Estimation of genetic parameters in barley (Hordeum vulgare L.)

Summary Four exotic and four indigenous strains of barley were used for making diallel crosses. The sets of parents and crosses making full, half and quarter diallel were analysed in a randomized block design for plant height, number of effective tillers, ear length, grain yield per plant, 100 grain weight and number of grains per ear.The three alternatives of diallel were similar with respect to the estimates of degree of dominance, general combining ability and specific combining ability, indicating that all these three methods of diallel were equally efficient. However, as the number of entries are minimum in quarter diallel, it would be economical in terms of cost, time and labour to estimate genetic parameters by this method. Average degree of dominance was found in the range of overdominance. The ranking of parents on the basis of their array mean was similar to the ranking based on gca effects. Similarly, the ranking of crosses on the basis of per se performance was similar to the ranking based on sca effects. This suggests that the selection of best general combiner or best cross combinations may be easier and more effective through array mean for per se performance rather than through high gca and sea effects, respectively. From among 56 crosses, IB-226 X X C-164 was the one which showed superiority for maximum number of characters followed by AB-12/59 X PTS-57. High sea effect for plant height, ear length, grain yield, 100 grain weight and number of grains per ear was the result of cross between parents having high X low general combining ability, indicating additive X dominance type of gene interaction. For number of effective tillers, high sca was produced by low x low general combiners, indicating dominance x dominance gene interaction.Part of a Ph. D. thesis submitted by senior author to Haryana Agricultural University, Hissar.     

A comparison of four experimental designs for the estimation of heritability

Summary The partial diallel cross, the complete diallel cross, and the designs known as North Carolina Experiments 1 and 2 are compared for their usefulness in estimating heritability. It is first shown that reliable values for the sampling mean and variance of heritability estimates are obtained from approximate expressions based on the moments of the chi-square distribution. These expressions are then applied to determine the optimum experimental designs for a range of situations.The main basis for discrimination is the amount of information per unit, defined as i = 1/(N var( ²)), where ² is the estimate of the heritability h ² and N is the number of units in the experiment, either individuals or families.The two parameters considered were the heritability of individuals and the heritability of full-sib families, and for each of these the partial diallel cross was the most preferred, followed in decreasing order of preference by design NC2, the complete diallel, and design NC1.It is first shown that there is no optimum number of parents for a partial diallel cross or male parents for designs NC1 and NC2. The number of crosses per parent for a partial diallel or dams per sire for designs NC2 and NC2 should generally be six or less. Any expansion should be in the direction of using more parents in the case of the partial diallel, or more male parents in the case of designs NC1 and NC2. For the two heritability parameters considered in this study it is inefficient to increase the number of replicates beyond two.     

Comparative genetic analyses of metric traits using diallel and factorial mating designs in bread wheat

Summary For studying the inheritance of metric traits, diallel cross and factorial mating designs are commonly used. Since factorial mating design is less restrictive in crossing plans, the genetic information drawn from it was compared with that from a diallel cross. The comparison was made using graphical, genetic components and combining ability analyses for grain yield, grain weight and spike length in a field experiment of bread wheat (Triticum aestivum L.). Analyses were made on a nine parent diallel cross and a 4 × 5 factorial mating design which was sampled from the diallel cross. In general, there was a high degree of agreement between the results obtained from factorial mating design and diallel cross analyses showing thereby that the former provides almost equivalent genetic information to the latter.     

Evaluation of circulant partial diallel crosses in maize

Summary The present study was conducted in maize (Zea mays L.) on crosses among 20 diverse parents. The materials were evaluated in four different environments for eight characters. Combining ability analysis was carried out following diallel and partial diallel crosses. The number of crosses per parent (s) varied from 3 to 19 and the results were studied to identify the critical value of s that would provide an adequate information with minimum resources. The S₅ partial diallel was as good as the S₁₉ for the detection of differences among general combining ability (GCA) effects. Even the S₃ gave adequate information in the case of characters with high heritability. However, partial diallel analysis was less efficient in detecting the differences due to specific combining ability (SCA) effects. These results varied with environments, and characters with low heritability were more prone to misinterpretation. GCA effects showed fluctuations in partial diallel analysis which were more pronounced in S₅ and S₃, particularly for characters with low heritability. The average standard error of difference between GCA effects increased with a decrease in s, with a steep slope for s < 7. The partial diallel analysis was more efficient for the estimation of the variance component of GCA than for SCA, as the estimate of SCA was biased upwards. Estimates of broad sense heritability obtained from the partial diallels agreed with the full diallel analysis better than the narrow sense estimates. Smaller partial diallels gave erratic estimates of heritability, particularly for the characters with low heritability.     

Fixed effect genetic analysis of a diallel cross in dry beans (Phaseolus vulgaris L.)

Summary A full diallel cross among four diverse homozygous strains of dry edible beans (Phaseolus vulgaris L.) was evaluated for yield, protein content, and culinary quality traits in the F₂ and F₃ generations in two locations. Interpretation of diallel effects [Method 1 Model I] using a fixed-effect genetic model made it possible to combine data from two generations into a single analysis and quantify the relative contributions of additive and dominance genetic effects to general (GCA) and specific (SCA) combining abilities. GCA was found to arise from three potential sources: additive effects, dominance interactions at homozygous loci, and average dominance interactions in hybrids involving the parent in question. SCA was found to be a function solely of dominance. Additive effects were the primary determinant of GCA and were highly significant. Specific dominance interactions were significant for seed yield, cooked bean moisture content, and texture but not for protein content. Texture was the only trait for which the additive-dominance model failed to provide an adequate fit to the data, suggesting that texture is significantly affected by epistatic interaction. One cross (Brazil-2 × Sanilac) was identified that exhibited a large heterotic effect for seed yield although the parents' additive effects were nonsignificant. Such a nicking effect was attributed to complementation between the two parents.Cooperative investigations of the Agricultural Research Service, U.S. Department of Agriculture and Michigan State University, East Lansing, MI 48824. Part of a thesis submitted by the senior author in partial fulfillment of the requirements for the Ph.D. degree at Michigan State University. Approved for publication by the Michigan Agricultural Experiment Station as Journal Article No. 11791. Research supported by USAID under a Title XII Bean/Cowpea CRSP and cooperative with Washington State University, Pullman, WA99164     

Combining ability in the F1 and F2 generations of diallel cross in hexaploid wheat (Triticum aestivum L. em. Thell)

The F(1) and F(2) progenies of a ten-parent diallel cross (excluding reciprocals) of hexaploid wheat (Triticum aestivum L. em. Thell) were analyzed for combining ability for quantitative and quality traits. The results indicated significant differences among the parents for general combining ability (gca) and crosses for specific combining ability (sca) for all the characters studied. The gca and sca components of variance were significant for all the traits. However, the gca component of variance was predominant indicating the predominance of additive gene effects for the traits studied. Among the parents Durgapura 65, HD 2285, Lok-1, Raj 1972 and HD 2329 were the best general combiners for grain yield and average to high combiners for tillers per plant, grain yield per spike, grains per spike and 1000-grain weight. The best specific crosses for grain yield were Sonalika x WH 157, HD 2428 x Durgapura 65, Durgapura 65 x Sonalika, HD 2428 x Lok-1 and CPAN 3004 x Raj 1972. The parent Raj 1972, Lok-1 and HD 2285 were the best general combiners for grain yield and protein content, however, Raj 3077 was the best general combiner for protein content. The most suitable specific crosses for protein content were HD 2329 x HD 2285, HD 2428 x Raj 1972 and CPAN 3004 x WH 157. Most of the specific crosses for grain yield as well as protein content involved high x average, average x average and average x poor general combiners. To ensure further increase in grain yield along with high protein, combinations of desirable yield components is advocated. Inclusion of F(1) hybrids showing high sca and having parents with good gca, into multiple crosses and/or bi-parental mating, or diallel selective mating could prove a worthwhile approach for further improvement of grain yield in bread wheat.     

Detection of genic interactions by analyzing the F2 generation of diallel crosses

Summary The ability of the Hayman and Jinks method of analysis of diallel crosses to detect genic interactions was studied in peanuts. Six traits, measured in the F₂ generation of a diallel cross of four cultivars, were analyzed. In F₂ analyses of the diallel, least squares estimates of variance components D, H₁ , H₂, F, E, were used as an additional criteria for the adequacy of the diallel model. They were found to be in substantial agreement with the tests based on W_r and V_r values, and probably more reliable. The regression of W_r on V_r was shown to be unsuitable to detect duplicate gene type of interactions; it was detected, however, by the ratio of the mean within-F₂ — family variance and the variance among the parents. Using the different criteria, duplicate genes type of interactions was detected for two traits, complementary genes type was detected for one trait and three traits were found to fit the additive-dominance model without any genic interactions.     

Changes in the levels of seven proteins involved in polypeptide folding and transport during endosperm development of two barley genotypes differing in storage protein localisation

The Russian barley cultivar Nevsky lacks ₃ hordein and accumulates most of its hordein in the lumen of the endoplasmic reticulum and only a minor portion in the vacuole. In wild type barley and all other temperate cereals, storage proteins are deposited in the vacuole. F1 crosses revealed that the Nevsky phenotype is recessive; but the extent of hordein accumulation in the endoplasmic reticulum in F2 endosperm lacking ₃ hordein was very much less than in the Nevsky parent. In order to study the Nevsky endosperm phenotype we have measured the levels of seven proteins and two mRNAs involved in protein folding in the ER lumen or ER to Golgi transport during endosperm development. The protein levels were unaltered in Nevsky as compared to the wild-type variety Bomi. When the levels of these seven proteins were correlated with the rate of hordein accumulation, four of these (HSP70, PDI, Sar1p and Sec18p) were consistently up-regulated with hordein synthesis. Accumulation of hordein in the endoplasmic reticulum appears to be determined by the absence of ₃ hordein, or the product of a gene closely linked to it, plus one or more other recessive genes.     

Analysis of Partial Diallel Crosses in Incomplete Blocks

With a large number of lines in a diallel cross experiment, the number of crosses becomes unmanageable to be accommodated in homogeneous blocks. To overcome this problem, a sample of crosses, known as partial diallel cross (PDC) is often used. The selection of a PDC is based on the criterion of high efficiency for the estimation of general combining ability (gca) effects. Even with a moderately large number of crosses, the use of incomplete blocks is necessary to obtain homogeneous experimental units. The analysis of data from a general PDC grown in general incomplete block designs is being described. An iterative scheme is being developed for obtaining a generalized inverse of the information matrix used in estimating gca effects. Properties such as connectedness and efficiency of mating designs embedded in environment designs are being examined. The paper also examines the universal optimality of some designs in a class of designs. An illustration of the numerical procedure is also presented.     

The HLE hybrid dysgenesis syndrome in the midgeChironomus thummi: Differences in the dysgenic potential between strains

After crossing experiments between twoChironomus subspecies the HLE hybrid dysgenesis syndrome occurs non-reciprocally in the F1 hybrids if the males of each of the fourCh. th. thummi strains (HL, So, G, Kr) are crossed to females ofCh. th. piger, strain E. In hybrid egg masses, differences in the strength of the abnormal trait reduced egg hatch were found between the differentpiger E ×thummi interstrain crosses. From the results it is concluded that eachthummi strain has a specific hybrid dysgenesis potential which determines the severity of the HLE syndrome in the hybrids. The HLE syndrome occurs also in the hybrids of crosses of differentCh. th. thummi strains. In these hybrids the syndrome is only obtained if the male parent descends from a strain which has a higher HLE hybrid dysgenesis potential than that strain from which the female parent is derived. These results demonstrate that the HLE syndrome is not restricted to the hybrids of thethummi × piger subspecies cross. In theCh. th. thummi interstrain crosses the hybrids of that cross direction are normal, which is reciprocal to the cross direction affected by the HLE syndrome. This is in contrast to thethummi × piger subspecies crosses, where the hybrids of one cross direction always show the HLE syndrome and the hybrids of the reciprocal cross direction the Rud syndrome. Thus, the present study also demonstrates that the HLE and Rud hybrid dysgenesis syndrome inChironomus are independent from one another.     

Segregation of random amplified DNA markers in F1 progeny of conifers

Summary The recently developed approach to deriving genetic markers via amplification of random DNA segments with single primers of arbitrary nucleotide sequence was tested for its utility in genetic linkage mapping studies with conifers. Reaction conditions were optimized to reproducibly yield clean and specific amplification products. Template DNA from several genotypes of Douglas-fir (Pseudotsuga menziesii) and white spruce (Picea glauca) were tested against eight ten-base oligonucleotide primers. Most of the tested primer/parent tree combinations yielded polymorphic PCR products (RAPD markers). Selected primers were then used in PCR reactions with template DNA isolated from offspring in Douglas-fir and black spruce diallel crosses among the same parental lines. The diallel study confirmed the appropriate inheritance of RAPD markers in the F₁ generation. The value of these dominant RAPD markers for genetic linkage mapping in trees was established from both theoretical and applied perspectives.     

The response of different genotypes of Fragaria × ananassa and their seedling progenies to in vitro micropropagation and the effects of varying the concentration of 6-benzylaminopurine in the proliferation medium

Two experiments are reported. In the first, six different strawberry genotypes were each grown in vitro on media containing five different concentrations of 6-benzylaminopurine (BAP) ranging from 0.5 M to 8.0 M. Responses of the genotypes to different BAP levels varied and the optimum levels for maximum proliferation were between 1 M and 8 M. In the second experiment the six genotypes were split into two groups of three and all crosses between the two groups were made. Seedlings from the nine progenies were then grown on media with the same five levels of BAP. Of the 12% of seedlings that did not proliferate, most were on media with the two lowest BAP levels (0.5 and 1.0 M). Seedlings from crosses involving the cv. Bogota proliferated least, while the three progenies with the breeding line ES652 as a parent had the highest mean proliferation rates. One cross, Bogota × Providence, had a particularly low mean proliferation rate. It was concluded that to optimize proliferation rate the medium must be tailored to the genotype. The best medium could not be reliably predicted from knowledge of the pedigree of a genotype.     

Cytogenetics of the F1 hybrid between cassava and ceara rubber,and its backcross

Cytogenetical studies of the F₁ hybrid between the commercially cultivated tuber crop, cassava (Manihot esculenta Crantz.) and the closely related wild speciesManihot glaziovii Muell. (Ceara rubber) used as donor specles for Cassava mosaic discase and drought-resistant genes and back crosses (to cassava parent) were made. The contrasting parental characters showed partial to total dominance in the F₁ hybrid, while the back cross plants were similar to cassava in most of their characters. Eleven of the twelve backeross plants exhibited resistance to Cassava mosaic under field conditions. Karyological similarities and differences as resolved on the basis of a comparative study of the karyotypes of the cassava parent and coara rubber were corroborated by the study of chromosomal pairing in the F₁ at pachytene. Major chromosomal differentiation in the two species involved three chromosomes of their haploid complement which were represented by three heteromorphic bivalent associations in F₁ each consisting of a probably basic chromosomal type and a derived type. Pachytene analyses of three back cross plants provided direct proof for random transmission of marker chromosomes of both the parents through male gametes of the F₁ hybrid. An increase in the chiasma frequency in the back cross plants over the F₁ hybrid at metaphase I stage was also observed. Pollen fertility of the backeross plants showed considerable variation.     

Improvement of β-amylase thermostability in transgenic barley seeds and transgene stability in progeny

The genetic improvement of enzymes important in the brewing process is one of the main goals of barley biotechnology. For the improvement of -amylase thermostability in barley seeds, we have already constructed a mutant thermostable -amylase gene, using site-directed mutagenesis and random mutagenesis to achieve the substitution of seven amino acids of the original barley -amylase. This sevenfold-mutant barley -amylase showed a thermostability increased by 11.6 °C compared to the original enzyme. In the present article, a thermostable -amylase gene under the control of the barley -amylase promoter was introduced to barley protoplasts, and fertile plants were generated from 9 independent transgenic lines. Subsequent analyses indicated that the thermostable -amylase gene was expressed and -amylase activity derived from both native and modified genes was detected in the seeds of 6 transgenic lines. The transgene was stably transmitted to progeny, and thermostable -amylase was synthesized in T₄ seeds, demonstrating that our strategy is applicable for the improvement of seed quality for industrial utilization.     

The inheritance of genetic markers in microspore-derived plants of barley Hordeum vulgare L.

Summary Biochemical, molecular and morphological markers have been used to monitor the segregation of alleles at major gene loci in microspore-derived lines of four spring barley crosses and their parents. Significant deviations from the expected Mendelian ratios were observed for four of the ten markers studied in the cross. Distorted ratios were associated with loci located on chromosomes 4H and 6H. The differential transmission of alleles was in favour of the responsive parent (Blenheim) used in the anther culture studies. For the -Amy-1 locus on chromosome 6H, the preferential transmission of Blenheim alleles was most pronounced in the haploid regenerants that were colchicine treated. These results are discussed in relation to the genetic control of androgenetic response in barley and with respect to the exploitation of another culture in barley improvement.     

Evidence on the nature and origins of endosperm dosage requirements in Solanum and other angiosperm genera

Success of seed development following sexual crosses is primarily dependent on proper endosperm function and development. The failure to produce triploids, or triploid block in 4x×2x crosses served as the impetus for numerous studies of embryo and endosperm to attempt to explain cross failure. Early explanations were based upon a concept of a 232 ploidy balance between maternal tissue, endosperm, and embryo. Subsequent studies done with maize demonstrated that normal endosperm development in intraspecific maize crosses is dependent solely on having a 21 maternal to paternal genome dosage in the endosperm. These results have been modified and extended to solanaceous species in the form of an endosperm dosage system in which empirically determined factors must bear the same 21 relationship for crosses to succeed. Crossing behavior of these species suggest that the system is polygenically controlled and regulates both interspecific and intraspecific crosses. Endosperm dosage systems explain many aspects of species evolution, but the system appears to have originated as an ancient means of ensuring diploid fidelity.     

Genetic diversity for RFLPs in European maize inbreds

Summary Restriction fragment length polymorphisms (RFLPs) have been proposed for the prediction of the yield potential of hybrids and the assignment of inbreds to heterotic groups. Such use was investigated in 66 diallel crosses among 6 flint and 6 dent inbreds from European maize (Zea mays L.) germ plasm. Inbreds and hybrids were evaluated for seven forage traits in four environments in the Federal Republic of Germany. Midparent heterosis (MPH) and specific combining ability (SCA) were calculated. Genetic distances (GD) between lines were calculated from RFLP data of 194 clone-enzyme combinations. GDs were greater for flint x dent than for flint x flint and dent x dent line combinations. Cluster analysis based on GDs showed separate groupings of flint and dent lines and agreed with pedigree information, except for 1 inbred. GDs of all line combinations in the diallel were partitioned into general (GGD) and specific (SGD) genetic distances; GGD explained approximately 20% of the variation among GD values. For the 62 diallel crosses (excluding 4 crosses of highly related lines), correlations of GD with F₁ performance, MPH, and SCA for dry matter yield (DMY) of stover, ear, and forage were positive but mostly of moderate size (0.09r0.60) compared with the higher correlations (0.39r0.77) of SGD with these traits. When separate calculations were performed for various subsets, correlations of GD and SGD with DMY traits were generally small (r<0.47) for the 36 flint x dent crosses, significantly positive (r<0.53) for the 14 flint x flint crosses, and inconclusive for the 12 dent x dent crosses because of the lack of significant genotypic variation. Results indicated that RFLPs can be used for assigning inbreds to heterotic groups. RFLP-based genetic distance measures seem to be useful for predicting forage yield of (1) crosses between lines from the same germ plasm group or (2) crosses including line combinations from the same as well as different heterotic groups. However, they are not indicative of the hybrid forage yield of crosses between unrelated lines from genetically divergent heterotic groups.     

Comparison of chlorophyll fluorescence kinetics and photochemical activities of isolated chloroplasts in genetic analysis of Lycopersicon esculentum Mill. hybrids

In experiments with 5x5 diallel cross of tomato cv. Smrické contents of individual chlorophylls (Chl) and carotenoids (Car), Chl fluorescence kinetic parameters (Fv/Fm, qp, qN and II), activities of photosystem 2 and of the whole electron transport chain (Hill reaction) were compared during plant ontogeny with the aim to detect genetically determined differences among genotypes. The differences in Chl fluorescence kinetic parameters and Hill reaction activities were more affected by leaf development than by plant genotype, although the differences between reciprocal crosses were found in the former ones. Some additive genetic effects may participate in the genetic determination of qN. Statistically significant non-additive genetic effects in inheritance of the total Chl content were found. Detailed pigment analysis revealed additivity together with differences between reciprocal crosses in individual Car and Chl. The highest general combining ability was found for the parental line AM.     

Induction of glycanolytic activities by Streptomyces sp. QM-B814 growing on different glucopolymers

The actinomycete Streptomyces sp. QM-B814 is able to use polymeric glucans as sole carbon source by the induction of specific extracellular enzymatic systems. Carboxymethylcellulose and barley -glucan were the most effective inducers of endoglucanase activities towards -(1,4) and mixed -(1,3)-(1,4) linkages. Avicelase activity was induced by Avicel and carboxymethylcellulose. The higher activity towards filter paper (cellulase) was measured in cultures growing in Avicel, carboxymethylcellulose, and barley -glucan. Laminarinase activity was induced only by laminarin and lichenan failed to induce significant activity levels against any of the substrates tested.     

Allozyme marker loci associated with favorable alleles for grain yield in maize

Summary The evaluation of germplasm to identify its potential as a source of new favorable alleles is a time-consuming phase of maize hybrid breeding programs. The objective of this paper was to study the relationship between allozyme diversity and quantitative estimators of the relative number of new favorable alleles for grain yield, present in donor lines but not present in the elite hybrid. Twenty-two maize inbred lines representing heterotic groups from the United States (US) and Yugoslavia (YU) were used as donors to estimate the presence of new favorable alleles for grain yield improvement for the hybrid B73 x Mo17. In a second experiment, a 15-line diallel was grown, and 13 single crosses differing in allozyme relatedness measure (ARM) and heterotic grouping were considered as targets to be improved by the remaining 13 lines. Minimally biased estimates of new favorable alleles for grain yield (G) and ARM values were made for all donor lines within each target hybrid. Donor lines were grouped in four allozyme-pedigree classes for each target hybrid to compare the effect of allozyme diversity with pedigree diversity. Pedigree dissimilarities had significant effects on G estimates. Dissimilar pedigree classes had higher G estimates than similar pedigree classes. Allozyme differences between donor inbred lines and target hybrids had inconsistent effects on G estimates. Significant differences in G estimates among allozyme classes were found for 31% of the target hybrids. Classes with similar allozymes had higher G estimates more frequently than classes with disimilar allozymes. Correlation coefficients between G estimates and ARM values were low and not significant for 12 of the 14 target hybrids.This project was partially supported by the USDA and Republic Funds for Scientific work of Serbia through funds available to the United States-Yugoslav Joint Board of Scientific and Technological Cooperation, Project JFP 662