Comparison of two crossing and four selection schemes for yield, yield traits, and slow rusting resistance to leaf rust in wheat

The most important breeding objectives in crop improvement are improving grain yield, grain quality, and resistances to various biotic and abiotic stresses. The objectives of our study were to compare two crossing and four selection schemes for grain yield, yield traits, and slow rusting resistance to leaf rust (Puccinia recondita) based on additive genes in wheat (Triticum aestivum), and to identify the most efficient crossing and selection methodologies in terms of genetic gains and cost efficiency. Segregating populations were derived from 18 simple crosses and the same number of top (three-way) crosses. Half of the crosses were derived from Yecora 70 and the other half from Veery #10 as the common leaf rust susceptible parents. The four selection schemes were: pedigree, modified bulk (F₂ and F₁-top as pedigree, selected lines in F₃, F₄, F₂-top, F₃-top as bulk; and pedigree in F₅ and F₄-top populations), selected bulk (selected plants in F₂, F₃, F₄, F₁-top, F₂-top and F₃-top as bulk; and pedigree in F₅ and F₄-top populations), and nonselected bulk (bulk in F₂, F₃, F₄, F₁-top, F₂-top and F₃-top; and pedigree in F₅ and F₄-top populations). A total of 320 progeny lines, parents and checks were tested for grain yield, other agronomic traits and leaf rust resistance during the 1992/93 and 1993/94 seasons in Ciudad Obregon (Sonora State, Mexico) which represents a typical high yielding irrigated site. The influence of the type of cross and the selection scheme on the mean grain yield and other traits of the progenies was minimal. The selection of parents was the most important feature in imparting yield potential and other favourable agronomic traits. Moreover, the highest yielding lines were distributed equally. Progeny lines derived from Veery #10 crosses had significantly higher mean grain yield compared to those derived from the Yecora 70 crosses. Furthermore, a large proportion of the highest yielding lines also originated from Veery #10 crosses. Mean leaf rust severity of the top cross progenies was lower than that of the simple cross progenies possibly because two parents contributed resistance to top cross progenies. Mean leaf rust severity of the nonselected bulk derivatives was twice that of lines derived from the other three schemes. Selected bulk appears to be the most attractive selection scheme in terms of genetic gains and cost efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

Studies in the breeding of self pollinating cereals

The trials described in this paper have led to the identification fo a number of promising parental combinations suitable for detailed pedigree sclection. Although these include crosses involving the highest yielding parents which would probably have been studied without this evidence, some of the crosses identified would not have been considered had these experiments not been conducted. This information has been obtained from a series of trials conducted in F₂ and F₃.     

Breeding for yield and quality in durum wheat

Three populations of an intervarietal durum wheat cross IWP5308/PDW208, F₅, F₅BIP₁ (population derived after intermating in F₂) and F₅BIP₂ (population derived after intermating in BIPF₁), were evaluated under three different agronomic environments for mean performance and stability of genotypes for grain yield, yield components and protein content. Though the biparental progenies indicated a higher mean performance, they did not differ significantly from progenies of the pedigree method for almost all characters. The biparental progenies, however, produced a higher number of stable genotypes for grain yield per plant, grains per ear and protein content. The F₅ population had a higher number of stable genotypes for 1000 grain weight and number of tillers per plant. The BIP progenies also had a higher number of genotypes with above average mean performance, and many were significantly higher than the checks WH896 and WH542, compared with F₅ progenies. Hence, in spite of high G x E interactions, the use of cycles of biparental mating and selection of top yielding lines on the basis of yield components can enable selection of stable genotypes with high protein content. Number of tillers per plant and 1000 grain weight were the yield component characters which made maximum contribution to phenotypic stability of the genotypes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Efficiency of Early Selection for Yield Performance in Wheat

Selection for yield in early generations is difficult, mainly due to a shortage of seed. In the breeding company Fr. Strube the breeder uses three-way crosses and first selects for yield between F₂′-derived lines in the F₄′. The generations are called F₁, F₂′, etc., because the first generation is already segregating. The efficiency of this selection was checked in the F₅′ and F₆′. For this purpose F₃′-derived progenies, corresponding to poor and good yielding bulks in F₄′, were selected. In later generations the average yield of the group of strains with good yield in the F₄′ was 4 % (F₅′) and 3 % (F₆′) higher, respectively, than the mean of the strains with low yield in the F₄′. Strains with very low yielding potential can easily be discarded in the F₄′. The risk of discarding lines with a very high yield is minimal. Selection for yield in the F₄′ was as effective as in the F₅′. The importance of testing F₂′-derived bulks in the F₄′ is not only based on the assessment of yield but also on the possibility of reliable intensive selection through recording visual characters (height, resistance against diseases) in a second environment in parallel to the single plant progenies.     

Selecting superior spring wheat crosses in early generations

Summary Three methods were evaluated as potential aids in identifying superior crosses early in the breeding program: (1) F₁ yield, (2) F₂ yield, and (3) midparental yield. Two sets of spring wheat (Triticum aestivum L. em. Thell.) crosses and parents were evaluated in replicated single row plots in F₁ and in replicated multi-row plots in F₂ over a period of five years. On the basis of F₁ yields, two of the highest yielding and two of the lowest yielding crosses in each set were chosen for a more detailed selection study in subsequent generations.In Set 1, random F₂ head selections for each of the selected four crosses were evaluated in 3-row yield plots in F₄. In Set 2, F₂ plant selections were made on the basis of head weight and the highest yielding 10% evaluated for yield in F₄.Lines of crosses identified as high yielding in F₁ had significantly greater mean yields in F₄ than those of crosses that were low yielding in F₁. The high yielding crosses had three to four times as many lines yielding in the top 10% in F₄ than did the low yielding crosses.Correlations for yield were obtained between F₁ and F₂ in different years, F₂ and midparents in the same year, lower or no correlations between F₁ and midparents in the same year, and no correlations between F₁ and midparent and F₂ and midparent in different years. It is suggested that larger plot sizes rather than single rows be used to evaluate parents.Midparental yield, F₁ yield and F₂ yield tests are recommended as a progressive set of screening tests for a given set of crosses to effectively maintain the superior crosses in the breeding program.     

Honeycomb selection for yield in early generations of spring wheat

Summary The effectiveness of the honeycomb selection method for yield in spring wheat (Triticum aestivum L.) was evaluated using progenies from two wheat crosses, Glenlea x NB131 and Glenlea x Era. Honeycomb selection was carried out in the F₂ and F₃ generations, grown at the University of Manitoba in the summers of 1980 and 1981, respectively. In both generations, divergent selection was made for both high and low yield. Plants selected in the F₃ generation were entered in an F₄ yield test in the summer of 1982. Results of the experiment showed that honeycomb selection for yield in the F₂ and F₃ generations was effective in identifying parents of high- and low-yielding lines. F₃ plants from highyielding F₂ selections gave higher yields than those from low-yielding F₂ selections by 11.5% and 13.0% for Glenlea x NB131 and Glenlea x Era crosses, respectively. The F₄ yield test showed that high yielding selections from both crosses significantly outyielded by 8.9% low yielding selections and by 14.4% the unselected composite lines. It is concluded that the honeycomb selection method can be used for early generation selection in spring wheat.     

Tartary buckwheat breeding (<Emphasis Type="Italic">Fagopyrum tataricum</Emphasis> L. Gaertn.) through hybridization with its Rice-Tartary type

Tartary buckwheat (Fagopyrum tataricum) is a highly nutritious crop that is widely grown in Asia, but the flour contains a large portion of the hull if it is ground with industrial processing since the hull is very hard to remove as it adheres to the testa layer of the groat. Rice-Tartary, a particular type of Tartary buckwheat, has seeds with a loose hull and the presence of splits on the sides of the seeds that make dehulling easily. The present study reports on the first attempt of crop improvement in Tartary buckwheat breeding through hybridization. Hybrids were obtained by hybridization of crosses between Tartary buckwheat and Rice-Tartary. Additional crosses were made among selected progenies of these crosses. Based on progeny analysis of the F₁, F₂, and F₃, the character of Rice-Tartary, as exhibited as the presence of splits on the sides of the seeds, is controlled by one gene which is homozygous recessive. A Tartary buckwheat breeding program has been conducted for 6 years based on these crosses. Advanced lines with easy dehull and yield potential are at the stage of ready for yield trials. Production of easy dehulling Tartary buckwheat in the future could boost Canada’s domestic and international markets.     

Effectiveness of early-generation testing applied to upland rice breeding

A plant breeding program is a long-term investment. Therefore, periodic assessment of the effectiveness of a breeding strategy is essential to maximize genetic gains per unit of time and resource invested. In this work, we assessed the effectiveness of the early-generation testing (EGT) approach used in the upland rice (Oryza sativa L.) breeding program at Embrapa (Brazilian Agricultural Research Corporation), Brazil, estimating the genetic progress achieved for three traits in two distinct phases, spanning 15 years. In the first phase (from 2003 to 2010), it was used the bulk method within F₃ progenies with prior testing of F₂ crosses, while in the second phase (from 2010 to 2017), it was used the bulk method within F₂ progenies. The dataset comprised 70 yield trials, involving 1884 F_3:5 progenies (phase I) and 925 F_2:4 progenies (phase II) from an elite population, and 10 check cultivars, evaluated for grain yield (GY), plant height (PH) and days to flowering (DTF). For estimating the genetic gain, we adapted a generalized linear regression method to compute bi-segmented linear regression coefficients. Desirable genetic gains were achieved only for GY in both phases of the breeding program, with 78.75 kg ha^?1 year^?1 (2.68%) in the first phase, and 53.78 kg ha^?1 year^?1 (1.54%) in the second phase. These results show the effectiveness of EGT, especially via bulk method within F₃ progenies with prior testing of F₂ crosses, applied to upland rice breeding. Some refinements are discussed in the method to make it more cost-effective and more efficient in achieving genetic gains.     

Inheritance of parthenocarpy in pickling cucumbers (Cucumis sativus L.) and linkage with other characters

Summary The inheritance of parthenocarpy in the pickling cucumber has been studied with progenies of an incomplete diallel cross between lines with different degrees of parthenocarpy and with two sets of F₁, F₂ and B₁ of crosses between parthenocarpic and non-parthenocarpic lines. The inheritance of parthenocarpy could be explained by three independent, isomeric major genes with additive action, together with a non-allelic interaction of the homozygote-heterozygote type. Indications have been found for linkages between genes that govern parthenocarpy, femaleness and the spined/hairy fruit character. Consequences for breeding parthenocarpic varieties are discussed.     

Genetics of Resistance in Vicia sativa L. to Orobanche crenata Forsk.

The genetics of resistance of common vetch (Vicia sativa L.) to broomrape (Orobanche crenata Forsk.) was studied for two years by using the P₁, P₂, F₁, BC¹, BC₂, F₂ F₃, and F₄ generations obtained from crosses between resistant and susceptible lines. Resistant lines were selected by screening a world collection m a naturally infested plot. Resistance was tested both under field and greenhouse conditions. The best index to measure resistance was the number of emerged broomrapes per host plant. The results fit the additive-dominance model. The main component of the variation was additivity; dominance and interaction effects seemed to depend on the environment. When dominance is expressed, a low number is dominant over a high number of broomrapes per host plant.     

Evaluation of Pedigree and Single Seed Descent Selection Methods for Cultivar Development in Cowpea (Vigna unguiculata L. Walp)

Three crosses of cowpea (Vigna unguiculata [L.] Walp) were used to assess the effectiveness of two selection procedures in cowpea. The selection procedures were pedigree selection (PD) and single seed descent selection (SSD). Lines developed through each procedure were yield tested and selected at F₆ and F₇ for PD and SSD, respectively. The selected lines for both procedures were compared at F₈ for grain yield per plant (YLD), number of pods per plant (NPO), and pod development period (PDP), in two locations. Lines developed through both procedures differed more in their population means than in the performance of their top 10% or highest yielding lines. There was no consistent relationship between magnitude of genetic variability and percentage of superior lines obtained from each procedure. Both procedures have been effective in producing superior genotypes for yield and number of pods. The PD method produced superior transgressive segregants in two out of the three crosses for PDP. SSD allowed a more rapid generation than PD.     

Studies on the breeding of self-pollinating cereals

Summary The limitations of the pedigree and mass methods of selection in the improvement of self pollinating cereal crops are discussed and the value of quantitative data in the early generations of selection is emphasised. Two systems of selection which provide such data have been introduced at the Cambridge Plant Breeding Institute.In one, known as the F₂ progeny method of selection, the progenies of selected F₂ plants are grown in yield trials without further selection during F₄, F₅ and F₆ after which single plant selections are made within the more promising progenies.In the other system, known as the pedigree trial method of selection, normal pedigree selection is carried out in F₂ and F₃. In the F₄ generation single plants are selected from the better families for continued pedigree selection and within each of these families the remaining plants are bulked to give grain for a yield trial in the following year. This process is repeated in F₅ and F₆, the grain for trial being in each case obtained from the progeny rows and not from the preceding trial.The use of these methods is illustrated by reference to wheat hybrids grown at Cambridge, and the data obtained is shown to be useful in comparing the value of different parental combinations as well as in assessing the relative merits of the selections within each cross.     

Inheritance of tomato leaf curl virus resistance in Lycopersicon hirsutum f. glabratum

Summary Inheritance of resistance to tomato leaf curl virus (TLCV) was studied in the progenies derived from interspecific crosses between TLCV resistant Lycopersicon hirsutum f. glabratum line B 6013 and five susceptible cultivars (HS 101, HS 102, HS 110, Pusa Ruby and Punjab Chhuhara) of L. esculentum. P₁, P₂, F₁, F₂, B₁ and B₂ progenies of the five crosses were artificially inoculated with local strains of TLCV by means of the vector whitefly, Bemisia tabaci (Genn.). and the disease reaction was studied in all the crosses. Reaction of parents, F₁, F₂ and backcrosses suggests that resistance derived from L. hirsutum f. glabratum B 6013 is based on two epistatic genes, one from the wild parent and one from the cultivated one, resulting in a 13:3 segragation in the F₂.     

Improvement of large-seeded common bean cultivars under sustainable cropping systems in Spain

Approaches are needed to broaden the genetic base and improve earliness and yield potential of large-seeded beans under sustainable cropping systems. The objective of this research was to develop adapted dwarf bean populations having a commercial seed quality and yield suitable for the production in the South of Europe. The original base populations were produced from crosses between genotypes within each Mesoamerica, Nueva Granada and Peru races, and between Peru and Nueva Granada, and Mesoamerica and Nueva Granada races. Visual mass selection for plant performance was practised in the F₂ and F₃ generations. In the F₄ and F₅, single plants were harvested under two cropping systems (sole cropping and intercropping with maize). From F₄, selection was based on precocity, combined with seed yield and seed commercial type. The F_4:7 selected lines from each original population were compared with their parents and five checks at four environments and two cropping systems. Differences among environments, populations, parents and checks were observed for all traits. Under intercropping with maize, there was a 50% reduction in seed yield. Yield of parents and checks belonging to Andean South American races, intraracial (Nueva Granada × Nueva Granada) and interracial (Nueva Granada × Peru) populations, was higher than that of those of Middle American origin. Intraracial crosses within large-seeded Andean South American (Peru race) and Middle American gene pools (Mesoamerica race) did not produce lines yielding more than the highest yielding parent. Only two large-seeded lines selected from crosses between small- and large-seeded gene pools out-yielded the best parent and check cultivar.     

The inheritance of resistance to Empoasca kraemeri Ross & Moore in the common bean,Phaseolus vulgaris L.

Summary Genotypes of Phaseolus vulgaris differ in their level of resistance to Empoasca kraemeri, but the speed with which these differences can be accumulated into a resistant variety will depend on whether they can be detected in single F₂ plants, and on whether the genetic control of resistance is additive. Thirty-eight homozygous genotypes were therefore grown in small plots with and without insecticidal protection, and also in simulated F₂ populations containing one plant of each genotype. In addition, the F₂ and F₃ generations of crosses were studied. Both generations of a 13 parent diallel were grown with and without insecticidal protection. Six crosses, involving some of the parents in the diallel and two additional genotypes, were studied in more detail. Protected and unprotected F₂ plants from these crosses were harvested individually, and the resulting F₃ families were grown with and without protection. In all the expriments, the level of damage to the unprotected plants and the vigour of the protected plants was assessed visually by means of a numerical score, and the seed yield was measured.The performance of a single homozygous plant was a reasonable predictor of the performance of the same genotype in a small plot, but in the six crosses studies in detail, the correlations between F₂ and F₃ performance were low. In the analysis of the diallel cross, the significant differences among crosses were due to specific combining ability and not general combining ability. These results indicate that the genetic control of resistance is largely non-additive, and that F₂ selection will probably be ineffective. However, F₃ selection should be effective, and transgressive segregants for resistance should be obtainable.In some of the six crosses, the F₃ families differed both in general vigour and in narrow-sense resistance to E. kraemeri, as defined by Galwey & Evans (1982). The visual assessment of damage symptoms in each plot was related to the subsequent seed yield of the plot, but was more heritable than yield. The relationship between these two variables was investigated by the genetic correlation coefficient and by an alternative method, and was found to be partly genetically determined, but partly due to the influence of environmental factors on both variables. This indicates that screening for E. kraemeri resistance should be done by means of a visual damage score in a carefully chosen, uniform environment.     

Studies on the breeding of self-pollinating cereals

The behaviour of 19 varieties of spring wheat under different environmental conditions is discussed in terms of the following five characters: weight per grain, grains per spikelet, spikelets per ear, ears per plant and yield of grain per plant. The first four of these characters (the components of yield) are shown by correlation analyses to be largely independent of each other.Four varieties, chosen because they each showed extreme values of one or more of the components of yield, were crossed in all possible combinations. The F₁ and F₂ of this diallel set of crosses was analysed for each character according to the method of Jinks and Hayman. The results are discussed in relation to crop improvement and predictions are made for each character as to which cross or crosses are likely to be the most profitable for the breeder. The future handling of this material in order to test these predictions is outlined.     

Induced polygenic variation and genetic architecture in Indian mustard

Summary The F₁ seed obtained from an eight-parent diallel (excluding reciprocals) in Indian mustard was treated with 100 Kr of gamma rays, grown and selfed to obtain F₂M₂. The mean was decreased in the majority of the crosses in an irradiated set whereas the coefficient of variation increased. Combining ability analysis revealed the change in relative ranking of the parents and crosses and suggested the genotype-specific responses to irradiations. Components of combining ability and heterosis were also changed considerably. In some cases irradiation resulted into an increase in additive genetic variance.     

A proposed mechanism for loss of heterozygosity in rice hybrids

A phenomenon, loss of heterozygosity (LOH), was discovered in hybrid plants involving a selected plant (named AMR) of the Chinese rice cultivar ‘ZhongxinNo. 1’ as one parent. In these hybrids and some of their progenies, somatic variations were manifested by molecular genotypes and/or morphological phenotypes in vegetative parts of the same plant. Random amplified polymorphic DNA (RAPD) markers for the parents have been followed through the F₃ generation. RAPD markers were uniformly present or absent in all plants within some or all F₂ panicle rows derived from F₁ hybrids involving AMR. In contrast, RAPD markers segregated in the Mendelian manner for dominant markers in panicle rows derived from control hybrids. Certain F₂ panicle rows from F₁hybrids involving the special rice became fixed for all assayed RAPD markers. Genotype fixation was confirmed by molecular assays and field observations of the F₃ progenies. We propose a new biological mechanism, called ‘assortment mitosis,’ as being responsible for the observed phenomenon. The use of this mechanism in plant hybrids allows the development of uniform progenies as early as the F₂ generation. Therefore, the time required to obtain fixed non-parental type progenies for subsequent performance trials can be drastically shortened. Utilizing this mechanism in plant breeding represents a new approach and requires the modification of conventional plant breeding procedures. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic analysis of resistance to <Emphasis Type="Italic">Fusarium</Emphasis> root rot in common bean

Fusarium root rot (FRR) is a major disease of common bean worldwide. Knowledge of the inheritance of resistance to FRR would be important in devising strategies to breed resistant varieties. Therefore, a 12 × 12 full diallel mating scheme with reciprocal crosses was performed to generate 132 F₁ progenies, which were then advanced to the F₃. The progenies were evaluated for resistance to FRR under green house conditions in Uganda. General combining ability (GCA) effects were highly significant (P ≤ 0.01) for disease scores. Specific combining ability effects were not significant (P > 0.05) in the F₁, but were highly significant (P < 0.01) in the F₃ generation. These results indicate that resistance to FRR was governed by genes with additive effects in combination with genes with non-additive effects. Reciprocal differences were also significant (P = 0.01) at F₁ and F₃, primarily reflecting a large influence of maternal effects in both these generations. In fact, susceptible parents did not differ significantly (P > 0.05) for disease scores when used as paternal parents in the F₃, but differed strongly as maternal parents (P = 0.0002). Generally, the progenies were distinctly more resistant when the resistant parent was used as the female in crosses, especially as observed in the F₃. The maternal effects were strong in the F₃ generation, suggesting a complex form of cytoplasmic–genetic interaction. The non-maternal reciprocal effects in the F₃ were significant (P < 0.05) in both the resistant × resistant diallel, and in the resistant × susceptible crosses. Mid-parent heterosis (MPH) occurred in most crosses, with average heterosis approximately equal in each of the three generations, indicating that epistasis was probably more influential than dominance of individual genes. Gene-number formulas indicated that several genes were involved in resistant × susceptible crosses. Among resistant × resistant crosses, many produced continuous distributions of F₁ progeny scores, suggesting polygenic inheritance, while bi-modal distributions were characteristic of the F₃ distributions, and fit expected ratios for two or three loci segregating in each cross. Dominant forms of epistasis favoring resistance were strongly indicated. Parent–offspring heritability estimates were moderate. Overall, the results indicate that resistant parents contain a number of different resistance genes that can be combined with the expectation of producing strong and durable resistance. The lines MLB-49-89A, MLB-48-89, RWR719 and Vuninkingi, with large and negative GCA effects, contributed high levels of resistance in crosses and would be recommended for use in breeding programs.     

The inheritance of curding periods in early summer and autumn cauliflower

Two series of diallel crosses were made between (a) eight varieties of autumn cauliflowers and (b) six inbred lines from a single variety of early summer cauliflowers in order to study the inheritance of curding periods. F₁'s and parents from each diallel were grown in separate randomised block experiments and the analyses presented as conventional Wr Vr graphs.The array points from the diallel analysis of autumn cauliflowers were spread along a regression line of unit slope thus indicating that the gene system controlling curding periods was additive with no evidence of gene interaction in these crosses. Varieties of early/mid period maturity with the exception of Veitch's Autumn Giant possessed most dominant polygenes while late maturity was controlled by recessive polygenes. The diallel analysis of curding periods for the early summer cauliflower inbreds indicated the presence of some interaction which was due to the influence of two lines. On reanalysis without these lines the regression of Wr Vr agreed with a slope of 1, although there was a higher degree of dominance than was shown by the autumn cauliflowers. Analysis of the early summer cauliflower lines for numbers of leaves produced before curding indicated an association between low leaf numbers and early curding, and between high leaf numbers and late curding.None of the F₁ hybrids in either series of crosses curded earlier than the earlier curding parent.