Crossability of tetraploid and hexaploid wheats with ryes for primary triticale production

Summary Crossability of wheat and rye was investigated during thirteen crop cycles in two contrasting locations to 1) evaluate tetraploid and hexaploid wheat parents in crosses with rye, 2) identify genotypes with high crossability and 3) assess the impact of environment on seed development. The majority of the tetraploid wheats crossed with rye had seed set around 20%, but very low embryo viability. Several wheat genotypes with seed set above 50% were identified. The hexaploid wheats crossed with rye showed poor seed set, but plant recovery was relatively high. The majority of the hexaploid wheats with highest seed set (20–30%) were from China. The results suggest differences in crossability between the rye populations, and wheat species by rye interactions. The crossability of the tetraploid and hexaploid wheats was affected by climate in the two locations.     

Parental effects on crossability,embryo differentiation and plantlet recovery in wheat x rye hybrids

Summary One hexaploid wheat cultivar (Triticum aestivem) and two tetraploid wheat lines (T. durum) were crossed with seventeen inbred lines of rye (Secale cereale). Seed set, degree of hybrid embryo differentiation at the time of excision for in vitro culture and recovery of amphihaploid plantlets from various embryo categories were studied. Degree of embryo differentiation was predominantly determined by maternal wheats, paternal rye genotypes appearing to be of minor importance. T. aestivum x rye hybrid embryos were superior to those produced from T. durum for degree of differentiation. The proportion of plantlets developing from differentiated embryos was high for all wheat parents, whereas undifferentiated embryos were mostly unsuitable for plantlet production. The results revealed that cross-incompatibility in hexaploid wheat x rye crosses was due to failure of fertilization, while in tetraploid wheat x rye crosses it was caused by lack of embryo differentiation. Correlation analyses showed that seed set provided a criterion to predict the amphihaploid plantlets to be expected from a particular wheat x rye combination.     

Crossability and embryo rescue enhancement in wide crosses between wheat and three Agropyron species

Summary Soft winter wheat lines were crossed with Agropyron intermedium, A. elongatum and A. trichophorum using pollen from single plants of Agropyron spp. to pollinate wheat spikes. Not only species but also individual plants within varieties of Agropyron species differed in percent seed set with a wheat genotype. In two arrays of crosses between two phenotypically different plants of A. elongatum and nine wheat lines, one Agropyron plant gave higher seed set (overall=27.1%) than the other (overall=3.7%). The differences were significant in seven of the nine cross combinations. Results are consistent with the hypothesis that these two plants differ in their crossability as pollen parents with wheat, and suggest the possibility of occurrence of crossability genes in wheatgrasses. The success rate of hybrid embryo rescue was higher (87.5%) with cold treatment (4°C) than without cold treatment (75.0%) of excised embryos on culture media. Results underscore the significance of genotype of the alien species for crossing with low crossable wheats, and of the physical factors for improving embryo rescue in wide crosses.Contribution No. 11,825, Purdue Agric. Exp. Stn., West Lafayette, Indiana 47907, USA. The research was supported in part by Public Varieties of Indiana.     

Transfer of the recessive crossability allele kr1 from Chinese Spring into the winter wheat variety Martonvásári 9

Summary The recessive of crossability allele kr1 was transferred from the spring wheat variety Chinese Spring (CS) into the winter wheat variety Martonvásári 9 (Mv9) by backcrossing the Mv9 × CS hybrids with Mv9. The Mv9 variety possesses dominant Kr1 alleles and is heterogeneous at the kr2 locus, so that some individual plants carry recessive kr2 alleles. The selection of plants possessing the recessive kr alleles from the (Mv9 × CS)Mv9 BC₁ generation was carried out according to the seed set achieved when pollinated with rye (Secale cereale L. cv. Mercator). The partial dominance of the Kr alleles made it possible to differentiate between plants heterozygous at the Kr1 locus and Kr1Kr1 homozygous dominant plants. Two selfed consecutive progenies were tested by pollination with rye to select the homozygous recessive kr1kr1kr2kr2 plants and to check the result of the selection after each backcross.As a result of three backcrosses with Mv9 and two selfings after each backcross the selected progenies had 61.6% seed set with rye tested on sixty individual plants. These data confirm that after the third backcross the selected Mv9 kr1 line carries necessive crossability alleles Kr1 and Kr2, but the genotype is 93.75% Mv9.     

The site of action of the crossability genes (Kr 1, Kr 2) between Triticum and Secale. I. Pollen germination,pollen tube growth,and number of pollen tubes

Summary Seven genotypes of common wheat (Triticum aestivum L.) were crossed with rye (Secale cereale L.) in order to find the site or sites of action of the crossability genes, Kr ₁ and Kr ₂, of wheat. The data obtained, by fluorescence microscopy, were compared to the controls (wheat x wheat). The results indicate that the crossability genes have little effect on pollen germination and on the time taken for the pollen tubes to reach the microphyle, irrespective of their crossabilities with rye. The number of pollen tubes reaching the microphyle is, however, affected by the Kr-genes, as high crossable genotypes have more pollen tubes than the low crossable ones. There was a high correlation between the mean number of pollen tubes at the micropyle with seed set, which also reflects the crossability. The Kr-genes seem to manifest themselves in the retardation and inhibition of pollen tube growth between the style base and the top of the embryo sac, where the effect is most distinct in the low crossable genotypes.     

The site of action of crossability genes (Kr 1, Kr 2) between Triticum and Secale. II. Proportion of pistils containing pollen tubes and effects of alternate pollination on seed set

Summary Different wheat genotypes (T. aestivum) were crossed with rye to ascertain the site or sites of manifestation of the crossability genes, Kr ₁ and Kr ₂. By using fluorescence microscopy, it was found that the order of increasing proportion of wheat micropyles containing pollen tubes is strongly correlated with the levels of crossability with rye. High crossable genotypes have more micropyles containing pollen tubes than those of the low crossable ones. Most of the inhibition or retardation of pollen tubes occurred between the style base and top of the embryo sac, expecially with the low crossable genotypes where both Kr-genes are present. The results also indicate that Kr ₁ is a greater inhibitor than Kr ₂. Seed set is also highly correlated with the number or proportion of micropyles having pollen tubes. Alternate pollinations seem to support the view that rye pollen tubes do not reach the micropyles of the low crossable genotypes, and hence when repollinated with wheat selfed seeds are produced.     

Genetic control of crossability of triticale with rye

Limited genetic knowledge is available regarding crossability between hexaploid triticale (2n= 6x= 42, 21″, AABBRR, amphiploid Triticum turgidum L.‐Secale cereale L.) and rye (2n= 14, 7″, RR). Our objectives were to determine (1) the crossability between triticales and rye and (2) the inheritance of crossability between F₂ progeny from intertriticale crosses and rye. First, ‘8F/Corgo’, a hexaploid triticale, was crossed as a female with two landrace ryes, ‘Gimonde’ and, ‘Vila Pouca’ and two derived north European cultivars, ‘Pluto’ and ‘Breno’. These crosses produced 21.7, 20.9, 5.9, and 5.6%, seed‐set or crossability, respectively, showing that the landrace ryes produced higher seed‐set than the cultivars. Second, ‘Gimonde’ rye was crossed as a male with four triticales for 3 years. The control cross, ‘Chinese Spring’ wheat × rye, produced 80‐90% seed‐set. Of the four triticales, ‘Beagle’ produced 35.7‐56.8% seed‐set. The other three triticales produced less than 20% seed‐set, showing that the triticales differ in crossability with ‘Gimonde’ rye. Third, six FiS from intertriticale crosses (‘8F/Corgo’בBeagle’, ‘Beagle’בCachirulo’, ‘Lasko’בBeagle’, ‘8F/Corgo’בCachirulo’, ‘Lasko’בCachirulo’, ‘Lasko’ב8F/Corgo’) were crossed to ‘Gimonde’ rye. Results indicated that lower crossability trait was partially dominant in the two F₁S from crosses involving ‘Beagle’(high crossability) with‘8F/Corgo’ and ‘Cachirulo’(low crossability) and completely dominant in the ‘Beagle’בLasko’ cross, as it happens in wheat. Fourth, segregants in four F₂ populations (‘Lasko’בBeagle’, ‘8F/Corgo’בBeagle’, ‘Lasko’ב8F/Corgo’, and‘8F/Corgo’בCachirulo’) were crossed with rye. Segregation for crossability was observed, although distinct segregation classes were blurred by environmental and perhaps other factors, such as self‐incompatibility alleles in rye. Segregation patterns showed that ‘Beagle’, with high crossability to rye, carries either Kr1 or Kr2. The three triticales with low crossability with rye were most likely homozygous for Kr1 and Kr2. Therefore, it is likely that the Kr loci from A and B genomes acting in wheat also play a role in triticale × rye crosses.     

How wide can a wide cross be?

Summary Wide crosses in wheat are reviewed in relation to various factors that facilitate wide crossing to show that wide crosses can be as wide as one can make them. Included in this review is a particular reference to wheat-wheatgrass (Agropyron complex) crosses and an update on wide crosses of wheat with various genera of Agropyron complex. Hybrid seed set is too variable to predict whether a wide hybrid, where no seed was obtained in one attempt, will not be possible. High crossability genes seem to facilitate not only fertilization but also seed development, enabling embryo rescue. Variability for crossability occurs not only in wheat but also in alien species. Contrary to conventional thinking, several wide hybrids with wheat can be produced when species with lower chromosome numbers are used as female parents. Pre-and post-fertilization barriers to wide crosses do not appear to be equally strong, and can be overcome by the development and application of various technologies. Considering these aspects of wide hybridization, and based on recent successes in producing previously unsuccessful and very wide hybrids, it is concluded that how wide cross between plant species can be made is an open question and that many new and wider hybrids can be produced in future.     

The development of cytogenetic research at the Plant Breeding Institute Halle/Hohenthurm with special reference to aneuploidy in cereals

Summary The historical development of cytogenetic research in cereals performed at the Plant Breeding Institute of the Martin-Luther-University Halle-Wittenberg from its beginning in 1935 until 1992 is reviewed with special reference to polyploidy, alien introgression and aneuploidy.Th. Roemer founded 1935 in the framework of his Institute a Department of Mutation Research which, in 1937, was extended to a Department of Cytogenetics with R. Freisleben as the first head. Research highlights of this period were the introduction of mutation breeding, the development of autotetraploids in barley and linseed, the discovery of the crossability genes in wheat and the performance of wheat-rye crosses.The main objective in the period between 1950–1960 was the analysis of the relationships between chromosome behaviour and seed set in tetraploid rye and octoploid triticale.Since 1961 the Cytogenetics Research Group was headed by D. Mettin; he was followed by W.D. Blüthner in 1983. The research activities in this period concerning aneuploidy in rye and wheat and alien introgression are being reviewed under the following headings: Cytogenetics of rye; work with wheat aneuploids; contributions to the IR introgression into wheat; alien introgressions into wheat to improve disease resistance and grain quality; the exploitation of molecular markers.     

Effect of D-Genome Chromosome Substitutions on Hybrid Seed Development and Viability in T. turgidum var. durum X S. cereale Crosses

Triticum turgidum var. durum cv. ‘Langdon’ and the set of D-genome disomicsubstitutions in ‘Langdon’, produced at Fargo, U.S.A., were grown in a temperature controlled greenhouse and crossed with diploid spring rye (Secale cereals), to determine the effect of each substitution on 1. the crossability with rye, and 2, the viability of the resulting hybrids kernels. None of the disomicsubstitutions lines, with the possible exception of the 5D (5Bj line, gave an appreciable improvement in hybrid kernel set, -development, and -viability over the control, ‘Langdon’ The post-zygotic barrier to endosperm and embryo development, which operates in crosses between durum wheat and rye, could therefore not be suppressed by any specific chromosome of the D-genome.     

Crossability of some bread wheat landraces and improved cultivars from western Himalayas with rye

Summary Crossability of 62 bread wheat accessions (14 landraces from Himachal Pradesh and 48 others) was examined with rye. The 3 rye cultivars did not differ in their relative crossability with 4 of the wheat accessions studied. On the other hand, the wheat cultivars differed greatly among themselves in their crossability with rye. Most of the wheat cultivars showed poor (<10%) crossability. Two of the 14 landraces from Himachal Pradesh were found to be free from the crossability inhibitors as they showed very high (>50%) crossability, whereas none of the other 48 cultivars studied was so.     

The importance of wild potato species resistant to the potato cyst nematode,Globodera pallida,pathotypes P4A and P5A,in potato breeding. II. The crossability of resistant species

Summary Over 2400 pollinations were made to investigate the crossability relationships between cultivated potatoes, and wild diploid species from Series Tuberosa and Circaeifolia, as well as wild polyploid species in Series Tuberosa and Longipedicellata resistant to potato cyst nematode, Globodera pallida pathotypes P₄A and P₅A. Wild diploids in Series Tuberosa crossed easily with cultivated diploid species, except with Solanum lignicaule where most pollinations failed, and seed set was extremely low (0.2 seeds per pollination or less). It is suggested that this species is 1EBN. S. capsicibaccatum is clearly isolated from Series Tuberosa, but can form hybrids with S. lignicaule, which can act as a bridging species to S. tuberosum haploids. S. gourlayi and S. oplocense can be crossed with both subspecies of S. tuberosum, but S. papita, Series Longipedicellata is reproductively isolated from the tetraploid cultigens. The crossability data are discussed in the light of germplasm utilisation for breeding potato varieties resistant to potato cyst nematode.     

The production and behaviour of hexaploid hybrids between Hordeum vulgare and H. Bulbosum

Summary To produce hexaploid (or other polyploid) hybrids, diploid or tetraploid Hordeum vulgare was crossed with hexaploid or octoploid H. bulbosum, and perennial triploid hybrids between the two species were treated with colchicine. The crosses did not yield viable plants: seedset was low, the seed aborted and embryo culture was unsuccessful. The colchicine treatments geve rise to plants in which hexaploid chromosome numbers were observed. At the hexaploid level chromosomal instability occurred, resulting in chromosome elimination.The colchicine-treated triploid hybrids showed in the first years after the treatment better fertility after open flowering than untreated plants, but the level of fertility remained very low. The offspring consisted of haploid, diploid and approximately triploid plants like H. vulgare, tetraploid and approximately tetraploid plants like H. bulbosum, and plants with hybrid morphology and unstable chromosome number, which were highly sterile. Thus the crossing barrier between H. vulgare and H. bulbosum could not be broken down at higher ploidy level.     

Crossability percentages of bread wheat collections from Tibet,China with rye

Summary The crossability percentages of 282 accessions of wheat (Triticum aestivum L.) collected in Tibet, China with rye (Secale cereale L.) have been tested. Five collections have a similar to and 277 accessions have a lower crossability percentage than Chinese Spring or are non-crossable with rye. The accessions with high crossability percentage occur along the highway near Lhasa. No landraces with higher crossability than Chinese Spring and rare landraces with similar crossability to Chinese Spring indicated that the landraces in Tibet region are different from those in Sichuan, Shaanxi and Henan provinces in the distribution frequency of high crossability, and there is no distribution of recessive kr4 alleles.     

Use of Vaccinium octoploids to facilitate 4x-6x gene transfers

The use of an 8x Vaccinium produced through doubling a tetraploid hybrid with colchicine was studied as a means of facilitating gene exchange between V. corymbosum L. and V. ashei Reade. Analysis included meiotic observations and crossability studies of the 8x plant, as well as attempting to develop 6x breeding lines. Meiotic analysis revealed the presence of micronuclei in Telphase II products and polyspory in sporads. In the crossability studies five 4x highbush plants were pollinated with 8x o2 4x pollen. Data gathered included: % fruit set, average weight/fruit, average number of seeds/fruit, average number of seedlings/pollination, and number of 6x seedlings. Octoploid pollinations were significantly lower than 4x pollinations in all parameters. One 6x plant was produced from the 4x-8x crosses but was found to be mitotically unstable, having somatic cells with chromosome numbers ranging from 48 to 168.     

The introduction into bread wheat of a major gene for resistance to powdery mildew from wild emmer wheat

Summary A new source of resistance to wheat powdery mildew caused by Erysiphe graminis has been transferred to hexaploid bread wheat, Triticum aestivum, from the wild tetraploid wheat, Triticum dicoccoides. The donor was crossed to bread wheat and the pentaploid progeny was then self-pollinated. Plants having a near stable hexaploid chromosome complement were selected in the F₃ progeny and topcrossing and backcrossing of these to a second wheat cultivar to improve the phenotype was undertaken. Monosomic analysis of early backcross lines showed the transferred gene to be located on chromosome 4A. The gene has been designated Pm16.     

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.     

Crossability percentages of bread wheat landraces from Shaanxi and Henan provinces,China with rye

Summary The crossability percentages of 118 landraces of wheat (Triticum aestivum L.) from Shaanxi and Henan provinces, China with rye (Secale cereale L.) have been tested. 14 landraces possess a higher crossability percentage than Chinese Spring, 30 landraces have a similar to and 74 landraces have a lower crossability percentage than Chinese Spring or are non-crossable with rye. Most landraces with high crossability percentage occur in south part of Shaanxi, and the west and southwest of Henan, their geographical distribution in these regions fits up with that in the Sichuan province.     

Crossability percentages of bread wheat landraces from Sichuan Province,China with rye

Summary The crossabilities of 177 landraces of wheat (Triticum aestivum L.) from Sichuan Basin and its adjacent mountain ranges with rye (Secale cereale L.) have been tested. 16 landraces possess a higher crossability than Chinese Spring, 34 landraces have a similar and 127 landraces have a lower crossability than Chinese Spring or are non-crossable with rye. Most landraces with high crossability occur in Qinling Mountain and Dabashan Mountain Ranges in north of Sichuan and the valleys of Minjiang River, Fujiang River and Jialinjiang River in Sichuan Basin.     

Relation between wheat-rye crossability and seed set of common wheat after pollination with other species in the Hordeae

Summary Cultivars of common wheat (Triticum aestivum L. em. Thell.) of high wheat-rye (Secale cereale L.) crossability set more seed with pollen of other related species than did wheats of low wheat-rye crossability. This was found to be true for pollen parents from the genera Triticum, Aegilops, Secale, Agropyron and Elymus.