that the variation induced in DH production is likely to be similar to that found in DHs from wheat x Hordeum bulbosum crosses.     

Genetic variability and advance under four selection procedures in wheat pedigree breeding programme

Summary Four methods of generation advance (SPS, SSD, BP and MMS) were compared in F₃ and F₄ generations. In the F₃ generation, the SPS and SSD methods of generation advance proved superior to the BP and MMS methods for grain yield per plant and for at least one of the yield component traits. The F₃ SSD population did not differ significantly from the F₃ SPS for any of the traits. However, the F₃ SSD population retained more range and cv for different traits than with other methods of generation advance. F₄ progenies derived from F₃ SSD population were significantly superior for grain yield than lines derived from the other three F₃ populations. The MMS method of generation advance proved useful for increasing the 1,000-grain weight for which initial selection was made.Part of Ph.D. Thesis submitted by senior author to Haryana Agricultural University, Hisar     

North American study on essential derivation in maize: inbreds developed without and with selection from F2 populations

An essentially derived variety largely retains the characteristics of a parental or ancestral variety. A consensus has not been reached regarding the threshold for declaring essential derivation in maize (Zea mays L.), partly because benchmark data are lacking. Our objective in this study, commissioned by the American Seed Trade Association, was to determine the range of parental contribution among maize inbreds developed without and with selection. Seed companies in North America contributed existing proprietary data on the molecular marker similarity of 100 or more families, developed without selection from F₂ populations, with each of their parents. The companies also sent us seed samples of elite inbreds, developed with selection from F₂ populations, for analysis using 60 RFLP marker loci and 20 SSR marker loci. Among the families developed without selection, the average parental contribution was close to the expected value of 0.50 for F₂ populations. Specific families received up to 79% of their alleles from one parent. Although selection tended to increase the frequency of such transgressive segregants, it did not necessarily increase the maximum parental contribution in an F₂ population. Parental contributions were consistent between the elite inbreds and their early-generation families. We conclude that inbreds with 70% to nearly 80% of their genome derived from one parent can be obtained from an F₂ population. Further empirical data would be valuable particularly for backcross populations, which were not available in this study. Received: 21 June 2000 / Accepted: 28 July 2000     

RFLP markers and predicted testcross performance of maize sister inbreds

 Inbreds selfed from the same F₂ or backcross population are referred to as sister inbreds. In some situations, maize (Zea mays L.) sister inbreds may not have testcross data available for best linear unbiased prediction (BLUP) of single-cross performance. This study evaluated the usefulness of BLUP and restriction fragment length polymorphism (RFLP)-based coefficients of coancestry ( f ) in predicting the testcross performance of sister inbreds. Parental contributions (p) were estimated from 70 RFLP loci for 15 inbreds that comprised three sister inbreds selfed from each of five F₂ populations. Estimates of p were subsequently used to calculate RFLP-based f. Grain yield, moisture, and stalk lodging data were obtained for 2265 single crosses tested by Limagrain Genetics in multilocation trials from 1990 to 1995. Performance of the sister inbreds when crossed to several inbred testers was predicted from the performance of the tested single crosses and RFLP-based f. Correlations between predicted and observed performance, obtained with a delete-one cross-validation procedure, were erratic and mostly low for all three traits. Correspondence was poor between ranks for predicted and observed general combining ability of the sister inbreds. The results suggested that the proportion of the genome derived by a sister inbred from a given parental inbred does not solely determine its testcross performance. The failure of BLUP and RFLP-based f to consistently predict testcross performance indicated that actual field testing will continue to be necessary for preliminary evaluation of sister inbreds. Received : 17 March 1997 / Accepted : 18 April 1997     

The genetical basis of hybrid vigour in a highly heterotic cross of Nicotiana tabacum

The genetical control of F₁ heterosis, observed in a cross of desirable Nicotiana tabacum varieties, was investigated by analysing the data of the basic generations, triple test cross-families and random samples of doubled haploids (DH) and single-seed descent (SSD) lines. Analyses of the first-degree statistics revealed a complex control underlying the genetic variation, including the presence of epistasis, linkage, maternal effects and their interactions, in addition to the additive and dominance effects of the genes segregating in the cross. These analyses identified gene dispersion, directional dominance, and duplicate epistasis, as the main causes of heterosis. The triple test-cross analysis also confirmed the presence of non-allelic interactions and indicated that the dominance ratio, although inflated by epistasis, is consistently partial for all the traits. The extent of transgression in the recombinant inbred lines finally established unequivocally that, as in numerous other crosses, gene dispersion and unidirectional, but partial, dominance are the true causes of heterosis in this cross too.