Epistasis in maize (Zea mays L.)

Summary Three flint and three dent maize (Zea mays L.) inbred lines, their possible F₁ crosses, F₂ and backcross progenies, and all possible three-way crosses were evaluated in a three-year experiment for yield, ear moisture, and plant height. The purpose was to estimate genetic parameters in European breeding materials from (i) generation means analysis, (ii) diallel analysis of generation means, and (iii) analysis of F₁ and three-way cross hybrids. Method (i) was based on the F-metric model and methods (ii) and (iii) on the Eberhart-Gardner (1966) genetic model; both models extended for heterotic maternal effects.Differences among generation means for yield and plant height were mainly attributable to dominance effects. Epistatic effects were significantly different from zero in a few crosses and considerably reduced heterosis in both traits. Additive x additive and domiance x dominance effects for yield were consistently positive and negative, respectively. Significant maternal effects were established to the advantage of generations with a heterozygous seed parent. In the diallel analysis, mean squares for dominance effects were greater than for additive effects for yield and plant height but smaller for ear moisture. Though significant for yield and plant height, epistatic variation was small compared to additive and dominance variation. Estimates of additive x additive epistasis for yield were significantly negative in 11 of 15 crosses, suggesting that advantageous gene combinations in the lines had been disrupted by recombination in the segregating generations. The analysis of hybrids supported the above findings regarding the analysis of variance. However, the estimates of additive x additive epistasis for yield were considerably smaller and only minimally correlated with those from the diallel analysis. Use of noninbred materials as opposed to materials with different levels of inbreeding is considered the main reason for the discrepancies in the results.     

Combined linkage and linkage disequilibrium QTL mapping in multiple families of maize (Zea mays L.) line crosses highlights complementarities between models based on parental haplotype and single locus polymorphism

Advancements in genotyping are rapidly decreasing marker costs and increasing marker density. This opens new possibilities for mapping quantitative trait loci (QTL), in particular by combining linkage disequilibrium information and linkage analysis (LDLA). In this study, we compared different approaches to detect QTL for four traits of agronomical importance in two large multi-parental datasets of maize (Zea mays L.) of 895 and 928 testcross progenies composed of 7 and 21 biparental families, respectively, and genotyped with 491 markers. We compared to traditional linkage-based methods two LDLA models relying on the dense genotyping of parental lines with 17,728 SNP: one based on a clustering approach of parental line segments into ancestral alleles and one based on single marker information. The two LDLA models generally identified more QTL (60 and 52 QTL in total) than classical linkage models (49 and 44 QTL in total). However, they performed inconsistently over datasets and traits suggesting that a compromise must be found between the reduction of allele number for increasing statistical power and the adequacy of the model to potentially complex allelic variation. For some QTL, the model exclusively based on linkage analysis, which assumed that each parental line carried a different QTL allele, was able to capture remaining variation not explained by LDLA models. These complementarities between models clearly suggest that the different QTL mapping approaches must be considered to capture the different levels of allelic variation at QTL involved in complex traits.     

Optimum prediction of three-way crosses from single crosses in forage maize (Zea mays L.)

Summary Three-way cross means were predicted with formulae involving linear functions of general (GCA) and specific combining ability (SCA) effects estimated from single-cross factorials between genetically divergent populations. Data from an experiment with 66 single-cross and 66 three-way cross forage maize (Zea mays L.) hybrids was used for comparing the prediction formulae. The genotypic correlation (r) between observed and predicted three-way crosses increased with increasing , the weighting factor of SCA effects, for plant height and ear dry matter (DM) content. It displayed slightly convex curves for total and stover DM yield, ear percentage, and metabolizable energy content of stover. For Jenkins' method B, r was considerably less than 1.0 for all traits, indicating the presence of epistasis. The square root of heritability (h) of the predicted means decreased with increasing , the reduction being small with a greater number of test environments. Using the product r·h as a criterion of efficiency, none of the prediction methods was consistently superior and the differences among them were rather small (< 7.5%) for all traits, irrespective of the number of test environments. We recommend evaluating the GCA of a greater number of lines from each parent population in testcrosses with a small number of elite lines from the opposite population. All possible three-way or double crosses between both sets of lines should be predicted by Jenkins's method C. This procedure allows one to select with a higher intensity among the predicted hybrids and thus should increase the genetic gain.Extended version of a paper (Geiger et al. 1986) read at the sixth meeting of the EUCARPIA Section Biometrics in Plant Breeding held at Birmingham, UK, July 28–August 1, 1986     

Thiols in cadmium- and copper-treated maize (Zea mays L.)

Maize plants (Zea mays L. cv. Honeycomb F-1) were grown on quartz sand containing amounts of Cd or Cu which resulted in comparable internal contents in the roots. Fresh and dry weights and the content of Cd or Cu were measured in roots and shoots after eight weeks. In addition, cysteine, γ-glutamylcysteine (γEC), glutathione (GSH) and the thiols in heavy-metal-binding peptides (HMBPs) were determined in the roots. At low internal contents, Cd and Cu inhibited root growth to the same extent. Inhibition by Cu was enhanced, however, at high internal contents, indicating that Cu was more toxic than Cd. Separation of extracts from roots of Cd- and Cutreated plants on a Sephadex G-50 column resulted in HMBP complexes with relative molecular masses (M_rs) of 6200 and 7300, respectively. Separation of these HMBP-complexes using HPLC resulted in a distinct pattern of thiol compounds for each heavy metal. The accumulation of HMBPs was linearly dependent on the content of Cd at all values examined. In Cu-treated roots, HMBP accumulation was linearly dependent on the internal Cu content only up to 7.1 μmol·g^?1 dry weight. At internal contents which caused an enhanced inhibition of root growth, no further significant increase in the HMBP content was detected. At these internal Cu contents an increased transport of Cu to the shoot was measured. This result indicates that HMBPs are involved in reducing heavy-metal transport from roots to shoots.     

Biochemical composition of maize (Zea mays L.) pollen

Summary Pollen grains containing either the Wx, wx, Su ₁, su ₁, Sh ₂ or sh ₂ alleles were stored at 0, 1, 2, 3, 4 and 5 days at 2 °C. After each storage period, a portion of pollen from each genotype was analyzed for free amino acid content. Over all genotypes, storage significantly altered the content of all 16 amino acids measured. With increasing storage, a relatively consistent increase in aspartic acid, isoleucine, leucine, phenylalanine, ethanolanine, aminobutyric acid, NH₃ and lysine was found. A relatively consistent decrease in glutamic acid, proline, glycine and alanine occurred with increasing storage. No consistent response to storage was obtained with threonine-serine, valine, histidine and the unknown. Apparently, storage or stage of viability loss has a pronounced effect on amino acid metabolism in maize pollen grains. The experiment was designed so that comparisons free of genetic background effects could be made between alleles at each locus. Significant allele X storage interactions at each locus were found as follows: at the waxy locus, aspartic acid, glycine, alanine and ethanolanine; at the sugary locus, aspartic acid, alanine, ethanolanine and aminobutyric acid; and at the shrunken locus, aspartic acid, alanine, valine, leucine and ethanolanine. Amino acid metabolism is apparently influenced by the action of the alleles at these loci. The differences between the loci in the amino acids affected indicate the different areas of amino acid metabolism are influenced by each locus.Journal Series Paper No. 4425, Florida Agricultural Experiment Station.     

Biochemical composition of maize (Zea mays L.) pollen

Summary Mean percentages of sugars, water-soluble polysaccharides, starch, total carbohydrates and lipids were 40.1, 7.4, 28.6, 76.1, and 1.8 respectively. Differences among the mutants were found only for water-soluble polysaccharides with both wx and sh ₂ decreasing the percentage a small but significant amount. In terms of the various carbohydrates measured, no correlation was found between the expression of these mutants in the pollen and the kernel.

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Zusammenfassung Die Untersuchung des Kohlenhydrat- und Lipid-Gehaltes im Pollen der Endosperm-Mutanten waxy, shrunken und sugary vom Mais ergab einen gemittelten Gehalt an Zucker von 40,1%, wasserlöslichen Polysacchariden 7,4%, Stärke 28,6, Gesamtkohlenhydraten 76,1% und Lipiden 1,8%.Unterschiede zwischen den Mutanten wurden lediglich hinsichtlich der wasserlöslichen Polysaccharide gefunden, und zwar bei wx und sh ₂, die einen signifikant niedrigeren Gehalt aufwiesen.Hinsichtlich der verschiedenen analysierten Kohlenhydrate wurde keine Korrelation zwischen der Expression der Mutanten im Pollen und in den Karyopsen gefunden.

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Herrn Professor Dr. J. Straub (Köln-Vogelsang) zu seinem 60. Geburtstag gewidmet.

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Journal Series Paper No. 3621, Florida Agricultural Experiment Station.     

Heterosis among populations of maize (Zea mays L.) with different levels of exotic germplasm

Summary Thirteen maize (Zea mays L.) populations including five adapted, five adapted x exotic, two composites of adapted and exotic, and one exotic selected for adaptability were crossed in a diallel mating system. The parents and 78 crosses and nine check hybrids were evaluated for grain yield and plant height in five environments. The Gardner-Eberhart model Analysis II indicated that additive and nonadditive gene effects accounted for 60 and 40% of the total variation among populations, respectively, for grain yield and 86% and 14% of the total variation, respectively, for plant height. Components of heterosis were significant in the combined analysis for both traits. Adapted Corn Belt populations tended to have higher performance in crosses and greater values of variety heterosis than 50% adapted populations. Nebraska Elite Composite, Corn Belt x Mexican, and Corn Belt x Brazilian showed high mean yields in crosses, however, they were not among those with high estimates of variety heterosis. One exotic population (Tuxpeno x Antigua Grupo 2) and three adapted populations [307 Composite, NB(S₁)C-3, and NK(S₁)C-3] might be combined together to form a high-yielding population. It may be possible to synthesize two useful populations for reciprocal recurrent selection by grouping Tuxpeno x Antiqua Grupo 2, NB(S₁)C-3, and NS(FS)LFW-8 into one population and NK(S₁)C-3, KrugxTabloncillo, and 307 Composite in the other one.Contribution from the Department of Agronomy, University of Nebraska, Lincoln, NE 68583. Published as Paper No. 8011. Journal Series, Nebr. Agric. Exp. Station. Research was conducted under Project 12-049     

Hydroxylation reactions in roots of maize (Zea mays L.)

The authors succeeded in demonstrating accumulation of tyrosine and p-coumaric acid in three day-old roots of maize (Zea mays L.) fed with L-phenylalanine and cinnamic acid. Phenylpyruvic acid applied under the same conditions gave rise to phenylalanine, indicating the presence of the corresponding transferase activity. Even simultaneous application of inhibitors of transaminase activity—hydroxylamine and isonicotinic acid hydrazide—did not result in the formation of p-hydroxyphenylpyruvic acid.     

Hydroxylation reactions in roots of maize (Zea mays L.)

The authors succeeded in demonstrating accumulation of tyrosine and p-coumaric acid in three day-old roots of maize (Zea mays L.) fed with L-phenylalanine and einnamie acid. Phenylpyruvic acid applied under the same conditions gave rise to phenylalanine, indicating the presence of the corresponding transferase activity. Even simultaneous application of inhibitors of transaminase activity — hydroxylamine and isonicotinic acid hydrazide — did not result in the formation of p-hydroxyphenylpyruvic acid.     

Biochemical composition of maize (Zea mays L.) pollen

Summary Proline was the most abundant amino acid with a mean value of 186.28 moles/mg dry pollen. The other amino acids tested were below 33 moles/mg dry pollen. The mutant wx significantly increased aspartic acid, valine, histidine and an unknown but significantly decreased aminobutyric acid. The mutant sh ₂ significantly increased glutamic acid, proline, lysine, histidine and an unknown but significantly decreased aspartic acid and aminobutyric acid. The effect of su ₁ was altered by the genetic background involved. In one genetic background, su ₁ did not significantly increase any amino acid but significantly decreased alanine and aminobutyric acid. However, in a distinctly different background, su ₁ significantly increased aminobutyric acid but significantly decreased aspartic acid and glutamic acid. Apparently the genetic background is capable of producing major shifts in the amino acid pattern in addition to the action of these mutants.The fatty acids, palmitic and linolenic were the most common with percentages of 54.1 and 34.4 respectively. The mutants tested did not affect the fatty acid distribution.

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Zusammenfassung Prolin war die am reichlichsten vorkommende Aminosäure mit einem mittleren Gehalt von 186,28 Mikromol per mg trockenen Pollen. Die anderen Aminosäuren erreichten weniger als 33 Mikrogramm per mg trockenen Pollen.Die Mutante wx zeigte einen signifikant erhöhten Gehalt an Asparaginsäure, Valin, Histidin, sowie einer nicht identifizierten Komponente, während der Gehalt an -Aminobuttersäure signifikant erniedrigt war. Die Mutante sh ₂ ist gekennzeichnet durch einen signifikant erhöhten Gehalt an Glutaminsäure, Prolin, Lysin, Histidin, sowie einer unbekannten Fraktion; der Gehalt an Asparaginsäure und -Aminobuttersäure war dagegen signifikant erniedrigt. Die Wirkung des mutierten Gens su ₁ wurde durch das übrige Genom, in dem es sich befand, geändert. In dem einen genetischen Milieu verursachte su ₁ keine signifikante Erhöhung des Gehaltes irgend einer Aminosäure, während der Gehalt an Alanin und -Aminobuttersäure signifikant erniedrigt war. In einem anderen genetischen Milieu jedoch zeigte su ₁ eine signifikante Erhöhung der -Aminobuttersäure; Asparaginsäure und Glutaminsäure waren signifikant erniedrigt.Offensichtlich ist das übrige Genom zusätzlich zu der Wirkung der genannten Mutanten in der Lage, wesentliche Verschiebungen im Verteilungsmuster der Aminosäuren zu verursachen.Von den Fettsäuren wurden am häufigsten Palmitin- und Linolen-Säure mit einem Gehalt von 54,1 bzw. 34,4% gefunden. Die untersuchten Endosperm-Mutanten zeigten keinen Einfluß auf die Fettsäureverteilung im Pollen.

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Journal Series Paper No. 3468, Florida Agricultural Experiment Station.     

Induced cytomictic diversity in maize (Zea mays L.) inbred

Mutation breeding has been used for improving oligogenic and polygenic characters, disease resistance and quantitative characters including yielding ability. The cytological stability of maize inbred lines is an important consideration in view of their extensive use in genetics and plant breeding research. Investigation in Zea mays L. confirms that the migration of chromosomes is a real event that cannot be misunderstood as an artifact produced by fixation or mechanical injuries. During present investigation, we found that out of six inbred lines of Zea mays L. viz. CM-135, CM-136, CM-137, CM-138, CM-142 and CM-213 at various treatment doses of gamma irradiations viz. 200, 400 and 600 Gy, some of the plants of inbred line CM- 138 at 200 Gy dose displayed characteristic cytoplasmic connections during all the stages of meiosis. Four plants from this treatment set were found to be engaged in a rare phenomenon reported as "Cytomixis". It elucidates that in inbred of Zea mays L., induced cytomixis through gamma rays treatment may be considered to be a possible source of production of aneuploid and polyploid gametes. This phenomenon may have several applications in Zea mays L. improvement in the sense of diversity and ever yield potential.     

The nucleolus organizer region of maize (Zea mays L.)

Maize plants were produced partially triploid for the heterochromatic segment of the nucleolus organizer region (NOR) or partially triploid or tetraploid for the site giving rise to the secondary constriction of the NOR. These partially hyperploid plants were characterized in terms of chromosome and/or nucleolar constitution by light microscopy at pachytene, diakinesis, and quartet stages of microsporogenesis. DNA's of the various partially hyperploid plants and appropriate controls were extracted and hybridized with ³H-rRNA. The heterochromatic segment of the NOR was found to contain most of the rRNA cistrons, but has little or no interaction with the nucleolus. In contrast with the heterochromatic segment, the site giving rise to the secondary constriction contains few rRNA cistrons but is active in nucleolar formation as viewed at pachytene, diakinesis and quartet stages.     

Monitoring transgene expression levels in different genotypes of field grown maize (Zea mays L.)

Transgenes in commercially available genetically modified plants are generally controlled by strong constitutive promoters to ensure a high level of expression at all stages of cultivation. Constitutive promoters however are influenced by a wide range of factors, and expression profiles of the transgenes in multiple genetic backgrounds have not yet been extensively studied. In this study a powerful expression profiling methodology for transgenic maize (Zea mays L.) is demonstrated on a large scale, analysing thousands of data points from three genotypes of herbicide and insect pest tolerant transgenic maize. Martonvásár inbred lines were crossed with LH244 maize line containing the MON 88017 events, and leaf tissue from the sixth backcross generation was sampled at four relevant phenological phases. Relative expression levels were determined using 18S rRNA as a reference and detailed statistical analysis performed. Expression levels of both transgenes are varied throughout plant development, and the interaction between the genetic background and phenophase are significant (p < 0.05). Expression is present at a significant level throughout all the phenological stages. We found that the genetic background has a significant (p < 0.01) effect on transgene expression levels in the case of the CP4epsps transgene, but not in the case of cry3Bb1, implying that the sensitivity of different constitutive promoter constructs to the effects of the genetic background is different.     

Recovery of maize (Zea mays L.) inbreds and hybrids from chilling stress of various duration: photosynthesis and antioxidant enzymes

The differences between two maize (Zea mays L.) inbred lines and their F1 hybrids in their response to chilling periods of various duration (1, 2, 3 or 4 weeks) and subsequent return to optimum temperatures were analysed by the measurement of the photosystem (PS) 1 and 2 activity, the photosynthetic pigments' content and the activity of antioxidant enzymes. The PS2 activity and the chlorophyll content decreased in plants subjected to 3 or 4 weeks of chilling, but not in those subjected to 1 or 2 weeks of chilling. This decrease was more pronounced in inbreds compared to their hybrids. The activity of superoxide dismutase did not much change with the increasing length of chilling period in the inbreds but decreased in the hybrids, the glutathione reductase activity increased in both types of genotypes but more in the inbred lines, while for ascorbate peroxidase and catalase the changes in parents-hybrids relationship did not show any specific trend. The PS1 activity and the carotenoids' content was not much affected.     

Ultrastructural studies on pollen embryogenesis in maize (Zea mays L.)

Maize anthers have been induced on modified N6 medium to produce embryoids. Different stages from the cultures were sampled and prepared for microscopical examination. The microspores at the onset of culture were in an early developmental stage, with the nucleus and numerous organelles centred in the middle, surrounded by many small vacuoles with a lipid content. The binuclear pollen grains contained small vesicles and much starch. The partially condensed vegetative nucleus indicated participation of the vegetative component in the formation of multicellular pollen grains (MPGs). Several MPGs have been observed which differed in morphology. We suggest, on the basis of these ultrastructural observations, that in maize mainly the vegetative cell contributes to the MPG which further develops directly into embryoids.     

Plot size for progeny selection in maize (Zea mays L.)

Summary Six progeny trials that included 147 half-sib progenies of maize (Zea mays L.) population ESALQ PB-5 were conducted for the purpose of studying plot size and its consequences in recurrent selection programs. The progenies were evaluated in three 7x7 duplicate simple lattice experiments using one-row plots of 5 m². At harvest each plot was partitioned into five sub-plots (sampling units), and data was collected from each sampling unit. At the same time and place the same progenies were evaluated in three 7x7 duplicate simple lattice experiments using 1-m² (linear row with 5 plants) plots. Data were collected for plant and ear height, ear diameter, total ear weight, and total grain yield. The data were combined by using adjacent sampling units, and the analyses were performed by considered five plot sizes in addition to those of the independent trials with 1-m² plots. The experiments with 1-m² plots were less efficient in discriminating for yield traits among progenies than those with 5-m² plots. The combination of plot size and number of progenies evaluated indicated that an optimum plot size for yield was between 3 and 4 m², or 15–20 plants per plot. With such sizes the expected gain was maximized for the four replications used in this study. If the total area covered by each progeny is constant, the maximum gain from selection, however, is attained by decreasing plot size and increasing the number of replications. The minimum size of plots is, however, limited by practical or theoretical criteria. Plot size affected the estimates of additive genetic variance, coefficient of heritability, and genetic coefficient of variation for all of the traits. No practical limitation was observed for conducting experiments with 1-m² plot.