High Frequencies of Fertilization and Haploid Seedling Production in Crosses Between Commercial Hexaploid Wheat Varieties and Maize

Nineteen commercial hexaploid wheat varieties were crossed with the maize F₁ hybrid ‘Seneca 60’. Fertilization frequencies ranged from 32.1 % to 47.5 % of pollinated florets (mean 39.5 %) in the 14 winter wheat varieties and from 40.7 % to 51.4 % (mean 47.8 %) in the five spring wheat varieties. In some cases only an endosperm was formed and the frequencies of embryo formation were therefore slightly lower, being 28.2 % to 45.9 % (mean 36.4 %) for winter wheats and 39.8 % to 48.6 % (mean 45.1 %) for spring wheats. Mean values were significantly higher in the spring wheats but no significant variation was found between varieties within the spring or winter categories. In the five spring wheats the mean yield of embryos, and hence the potential yield of haploid plants, was 3.4-fold higher than with the tetraploid Hordeum bulbosum clone PB179. For the 14 winter wheats the figure was 10.9-fold higher. These differences were highly significant (p < 0.001) in all varieties. A single 2,4-D treatment given to spikes one day after pollination with maize enabled embryos to be recovered from all 19 varieties. A total of 311 embryos were recovered from 950 florets (an average of 7.3 embryos per spike) of which 191 germinated, giving an average yield of one haploid plant for every 5.0 florets pollinated (4.4 haploid plants per spike).     

A simple technique for the identification of embryo-carrying seeds from wheat × maize crosses prior to dissection

Embryo rescue is an important step in the wheat x maize system of induced haploidy in wheat. On average, only one-third of the seeds carry embryos, but they all have to be dissected because no morphological distinction is available to distinguish between seeds with and without embryos. We here report a simple technique in which immature seeds from wheat × maize crosses are placed on a transparent surface and illuminated from above. The free-floating embryo, settled at the bottom of the seed, can be seen from below or in an image reflected on a plane mirror. Using this technique 97.8% of the embryo-containing seeds could be detected prior to dissection. The technique will be useful in reducing the time and labour involved in embryo culture, thereby scaling up haploid production.     

Improved Techniques for Haploid Production in Wheat using Chromosome Elimination

Wheat × maize and wheat × pearl millet crosses have proved efficient for haploid production using various genotypes of wheat; 22 and 27 % of florets produced embryos. In favourable conditions 6—9 haploid plants per spike were produced. The following simplifications or improvements in technique are recommended: 1. Only a single treatment with an aqueous solution of dicamba or 2,4-D (50–100 ppm) for embryo stimulation in vivo; 2. Application by spraying or dipping the spikes; 3. Application time two to four days after pollination; 4. Embryo rescue 15 to 18 days after pollination; 5. Crosses without emasculation are possible if pollination occurs 1–2 days before anthesis. More than 450 haploids and some doubled haploid (DH) lines (after colchicine treatment in vitro) were produced using these methods. No hybrid plants, chromosome additions or substitutions were found.     

Genetic Variability of Rubisco in Tetraploid Wheats (Triticum turgidum)

Nineteen tetraploid wheats (Triticum turgidum), differing in their agronomical behaviour and geographic origins were grown in two experiments and analysed for the following physiological traits: soluble leaf protein content, rubisco content, initial and potential carboxylase activities of this enzyme as well as its relative content in percentage of protein, the specific activity of the enzyme and its activation state. The results show a high genetic variability for all studied traits and more particularly for the initial carboxylase activity. High phenotypic correlation was observed between the rubisco content and its potential activities, in the first experiment. Correlation between souble leaf protein and rubisco content was significant in 9 genotypes of the second experiment.     

Variation in Doubled Haploid Plants of Wheat Obtained through Wheat (Triticum aestivum) × Maize (Zea mays) Crosses

Variation was investigated in 110 doubled haploid (DH) lines of wheat derived from wheat × maize crosses. Field observation revealed visible variations in 15 DH2 lines such as extreme dwarfism, low seed fertility, alteration of spike type and stripes. Six agronomic traits, i.e., heading date, spike number/ plant, culm length, spike length, seed fertility and grain weight were statistically analyzed in the DH2 and DH3 generations. Out of the 88 DH2 lines/DH3 groups, 26 %/64 % showed significant differences from the parental variety in the means of one or more traits. Ranges of the DH3 lines were larger than those of the DH2 lines, except for spike number/plant. Furthermore, analyses of variance within and between DH lines showed the presence of heterogeneity/heterozygosity in the DH2 lines/plants. These results indicated the occurrence of gametoclonal variation in the DH lines. It is considered that most of the variations detected were due to the colchicine treatment rather than to the 2,4-D treatment or in vitro culture.     

Production of Doubled Haploids by Anther Culture and Wheat X Maize Method in a Wheat Breeding Programme

Utilization of the doubled haploid method of breeding usually shortens the time to cultivar release, and methods of haploid production need evaluation in a breeding programme. Thirty-eight different three-way crosses were tested for anther culture response. On average 5.8 percent of the anthers cultured produced calli. Three crosses were found recalcitrant for callus induction. Overall, the anther culture method produced 0.6 plantlet per 100 anthers cultured. Five crosses with an average of 5.8 and 2.8 percent of anthers producing calli and plantlets, respectively, were compared using anther culture and wheat × maize crosses. Non-responsive genotypes for callus induction and plantlet formation in the anther culture method proved to be good parental material in wheat × maize crosses. The average percentages of embryo formation and plantlet production in wheat × maize crosses were 10.3 and 4.7, respectively. Anther-derived plants were cytologically unstable, whereas all the plants regenerated from wheat × maize crosses were haploids (n = 21 chromosomes). The chromosome numbers of the polyhaploids were doubled with a colchicine treatment. Improvement of the two haploid production methods to facilitate their efficient use in a breeding programme is discussed.     

Production of haploids in bread wheat, durum wheat and hexaploid triticale crossed with pearl millet

Pearl millet is an efficient alternative to maize as a pollen source for haploid production in bread wheat. To compare haploid production frequencies in other Triticeae species, the crossabilities of two genotypes each of bread wheat, durum wheat and hexaploid triticale with four pearl millet genotypes and a maize control were examined. Embryos were obtained from crosses of all three species with both pearl millet and maize. However, significant differences in crossability were found among the three species (10.5–79.8% seed development and 1.4–15.8% embryo formation), as well as among genotypes of durum wheat (7.2–23.7% and 2.1–6.4%) and hexaploid triticale (0.3–20.6% and 0.1–2.7%). Crossability of bread wheat with pearl millet was relatively high. Haploid plants were regenerated from crosses of all three species with pearl millet. As in the case of maize crosses, low crossabilities of durum wheat and hexaploid triticale with pearl millet can be attributed to the absence of D-genome chromosomes.     

Genetic Variation for Anther Callus Inducibility in Crosses of Highly Culturable Winter Wheats

Heritability studies suggest that variation among genotypes for callus inducibility should respond to selection. The objective of this study was to determine if crosses among parents with high callus initiation frequencies would produce segregation from which genotypes with improved callus inducibility could be selected. The winter wheat (Triticum aestivum L.) cultivars‘Centurk 78’, ‘Norstar’ and ‘Norwin’ were chosen as parents, based on their high callus initiation frequencies and diverse parentage. F₂ populations from crosses between each of these parents were evaluated for callus initiation frequencies and callus vigor as determined by growth rate and healthy appearance. Heterosis was detected for callus initiation in the‘Norwin’בNorstar’ cross and for callus vigor in all crosses. Broad sense heritability estimates of 0.25 and 0.37 for callus initiation in the ‘Centurk 78’בNorstar’ and ‘Norwin’בNorstar’ crosses, respectively, suggest this trait can be improved by selection. Heritability estimates for callus vigor were less promising, ranging from 0.11 to 0.24.     

不同小麦组合与玉米杂交产生单倍体的差异

实验采用３种类型、１５个小麦Ｆ１组合与玉米杂交种中单１２０进行杂交，研究不同小麦基因型单倍体产生的差异。     

AFLP analysis indicates no introgression of maize DNA in wheat x maize crosses

Amplified fragment length polymorphisms (AFLPs) were used to follow the possible introgression of maize DNA into haploids of wheat as a side‐effect of exploiting wheat x maize hybridization for haploid production. AFLPs were generated with 64 MseI/ EcoRI and 64 MseI/ PstI primer combinations, and the AFLP profiles of haploids were tested against those of maize and of the regular wheat varieties involved in the crosses. On average, 45.1 and 110.7 fragments were produced per assay with the MseI/EcoRI and MseI/PstI combinations, respectively. Different numbers of fragments were produced for wheat and maize: an average of 81 in the haploid, 80 in the wheat parent, and only 67.1 in maize. No evidence was found for introgression of maize into the wheat genome. Three unique AFLP fragments were detected in haploids, which were not present in the parental wheat genotypes. These ‘novel’ AFLP bands in the haploids could be caused by nucleo‐cytoplasmic interaction in the hybrid zygote. Such instability in the wheat genome is defined as temporal, as it was not detected in further generations when colchicine‐doubled progeny of the haploids was tested for the presence of polymorphic fragments.     

Durum wheat × maize crosses for haploid wheat production: Influence of parental genotypes and various experimental factors

To improve haploid plant production in durum wheat, the haplomethod involving intergeneric crossing with maize followed by embryo rescue was used. The influence of parental genotypes and various experimental factors were studied. Ten cultivars of Triticum turgidum ssp. durum (female parent) were crossed with eight genotypes of Zea mays. After pollination, plant stems were either maintained in situ or cut near the base and kept in a 2,4‐dichlorophenoxyacetic acid (2,4‐D)‐sucrose solution. Ten to 18 days after pollination, embryos were excised from developed ovaries and cultured on one of MS, MS/2, or B5 media. Haploid embryos and plants were obtained (78 green haploid plants regenerated in 0 year). The wheat genotype was significant for ovary development, embryo and plant formation, whereas the maize genotype was significant only for embryo formation. Detailed results of all crosses showed the best crossing partner for each wheat genotype. Cutting the plant stems after pollination gave better results than maintaining them in situ. The optimal stage for embryo rescue was 14 days and B5 and MS/2 media were more efficient than MS for embryo culture.     

Epistasis in Maize (Zea mays L.) III. Comparison of Single and Three-Way Crosses for Forage Traits

The importance of epistasis for forage performance and quality traits was studied in early maturing European maize (Zea mays L.) breeding materials. Sixty-six three-way crosses (3W), 33 of the flint x (dent · dent) and 33 of the dem × (flint · flint) type, were compared with the mean of their respective non-parental single crosses () using data obtained in six environments. For each of the 11 traits examined, at least 6 out of the 66 three-way crosses displayed significant (P < 0.05) epistatic deviations, calculated as – 3W. Averaged over crosses and environments, epistatic deviations were highly significant (P < 0,31] for 8 trails, but of minor importance except for grain yield. In general, the variance of epistatie deviations was appreciably smaller than the variance of genotype × environment interactions. Highly significant correlations occurred among epistatie deviations for silking date, dry matter content of ear, and dry matter yield of grain and stover. The reliability of predicting the performance of three-way crosses from 2W as opposed to assessing them directly was investigated as a function of the number of test environments. Disturbing e fleets of epistasis on prediction only became important under intense testing (> 3 or 4 environments), With fewer test environments, both procedures were almost equivalent, due to a smaller influence of genotype × environment interaction on the values, Epistasis is concluded to be of minor importance with regard to the optimum type of hybrid for forage use.     

Genetic Diversity for Adult Plant Resistance to Leaf Rust (Puccinia recondita) in Near-Isogenic Lines and in Indian Wheats

Variation for adult plant resistance in near-isogenic wheat lines carrying Lrl4b, Lrl4ab and Lr30 in a ‘Thatcher’ background indicated the possible presence of novel adult plant resistance genes effective against the Indian leaf rust population. Sixty-one wheats released for cultivation in India were grown in isolated nurseries. Each nursery was separately inoculated with one of four leaf rust pathotypes which had been selected to aid identification of resistance effective only in the adult plant stage. Seven distinct response groups were recognised and a minimum of six sources of adult plant resistance were postulated. In a group of 14 wheats, resistance was explained on the basis of the seedling response genes that were identified. Similar results for two years with pathotype 77-1 gave support to the reliability of field tests. Adult plant resistance (APR) sources were either race-specific or effective against all pathotypes used. Seedlings of cultivars with APR showed susceptible reactions. The possible presence of Lr34 in Indian wheats and its role in durable leaf rust resistance are discussed.     

三个超高产夏玉米品种的物质生产及光合特性

探明不同株高、穗位高具有超高产潜力(>15 000 kg hm^-2)夏玉米品种的物质生产及光合特性,有利于密植增产。本研究在多年高产试验基础上(连续3年达15 000 kg hm^-2以上),选用了登海661 (DH661,低秆低穗位)、登海701(DH701,中秆中穗位)和先玉335 (XY335,高秆高穗位) 3个具有超高产潜力的品种进行试验。在一般高产条件下,这3个品种间产量无显著差异,但干物质生产特性的模型解析表明,DH661的产量潜力高于DH701和XY335,其活跃生长期比XY335长近3周。开花后光合特征参数显示,DH661光合作用与光能利用效率明显较高,尤其是当XY335进入生理功能速衰期时(开花后28 d),DH661仍处于缓慢下降阶段,且具有较高的光合速率和光能利用效率,DH661的这一特性表明其具有更高的产量潜力。     

Interspecific hybridization of a multiploid mutant of durum wheat with rye and Triticum monococcum L. results in pentaploid hybrids

The multiploid mutant of durum wheat is a genotype that produces unreduced gametes. Our objective was to test the recovery of pentaploid hybrids in crosses of the mutant with rye and Triticum monococcum L. Compared with check crosses, the mutant had a two‐third reduction in percent seed set for rye crosses, but had only a slight decrease in crossability with T. monococcum. Pentaploid hybrids were associated with plump seeds of the mutant/rye cross, and with shrivelled seeds of the mutant/T. monococcum cross. We suggest that the endosperm balance number hypothesis explains the association of pentaploid hybrids with endosperm type. This association made for easy recovery of pentaploid hybrids from crosses to both species. Mature, plump seeds from the mutant/rye cross were germinated and pentaploid hybrids were recovered. One pentaploid hybrid was recovered for every 50.5 and 15.1 florets pollinated with rye and T. monococcum, respectively. Unreduced gametes in the multiploid mutant will facilitate interspecific hybridization by reducing the time to produce pentaploid plants.     

普通玉米3个籽粒品质性状的配合力分析

以10个普通玉米自交系及其90个杂交组合在北京和山西2个地点的试验数据,分析了蛋白质、淀粉和油分含量的配合力效应。结果表明：(1) 各性状的一般配合力(GCA和特殊配合力(SCA) 均方极显著,且GCA均方大于SCA均方,说明它们在遗传上主要受加性基因效应的影响。(2) 杂交种的3个品质性状与亲本GCA平均值和SCA极显著正相关,但与亲本GCA均值间的相关性相对较强,其相关系数(r)分别为0.708**(蛋白质)、0.756**(淀粉)和0.772**(油分),表明提高亲本GCA对改良杂交种品质具有更大的作用。(3) 自交系BT1和E28的蛋白质含量GCA较高,掖478、昌7-2、掖107和48-2的淀粉含量GCA较高,E28、黄C和P138的油分含量GCA较高。