Wheat Substitutions in Hexaploid Triticale*

Crosses between octoploid and hexaploid triticales have been made m breeding programs for several years, From an analysis of the progeny of such crosses where selections for an octoploid-like phenotype had been made, it was established that 149 out of 150 lines were hexaploid in chromosome number, C-banding and in situ hybridization demonstrated that all but five of the 62 lines analyzed in detail contained visible chromosomes or segments from the D genome. Only four lines had D-genome chromosome replacing rye chromosomes. All of the remaining 53 D-genome substitutions involved the replacing of wheat chromosomes from either the A or B genomes. This establishes the ease with which D-genome genes can be placed into triticale without the loss of rye chromosomes.     

Rapid Incorporation of D-genome Chromosomes into A and/or B Genomes of Hexaploid Triticale

Two series of progenies were developed from hybrids between octoploid (AABBDDRR) and tetraploid triticale ((AB)(AB)RR). One arose from the successive selfing of the F₁s, while the second was established after one backcross of the F₁ hybrids with the respective 8 × triticale parent. Altogether, 250 F₃ and BC₁F₂ lines were developed, of which 112 were karyotyped in the F₄/F₅ or in BC₁F₃/BC₁F₄ generations using C-banding and SDS-PAGE. The 112 lines represented 61 different karyotypes, of which 39 appeared to be stabilized, having pairs of homologous wheat chromosomes only, while 22 karyotypes exhibited 1—3 heterologous pairs. The frequency of karyotypically stabilized lines originating from the series with one backcrossing was much higher (79.5 %) than those derived from the successive selfing of the F₁ hybrids (51.7%). Six lines had the pure hexaploid triticale chromosome constitution. The frequency of disomic substitutions of D genome chromosomes for their homoeologous A and/or B genome chromosomes ranged from one to six per line with an average of 1.7. Except for 3B and 6B all possible D(A/B) substitutions were obtained. Chromosomes ID and 3D substituted for their homoeologues with the highest frequency, while the substitution of chromosome 4D for 4A or for 4B was the least frequent. D(R) substitutions were found in eight lines only. A complete set of 6x triticale lines was established in which chromosome ID was present in all possible combinations, i.e. single 1D(1A/1B/1R) disomic substitutions as well as disomic ID addition.     

Transfer of the Glu-D1 Gene from Chromosome 1D to Chromosome 1A in Hexaploid Triticale

To complement previously developed recombinant chromosomes 1R.1D, two series of translocations involving the Glu-D1 gene from chromosome ID to chromosome 1A were produced in hexaploid triticale. These series involve seven independent transfers of allele d encoding for high molecular weight glutenin subunits 5+10 and ten independent transfers involving allele a encoding for HMW glutenin subunits 2 + 12. The frequency of homoeologous recombination between chromosomes 1A and 1D was within the range observed for pairs of homologues in wheat, supporting earlier observations that homoeologous recombination in triticale is frequent. Recombined chromosomes 1A.1D can be used to introduce the Glu-D1 gene to durum wheats, and to manipulate the dosage of Glu-D1 in hexaploid triticale and bread wheat.     

Nitrogen and phosphorus uptake and utilization efficiency in D(R) substitution lines of hexaploid triticale

Two sets of disomic substitution lines, derived from the cultivars ‘Presto’ and ‘Rhino’ of triticale, with rye chromosome pairs replaced by their wheat D‐genome homoeologues, were tested in hydroponic culture for nitrogen and phosphorus uptake and utilization efficiency. The effect of a substitution on the amount of absorbed nutrients was predominantly negative and proportional to the effect on plant dry matter. Significant decreases were found for the substitutions 5D(5R), 6D(6R) of both cultivars, 2D(2R), 4D(4R) of ‘Presto’ and 3D(3R) of ‘Rhino’. On the other hand, the nitrogen utilization efficiency was significantly increased in all substitution lines, with the exception of the 1D(1R) ones. The differences in phosphorus utilization were generally positive, but less pronounced, and significant only in the lines 2D(2R) and 6D(6R). The data suggest that presence of both rye and D‐genome chromosomes is conducive for the best effect of the applied N and P fertilizers.     

Characterization of wheat-triticale doubled haploid lines by cytological and biochemical markers

Five wheat-triticale doubled haploid (DH) lines— M08, V209, DH220-14-2, DH696-3-4 and M16 —derived from anther culture of F₁s resulting from crosses involving hexaploid or octoploid triticale × hexaploid wheat, were characterized by cytological and biochemical markers. Cytological evidence from genomic in situ hybridization and C-banding indicated that DH lines M08 and V209 (2n= 42) each contained a pair of 1BL/1RS translocation chromosomes. DH220-14-2 (2n= 42) was also a translocated line with two pairs of chromosomes containing small fragments of rye. One of the translocation fragments carried the Sec-1R gene originating from the satellite region of 1RS; the origin of the other one remains unknown. DH696-3-4 (2n= 42) contained a 3D(3R) substitution. In M16 (2n= 44), three pairs of rye chromosomes, 3R, 4R and 6R, were present, 4R as an addition and 3D(3R) and 6D(6R) as substitutions. Biochemical, isozyme and storage protein markers confirmed the cytological conclusions. The advantages of transferring alien chromosomes or chromosome fragments into wheat and creating alien aneuploid lines by anther culture of hybrid F¹s are discussed.     

Grain quality and yield characteristics of D-genome disomic substitution lines in 'Langdon' (Triticum turgidum var. durum)

Sets of D-genome disomic substitution lines of ‘Langdon’ (Triticum turgidum var. durum) were used to study the effect of chromosome substitutions on grain yield and flour technological properties. In general, the substitution of any D-genome chromosome had a detrimental effect on grain yield and growth vigour (some lines were sterile). SDS-sedimentation, SE-HPLC and two-gram mixograph procedures were used to measure dough strength of the lines studied. Significant correlations were observed between protein concentration and grain yield and other quality parameters such as SDS-sedimentation value, the proportion of glutenin, dough mix time and peak resistance. Most of the quality characters were highly correlated with each other. Substitution of chromosomes 1D, 5D, 2D and 7D resulted in positive responses to SDS-sedimentation values, but only chromosome 1D had positive effects on the proportion of peak 1 (P₁%), measured by SE-HPLC. Besides the major influence of chromosome 1D on three major mixograph parameters (mixing time, peak resistance and resistance breakdown), chromosome 5D also exhibited significant effects on these mixing parameters. Principal-component analysis showed that the predominant effect on durum-wheat rheological properties was from chromosome 1D, whereas chromosome 5D had a major effect on grain hardness (50%) and increased the whiteness of the flour.     

Chromosomal location of resistance to Septoria tritici in seedlings of a synthetic hexaploid wheat, Triticum spelta and two cultivars of Triticum aestivum

Chromosomal location of resistance to two virulent Argentinean isolatesof Septoria tritici was studied in two wheat (Triticum aestivumL.) cultivars (Cappelle-Desprez & Cheyenne), a synthetic hexaploid(Synthetic 6x) and Triticum spelta in seedlings. Substitution lines of these(resistant or moderately resistant) genotypes into (susceptible) ChineseSpring were selected from a previous screening. For Synthetic 6x,resistance was clearly located in chromosome 7D. Chinese Spring with the7D chromosome substituted by Synthetic 6x showed almost completeresistance, similar to the level of Synthetic 6x. For the substitutions withCappelle-Desprez, Cheyenne, and T.spelta there were no lines with abehaviour similar to the resistant parent. However, some substitutions weremore resistant than the susceptible parent suggesting that severalchromosomes could be involved in the resistance of these genotypes toSeptoria leaf blotch.     

Effects of D-genome chromosomes on crossability of hexaploid triticale (X Triticosecale Wittmack) with maize

With the aim of producing polyhaploids of hexaploid triticale, 20 genotypes from a CIMMYT breeding programme and eight D-genome chromosome substitution lines of ‘Rhino’ were crossed with maize. In crosses between 20 triticale genotypes and maize, 15 lines produced embryos. Frequencies of embryo formation ranged from 0.0 to 5.4%, with an average of 1.1%. From a total of 200 pollinated spikes, 62 plants were regenerated. Most regenerated plants were polyhaploids with 21 chromosomes, and few aneuhaploids with 22 chromosomes were found. In crosses of triticale substitution lines with maize, all the lines produced embryos, while ‘Rhino’ produced no embryos at all. Higher frequencies of embryo formation were obtained in substitution lines with chromosomes 2D and 4D. These results suggest that D-genome chromosomes in a triticale genetic background have the effect of increasing the frequency of polyhaploid production in triticale x maize crosses.     

Effect of alien 5R(5A) chromosome substitution on ear-emergence time and winter hardiness in wheat-rye substitution lines

Ear emergence time and response to vernalization were investigated in 12 alien substitution lines in which a pair of chromosomes 5A of recipient spring wheat cultivars was replaced by a pair of chromosomes 5R of Siberian spring rye ‘Onokhoiskaya’. The recipients were 12 spring cultivars of common wheat, each carrying different Vrn genes. Spring rye ‘Onokhoiskaya’ had the Sp1 (now called Vrn-R1) gene for spring growth habit located on chromosome 5R, but its expression was weaker. The Vrn-R1 gene had no effect on growth habit, ear emergence time and response to vernalization in wheat-rye substitution lines. Ears emerged significantly later in the 5R(5A) alien substitution lines than in the recipient wheat cultivars with the Vrn-A1/Vrn-B1/vrn-D1 or Vrn-A1/vrn-B1/Vrn-D1 genotypes. No difference in ear emergence time was found between most of the 5R(5A) alien substitution lines and the cultivars carrying the recessive vrn-A1 gene. The presence of the Vrn2a and Vrn2b alleles at the Vrn2 (now called Vrn-B1) locus located on wheat chromosome 5B was confirmed.The replacement of chromosome 5A by chromosome 5R in wheat cultivars ‘Rang’ and ‘Mironovskaya Krupnozernaya’, which carries the single dominant gene Vrn-A1, converted them to winter growth habit. In field studies near Novosibirsk the winter hardiness of 5R(5A) wheat–rye substitution lines of ‘Rang’ and ‘Mironovskaya Krupnozernaya’ was increased by 20–47% and 27–34%, respectively, over the recurrent parents.     

Identification of wheat chromosomes involved with different types of resistance against greenbug (Schizaphis graminum, Rond.) and the Russian wheat aphid (Diuraphis noxia, Mordvilko)

Two sets of intervarietal chromosome substitution lines in the recipient,susceptible cultivar ‘Chinese Spring’ were screened to identify the wheat chromosomes involved with antixenosis, antibiosis and tolerance resistance to greenbug and Russian wheat aphid. The amphiploid ‘Synthetic’ and the cultivar ‘Hope’ were the donor parents. Antixenosis, antibiosis and tolerance were evaluated with conventional tests in controlled environmental conditions using a clone of greenbug biotype C and a clone of RWA collected on wheat. Antixenosis against greenbug was accounted for by several chromosomes in both sets of substitution lines with chromosome 2B contributing the highest level of this type of resistance. The highest levels of antixenosis against RWA were associated with the group of chromosomes 7 of the substitutions CS/Syn set and the chromosome substitutions 2B, 6A and 7D of the CS/Hope set. Antibiosis against both aphids species was accounted for by several different chromosomes. The highest levels of antibiosis for most of RWA resistance traits were recorded from the 1B substitution line of the CS/Hope set. More than one gene appears to determine antibiosis. Tolerance to both greenbug and the RWA was significantly associated with chromosomes 1A,1D, and 6D in the CS/Syn set of substitutions. These lines showed enhanced plant growth under aphid infestation. The highest levels of antixenosis, antibiosis and tolerance against the two aphid species occurred mostly in different substitution lines. Consequently, the different types of resistance for both pests seem to be partially independent. Since different genes seem to be involved in at least several traits of the resistance categories against the two aphid species, such genes could be combined in new cultivars of wheat to broaden their genetic base of resistance against the greenbug and the RWA. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spontaneous wheat/rye translocations from female meiotic products of hybrids between octoploid triticale and wheat

Summary Heptaploid hybrids between octoploid triticale and wheat were backcrossed as female parents with wheat to examine the rye chromosome distribution in the resultant progenies using genomic in situ hybridization (GISH). One hundred and one backcross (BC) seeds were examined and whole rye chromosome additions and substitutions, wheat/rye centric and noncentric translocations and rye telocentric chromosomes were detected. Dicentric wheat/rye translocated chromosomes were also observed. Comparisons were made with previous results on the rye chromosome distribution from male gametes of the same cross and differences were found, where in the female derived population a deficit of plants with more than two rye chromosomes was apparent relative to the anther derived population.     

Transfer of the Glu-D1 Gene from Chromosome 1D of Breadwheat to Chromosome 1R in Hexaploid Triticale

The use of hexaploid triticale as a crop for human consumption has been limited by its inferior bread-making quality. To ameliorate this problem, a segment of chromosome ID of breadwheat with the Glu-D1d allele encoding for high molecular weight glutenin subunits 5 7plus; 10 was translocated to chromosome 1R of the hexaploid triticale ‘Rhino’ through a combination of a centric break-fusion translocation followed by 5D(5B)-induced homoeologous pairing. The resulting recombinant chromosome 1R has a small interstitial segment of ID with the Glu-D1d allele. The maximum physical length of the translocated segment is estimated at about 16.5 % of 1DL. Frequency of translocations involving the long arms of homoeologous group-1 chromosomes in the analyzed progeny suggested that homoeologous recombination in triticale was substantially higher than that previously reported in hexaploid wheat.     

Detection of a 4DL chromosome segment translocated to rye chromosome 5R in an advanced hexaploid triticale line Bronco 90

Bronco 90 is an advanced line of hexaploid triticale and was reported to be a 2D(2R) chromosome substitution type. In F₁ hybrids of this triticale with bread wheat, however, a meiotic configuration of 16 bivalents and 10 univalents was frequently observed indicating the presence of an additional D(R) chromosome substitution or D/R translocation. To determine the chromosome constitution of Bronco 90, C-banding and fluorescent in situ hybridization techniques were applied to somatic and meiotic metaphase chromosomes. These analyses revealed that in Bronco 90, the terminal 7% of the long arm of rye chromosome 5R is derived from the long arm of chromosome 4D. This translocated chromosome (5RS.5RL-4DL) and telosome 4DL formed metaphase I bonds at a frequency of 71%, demonstrating the significance of small terminal chromosome segments for pairing. This novel rye-wheat translocation is probably generated by homoeologous crossing-over because the distal region of 5RL is known to be homoeologous to that of 4DL. Possible association of this translocation with the absence of hairy peduncle character in Bronco 90 is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns of meiosis in ABDR amphihaploids depend on the specific type of univalent chromosome division

Meiosis in wide F₁ hybrids in the tribe Triticeae is complex and irregular. Despite extensive research, the meiotic pathways in such hybrids remain largely unknown. In this study, the effect of the genotypes of disomic wheat–rye substitution lines on the regulation of meiosis in ABDR amphihaploids was analysed by comparing microsporogenesis and seed set under self pollination in hybrids of normal bread wheat (Triticum aestivum L.) and wheats with single chromosome substitutions 1R(1A), 1Rv(1A), 2R(2D)₁, 2R(2D)₂, 2R(2D)₃, 5R(5D), 5R(5A), 6R(6A) with rye (Secale cereale L.). Three types of meiotic pathways were identified in the hybrids: reductional, equational, and equational+reductional. During the reductional type division, chromosomes randomly moved toward the poles at Anaphase I (AI) and separated sister chromatids at AII. Meiosis ended with tetrad formation, and the resulting plants were sterile ((2R(2D)₁ × R, 2R(2D)₂ × R). In the equational type division, the chromosomes aligned along the equator at Metaphase I and at AI separated into chromatids that moved toward the poles. This meiotic pathway led to dyad formation and partial fertility (especially in hybrids of 1Rv(1A) and 6R(6A)). In the equational+reductional pathway, some chromosomes divided in an equational manner while others were segregated reductionally within the same microsporocyte and this led to formation of tetrads with groups of uncoiled chromosomes. Comparison of the frequencies of each of the three meiotic pathways in different hybrid combinations suggests differences in contributions of individual chromosomes to the control of the entire meiotic process.     

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.     

Hexaploid triticales from hybrids of 'Langdon' durum D-genome substitutions with 'Gazelle' rye

The formation of unreduced gametes in some hybrids between disomic D‐genome substitutions (DS) of durum wheat cv.‘Langdon’ and rye provides a convenient approach for the rapid introduction of D‐genome chromosomes into hexaploid triticale. Meiotic pairing at metaphase I and seed fertility in spontaneous and colchicine‐induced amphidiploids derived from F₁ hybrids between a set of ‘Langdon’ DS and ‘Gazelle’ rye were analysed. The purpose was to determine the effects of the substitution of D‐genome chromosomes for their A‐ and B‐genome homoeologues on hexaploid triticale and to select stable disomic D‐genome substitutions of hexaploid triticale. The results showed that the disomic substitutions with D‐genome slightly increased the frequency of univalents (1.0‐3.13) compared with the ‘Langdon’ control primary hexaploid triticale (0.76). Substitutions 2D(2A) and 3D(3B) were partly desynaptic. The substitutions 1D(1A), 1D(1B) and 7D(7B) exhibited high seed fertility but the others had decreased fertility. Except for 2D(2A), 5D(5A), 3D(3B) and 5D(5B), 10 of the 14 possible hexaploid triticale D‐genome disomic substitutions have been obtained. The results suggest that the poor compensation ability of some D‐genome chromosomes for their homoeologous A‐ and B‐genome chromosomes is a major factor affecting meiotic stability and fertility in the hexaploid triticale D‐genome substitutions.     

Molecular cytogenetic characterization and phenotypic evaluation of new wheat–rye lines derived from hexaploid triticale ‘Certa’ × common wheat hybrids

Hexaploid triticale contains valuable genes from both tetraploid wheat and rye and plays an important role in wheat breeding programmes. In order to explore the potential of hexaploid triticale ‘Certa’ in wheat improvement, two crosses were made using ‘Certa’ as female parent, and common wheat cultivars ‘Jinmai47’ (JM47) and ‘Xinong389’ (XN389) as male parents. The karyotyping of BCF_4:5 lines from Certa/JM47//JM47 and F_5:6 lines from Certa/XN389 was investigated using sequential fluorescence in situ hybridization (FISH). One 1B(1R) substitution line and five 1BL.1RS whole‐arm translocation lines were identified, one of which was found lacking ω‐secalin locus. Many structural alterations on wheat chromosomes were detected in the progeny. Great morphologic differences resulting from genetic variations were observed, among which the photosynthetic capability was increased while grain quality was slightly improved. Compared with both parents, the stripe rust resistance at adult stage was increased in lines derived from Certa/JM47//JM47, while it was decreased in lines derived from Certa/XN389. These newly developed lines might have the potential to be utilized in wheat improvement programmes.     

Chromosomal location in triticale of leaf rust resistance genes introduced from Triticum monococcum

This study was undertaken to understand the inheritance of leaf rust resistance in line TM16 of Triticum monococcum ssp. monococcum var. macedonicum Papag. which is the source of resistance transferred into hexaploid triticale lines (Tcl/Tm). Thirty-two secondary tetraploid genotypes were analysed cytologicaly to identify substitutions of A^m-genome chromosomes by their homoeologous A-genome chromosomes from a leaf rust susceptible hexaploid triticale accession. Plants with one (or more) substituted chromosomes were inoculated with leaf rust at two growth stages. The disease phenotypes of these lines indicated that a major resistance gene was located on the short arm of T. monococcum chromosome 2A^m. An additional gene on chromosome 6A^m had complementary effects in enhancing the effects of the gene on chromosome 2A^m.     

Host plant resistance in some wild wheats to the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae)

A total of 259 accessions of wild Triticum species originating from different countries, along with 91 triticale (6×)× bread wheat true-breeding derivatives, two bread wheat, and three triticale cultivars were screened for resistance to the Russian wheat aphid, a serious insect pest of the wheat crop. Twenty-four entries with low damage ratings on the basis of amount of leaf rolling and leaf chlorosis were retested along with resistant and susceptible controls. On the basis of leaf roll damage ratings, eight entries including four Triticum monococcum var. boeoticum (T. boeoticum), one T. monococcum var. monococcum (T. monococcum), two T. timopheevii var. araraticum (T. araraticum), and one triticale cultivar were significantly superior to ‘Karl’ (susceptible control) wheat. Among these, four accessions — three T. boeoticum and one T. araraticum— were significantly superior to all other entries and were equal to the resistant control (PI 372129) in resistance rating based on leaf rolling and leaf chlorosis (except T. boeoticum TA 202). The leaf chlorosis damage rating of all accessions were significantly lower than that of the susceptible check.     

Genetic control of esterase-6 isozymes in hexaploid wheat and rye

Summary The EST-6 leaf esterase phenotypes from euploid, nullisomic-tetrasomic and rye chromosome addition and substitution lines of common wheat were determined using polyacrylamide gel electrophoresis. Evidence is presented to demonstrate that Est-6 is a new set of genes, that are expressed in the leaf. The Est-6 gene set were clearly distinguished from the Est-5 genes which are expressed in the grain. The three homoeoallelic loci, Est-A6, Est-B6 and Est-D6, were located on chromosomes 3A, 3B and 3D. An Est-R6 gene was located on chromosome 6R is involved in rye. Some considerations concerning homoeology between homoeologous group 3 of wheat and the rye chromosome 6R are made.