The genotypic and phenotypic interaction of wheat and rye storage proteins in primary triticale

The creation of new triticale germplasm from its wheat and rye progenitors is an important source of potentially beneficial alleles. This research focused on the inheritance of glutenin and secalin alleles in triticale and their interaction in the hybrid offspring in terms of both subunit expression and gluten strength. Multiple crosses between five durum lines and two inbred rye lines were performed to create a set of 23 fertile amphidiploid lines. The SDS-PAGE banding patterns of the denatured HMW glutenins, LMW glutenins, HMW secalins and 75k γ-secalins (termed secaloglutenin) were compared to their exact parental plants, and their SDS-sedimentation was evaluated. Secaloglutenin subunits were simply inherited and expressed in all primary triticales. The few observed variations in banding patterns were accounted for within measured heterogeneity of the exact rye parental plants, for which the secalin alleles were thoroughly described. However, the possibility remains that mixed protein oligomers were formed between polymeric and monomeric storage proteins at the ultrastructure level. Furthermore, significant differences in the gluten strength of primaries derived from different durum parents were observed. This research suggests triticale of high gluten strength can be obtained by selection of parents with favourable glutenin alleles.     

Heterosis in Primary Hexaploid Triticale with Heterozygous Wheat or Rye Genome*)

Twelve primary hexaploid triticale (X Triticosecale Wittmack), synthesized from, three lines of tetraploid wheat (Triticum durum L., T. turgidum L.) and four inbred lines of rye (Secale cereale L.), were used to produce 18 crosses with homozygous wheat and heterozygous rye genome and 12 crosses with heterozygous wheat and homozygous rye genome. Parents and crosses of triticale, wheat, and rye were tested for two years (rye for one year only) in two-replicate block designs with 1 m²-plots. Data were assessed for plant height, grain yield and for yield-related traits. Performance of triticale crosses was considerably lower than that of the wheat and rye crosses. The amount of heterosis varied greatly between years. Positive and mainly significant heterosis was revealed in triticale generations F₁ and F₂. The average values were closer to those in wheat than to those in rye. For most characters a high level of heterosis was retained in tnucalt1 generation F₂. Heterozygosity of the wheat and rye genome both contributed to heterosis in triticale. However, gene action of the rye genome strongly depended on the homozygous wheat background: one wheat line almost completely suppressed and another greatly stimulated the heterotic effect of the rye genome. In the later case, the amount of heterosis was related to that in rye per se. Information from hybrid rye breeding may therefore be used when establishing gene pools for hybrid breeding in triticale.     

Pollen production in hexaploid triticale

Summary Comparative studies were made of the pollen characteristics of triticale, wheat, and rye. Measurements were made of the anther length, width, and percent extrusion; pollen viability; size and number of pollen grains per anther; and dispersal on 10-mm² slide area pollen traps. Triticale anthers were intermediate in length between and significantly different from both wheat and rye. Rye pollen grains per anther were four and two times greater in number than those of wheat and triticale, respectively. Pollen viability was not significantly different between species. Rye pollen grains were smaller than those of wheat and of some triticale cultivars. Simple correlations between anther length and anther width, pollen grains per anther, pollen grain trapped per 10 mm² slide area, and plant height were significantly positive.     

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.     

普通小麦 小黑麦和黑麦花粉粒及其叶表皮结构的扫描电镜观察

普通小麦、小黑麦和丹麦黑麦在花粉粒形状。花粉粒表面纹饰和叶表皮结构等方面存在显著差异。小黑麦的上述特征介于普通小麦和丹麦黑麦之间。因此,花粉粒形状及其表面纹饰特征和叶表皮结构特征可能适于作为种属分类的重要依据之一,对新物种的鉴定和确定物种间的亲缘关系也有参考意义。     

Screening for resistance to take-all in wheat,triticale and wheat-triticale hybrid lines

Summary Fifteen triticale and wheat-triticale hybrid lines were evaluated for resistance to the take-all fungus Gaeumannomyces graminis var. tritici and compared with five wheat and two rye lines in inoculated field and pot trials. The triticale and wheat-triticale hybrid lines varied in rye chromosome number and degree of resistance expressed. One line, Venus with seven pairs of rye chromosomes consistently showed levels of resistance intermediate between wheat and rye. A trend was observed where increasing rye chromosome content led to greater resistance but exceptions showed that variation within triticales could not be ascribed to rye chromosome content alone.     

A New Method to Produce 4× Triticales and their Application in Studying the Development of a New Polyploid Plant

F₁ hybrids of triticale × rye derived from commercial varieties were backcrossed to the respective triticale parent. Selfing of the backcross generation yielded a large number of 4× triticales containing a genetically balanced wheat genome. This indicates that the 28-chromosome F₁ plants with the genomic constitution of ABRR produced functional 14-chromosome gametes in high frequency each with a complete wheat and rye genome. The cytological mechanism leading to the formation of tetraploid triticales is described. The chromosomal constitution of the wheat genome in the progenies of 30 back cross plants was analysed by the C-banding technique. One offspring possessed a complete B genome of wheat. The production of tetraploid triticale through backcrossing in comparison to selfing the ABRR hybrid is largely independent of the genotype; it leads to new tetraploids in just three generations and it reduces the chance of translocations between the homoeologous wheat chromosomes. The possibility of studying the effect of different mixtures of chromosomes of the A and B genomes of wheat on the phenotype of the tetraploid triticale is discussed.     

Attempts at Interspecific Hybridization Between Phaseolus vulgaris L. and P. acutifolius A. Gray-Using Embryo Rescue

In diploid rye, two genes were detected which cause hybrid necrosis by complementary action if both are present in dominant condition. Moreover, these genes cause hybrid necrosis in triticale complementing with cither one of the two genes, Ne1 and Ne2 which are known to cause hybrid necrosis in wheat. It is suggested, that the two genes in rye are named Ner1 and Ner2 corresponding to the wheat gene with which they complement in triticale. The consequences of the presence of necrosis genes in rye populations for breeding of rye are discussed.     

Histological studies on the infection of triticale,wheat and rye byPuccinia recondita f.sp.tritici andP. recondita f.sp.recondita

Summary The reaction of eight triticales and of the respective wheat and rye parental lines to infection by the leaf rust fungi of wheat and rye were studied in the seedling stage. The histological observations indicated that wheat and triticale showed a typical nonhost reaction to the leaf rust of rye: sporelings of this fungus were arrested after the formation of primary infection hyphae and before the formation of extensively branched mycelium, mostly without necrosis of plant cells. The rye inbred lines were all susceptible to the rye leaf rust. The reaction of wheat and triticales to the wheat leaf rust was susceptible or resistant. The reaction of resistant lines could be early or late and complete or incomplete, but was associated with substantial necrosis of plant cells, and therefore entirely different from the nonhost reaction to rye leaf rust. In their reaction to wheat leaf rust the rye lines were similar to the resistant wheat and triticale lines. They did not show an important degree of nonhypersensitive early abortion as would be expected in a nonhost species. It appeared that genes for hypersensitive resistance in triticale may be contributed by either the wheat or the rye parental line.A screening of sixty wheat, rye and triticale lines confirmed the nonhost status of wheat and triticale to rye leaf rust and the hypersensitive or moderately susceptible reaction of rye to wheat leaf rust.     

Triticale,Still a Promise?

Experiments with primary triticale genotypes produced from defined wheat and rye parents provide a strong indication of a preponderance of specific genome combining ability in the phenotypic expression of triticale. Studying line families developed from specific cross combinations by means of the single seed descent method revealed that, in contrast to wheat, the distribution of genotypes in an F₂ population of triticale is strongly skewed, with the majority of them being inferior to the mid-parent value. This is the case even if recombination is restricted to one fourth of one of the parental components of triticale, i.e. wheat or rye. Correlations of yield components were found to deviate substantially from known types of correlations in wheat, an established natural allopolyploid species. These findings give a strong indication for a genomic inbalance in triticale being a major cause for slow advance in varietal improvement. New breeding strategies designed for the specific genetic structure of triticale are necessary to accelerate breeding progress and to help triticale ultimately to live up to its promise.     

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.     

Hardening of Some Winter Wheat (Triticum aestivum L.), Rye (Secale cereals L.), Triticale (×Triticosecale Wittmack) and Winter Barley (Hordeum vulgare L.) Cultivars During Autumn and the Final Winter Survival in Finland

Profound hardening is the prerequisite for survival of a cultivar over winter. In the present study, hardening abilities of 13 winter wheat, 10 rye, 3 triticale and 6 winter barley cultivars were determined in field samples collected during the autumns of 1990 and 1991 by measuring the amount of ion leakage from frost-treated leaf segments. The cultivars tested were selected from the Inter-Nordic Winterhardiness Project, in which Field Survival Indices were determined for a total of 23 winter wheat, 13 rye, 5 triticale and 11 winter barley cultivars in order to estimate winter survival potentials under Finnish conditions. The winter survival of the materials studied correlated well with their hardening ability assessed by using the electrolyte leakage method.     

Fluorescent in situ hybridization as an aid to introducing alien genetic variation into wheat

Summary Fluorescent in situ hybridization (FISH) has been used to assess the occurrence and frequency of wheat-alien chromosome pairing in a wheat/Thinopyrum bessarabicum hybrid and in wheat/rye hybrids with different levels of chromosome pairing by examining pollen mother cells at metaphase I of meiosis. The use of FISH to identify the presence and size of alien chromatin in a wheat background is also demonstrated.The value of FISH as an aid to the introgression of alien genetic variation into wheat is discussed.Abbreviations FISH fluorescent in situ hybridization - GISH genomic in situ hybridization - PRINS primer-induced in situ hybridization     

Dosage Response of Rye Genes in a Wheat Background

Wheat (Triticum aestivum L.) breeders often utilize alien sources to supply new genetic variation to their breeding programs. However, the alien gene complexes have not always behaved as desired when placed into a wheat background. The introgressed genes of interest may be linked to undesirable genes, expressed at low levels or not at all. The short arm of rye (Secale cereale L.) chromosome one (1RS) contains many valuable genes for wheat improvement. In order to study rye gene response to varying copy number, wheat lines were constructed which contained zero, two or four doses of 1RS. The meiotic behavior of rye chromosome 1R, and wheat/rye translocation chromosomes, 1AL/1RS and 1BL/1RS was studied in the F₁ hybrids between wheat lines carrying 1R or the translocation chromosomes. The IRS arm was transmitted at a very high frequency; 98 % of the F₂ plants had at least one of the chromosomes with a IRS arm. In addition, 44 % of the F₂ plants received at least one copy of the chromosomes from each parent. Analysis of the meiotic behavior of the IRS arm suggested that few euploid wheat gametes were formed. Therefore, most of the pollen must have contained IRS. It is unknown whether the lack of euploid wheat pollen could account for the high transmission frequency of the rye chromosomes. There may have been differential survival of the embryos receiving the rye chromosome as well.     

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.     

Morphological and physiological differences in the response of cereals to zinc deficiency

Greenhouse and growth chamber experiments were carried out using seven bread wheat (Triticum aestivum), three durum wheat (T. durum), two rye (Secale cereale), three barley (Hordeum vulgare), two triticale (x Triticosecale Wittmack) and one oat (Avena sativa) cultivars to study response to zinc (Zn) deficiency and Zn fertilisation in nutrient solution and in a severely Zn deficient calcareous soil. Visual Zn deficiency symptoms, such as whitish-brown necrotic patches on leaf blades, developed rapidly and severely in the durum wheat and oat cultivars. Bread wheat showed great genotypic differences in sensitivity to Zn deficiency. In triticale and rye, visual deficiency symptoms were either absent or appeared only slightly, while barley showed a moderate sensitivity. When grown in soil, average decreases in shoot dry matter production due to Zn deficiency were 15% for rye, 25% for triticale, 34% for barley, 42% for bread wheat, 63% for oat and 65% for durum wheat. Differential Zn efficiency among and within cereal species was better related to the total amount of Zn per shoot, but not to the Zn concentration in the shoot dry matter. However, in leaves of Zn efficient rye and bread wheat cultivars, the activity of Zn-containing superoxide dismutase was greater than in Zn inefficient bread and durum wheat cultivars, suggesting higher amounts of physiologically active Zn in leaf tissue of efficient genotypes. When grown in nutrient solution, there was a poor relationship between Zn efficiency and release rate of Zn-chelating phytosiderophores from roots, but uptake of labelled Zn (⁶⁵Zn) and its translocation to the shoot was higher in the Zn efficient rye and bread wheat cultivars than in inefficient bread and durum wheat cultivars. The results demonstrate that susceptibility of cereals to Zn deficiency decline in the order durum wheat > oat > bread wheat > barley > triticale > rye. The results also show that expression of high Zn efficiency in cereals was causally related to enhanced capability of genotypes to take up Zn from soils and use it efficiently in tissues. This revised version was published online in August 2006 with corrections to the Cover Date.     

Influence of selected wheat and rye genotypes on the direct synthesis of hexaploid triticale

Summary In the synthesis of primary hexaploid triticale, a cross-incompatibility barrier exists when tetraploid wheat (4X) is crossed with diploid (2X) rye. Fertilization may occur, however, abnormal endosperm development usually leads to premature embryo death. Four selected tetraploid wheat lines were crossed as females with seven open-pollinated rye lines and the resulting embryos were rescued in vitro 13–16 days after pollination. The wheat genotypes showed a major influence on crossability (seed set), embryo development and plant recovery. The highest efficiency of amphihaploid plant recovery (18.3 plants per 100 pollinated florets) was obtained from one 4X wheat line originally selected from the cross T. carthlicum × T. dicoccoides. Some of the 3X amphihaploid plants (ABR) derived from two wheat lines showed relatively high level of partial fertility presumably as a result of meiotic restitution. Correlation analysis showed that crossability (seed set), normal hybrid embryo development in vivo and embryo culturability were independent of each other.     

The Identification of a Gibberellic-Acid-Insensitive Gene in Secale cereale*

Among 16 dwarfing genes identified in wheat (Triticum aestiuvm L. em Thell.), four are known to be associated with insensitivity to the externally-supplied growth hormone gibberellin (GA). Rht1 and Rht2 (Reduced height 1 and 2, respectively) have been the most extensively used, because of their positive effect on yield. To increase the germplasm pool for dwarfism, a spring rye (Secale cereale) population (UC-90, CI-174) was selected because it contains high variability and any useful genes would benefit triticale and wheat as well. Seedlings of the CI-174 rye population were treated with 50 ppm of GA to identify any insensitive types. GA-insensitive and -sensitive seedlings were identified and, after three generations of selfing, GA-insensitive and -sensitive lines were fixed. Rye insensitive was crossed to a sensitive wheat and to rye and, reciprocally, insensitive wheat was crossed to sensitive rye. The results indicated that a GA-insensitivity dwarfism system similar to that originally found in wheat also operates in rye and appears to be under simple inheritance. Rye GA-insensitivity was expressed in triticale. Therefore, it is possible to transfer this new source of insensitivity and dwarfism into triticale and wheat.     

Production of polyhaploids of hexaploid wheat using stored pearl millet pollen

The effects of drying and freezing on viability of pearl millet pollen were examined with the aim of using stored pollen in polyhaploid production of hexaploid wheat. Freshly collected pollen of pearl millet line NEC 7006 with 55% water content, germinated at a frequency of 80%. Pollen that was dried for two hours to 6% water content showed 50% germination frequency and maintained similar frequencies after the freezing process. In crosses of hexaploid wheat variety Norin 61 with fresh pearl millet pollen, embryos were obtained at a frequency of 27.6%. In crosses with pollen stored at -196 °C, -80 °C and -20 °C for one month, embryo formation frequencies ranged from 27.5 to 17.4%. After five and twelve months of storage, the frequencies ranged from 29.7 to 14.6% at storage temperatures of -196 °C and -80 °C, and from 8.0 to 3.2% at -20 °C, indicating significant differences among storage temperatures. However, no significant frequency difference was found among pollen water contents at the time of collection. All plants regenerated from crosses with pearl millet pollen stored for five months were wheat polyhaploids. These results suggest that stored pearl millet pollen is an efficient medium for producing polyhaploids in hexaploid wheat. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.