Comparative genetic mapping of loci affecting plant height and development in cereals

Restriction fragment length polymorphism (RFLP) mapping data for genes determining dwarfness (GA insensitive and GA sensitive), vernalisation response and photoperiodic response in wheat, rye and barley were compared and their homoeologous relationships discussed. The GA insensitive Rht genes of wheat are not related to the GA insensitive dwarfing genes of rye or barley; however, homoeology is present for two members of the GA sensitive dwarfing genes of wheat (Rht12) and rye (Ddw1), located on the translocated segments of the long arms of chromosomes 5A and 5R, respectively. The comparative mapping of the Triticeae group 5 vernalisation response genes of wheat, rye and barley, and the group 2 photoperiodic response genes of wheat and barley, show that both gene families are located in homoeologous regions of the particular chromosomes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Classification of GA response,Rht genes and culm length in Japanese varieties and landraces of wheat

Summary The GA response, Rht genes and culm length of 133 Norin varieties, 6 breeding lines and 16 landraces of Japanese wheat were investigated. Out of 133 Norin varieties tested, 103 were GA-insensitive and 30 GA-responsive. The 6 breeding lines were all GA-insensitive. Out of 16 landraces tested, 10 were GA-insensitive and 6 GA-responsive. Among the 10 GA-insensitive landraces, only Daruma had a Rht1 genotype. The other 9 had a Rht2 genotype. None of the landraces tested carried both Rht1 and Rht2 or Rht3. Out of the 103 GA-insensitive Norin varieties, 22 carried only Rht1, another 79 carried only Rht2, and only Norin 10 and Kokeshikomugi carried both Rht1 and Rht2. No tested variety carried Rht3. Some Norin varieties carrying Rht2 showed tall culms comparable to that of the rht tester line Chinese Spring. These results suggest that these varieties had a nullifier or modifier gene(s) or height promoting genes in the background controlling the height-reducing effect of Rht2. Conversely, six GA-responsive Norin varieties were as short as Akakomugi which carries the GA-responsive Rht genes, Rht8 and Rht9. The also seemed to carry a GA-responsive Rht gene or genes, and moreover all but one may carry gene(s) other than the Akakomugi genes. The origin of Rht1 and Rht2 of Norin 10 was speculated on the GA-response and Rht genotypes of its related varieties and landraces.     

Identification,distribution and effects on agronomic traits of the semi-dwarfing <Emphasis Type="Italic">Rht</Emphasis> alleles in Bulgarian common wheat cultivars

Bulgarian common wheat cultivars released in the period 1925–2003 were studied using the gibberellic acid (GA) test and microsatellite analysis of the Xgwm261 locus on chromosome 2DS to identify the semi-dwarfing (Rht) genes. The old cultivars, isolated through selection from landraces, carried rare alleles (211- and 215-bp) at Xgwm261 locus, and those developed by hybridisation to foreign cultivars, carried the 165- and 174-bp alleles. Forty-two (55.3%) of 76 modern cultivars were GA-responsive. The 192-bp allele, diagnostic for Rht8, was observed in 64 (84.2%) modern cultivars, of which 37 carry Rht8 alone, and 27 possess a combination of Rht8 and a GA-insensitive allele viz. Rht-B1d (17); Rht-D1b (6) and Rht-B1b (4). The 174-bp allele is present in seven cultivars, only one of which is photoperiod-sensitive, and the rest are day-length insensitive. The 203-bp allele was found in six modern cultivars. Cultivars carrying the Rht8 allele are the most widespread and some of them have been cultivated for a long period. Cultivars with the `Saitama 27' allele (Rht-B1d) are the most productive and are second in distribution in the country. The recently observed trend for increasing the proportion of cultivars with GA-insensitive Rht genes is probably due to their combination with the 192-bp allele of Xgwm261 locus tightly linked to the Ppd-D1, to the break of the link between the 174-bp allele and ppd-D1, and to the introduction of other genes influencing flowering time.     

In vitro culture variation of wheat and rye caused by genes affecting plant growth habit in vivo

Summary The influence of genes affecting the plant growth habit in wheat (Rht8 and Ppd1) and rye (ct1 and ct2) on tissue culture response was studied using immature embryos. Whereas the semi-dwarfing gene Rht8 seems to promote only a minor effect, the day-length sensitive allele ppd1 determined a major increase in callus growth and regeneration ability. With regards to their tissue culture efficiency, the four alleles studied could be ranked as follows: ppd1>Rht8>rht8>Ppd1.In contrast to wheat, the GA insensitive semi-dwarfing genes of rye (ct1 and ct2) appear not to influence in vitro response.     

Pleiotropic Effects of Genes for Reduced Height (Rht) and Day-Length Insensitivity (Ppd) on Yield and its Components for Wheat Grown in Middle Europe

Under field conditions in Germany over three growing seasons the pleiotropic effects on yield and its components of four sets of near isogenic lines carrying the GA insensitive dwarfing alleles Rht1, Rht2, Rht3, Rht1+2, Rht2+3 or rht (tall) in four different genetical backgrounds were examined together with 24 single chromosome recombinant lines segregating for the GA sensitive dwarfing gene Rht8 and the gene for day-length insensitivity Ppd1 in a ‘Cappelle-Desprez’ background. For the GA insensitive semi-dwarfs it was shown that in all three years a higher number of grains per ear was accompanied by a lower grain weight. Depending on the climatic conditions in a particular year, the increase in grain number was sufficient to compensate for the reduction in grain size and resulted in higher yields. For the Ppd1 allele yield advantages were found for wheats grown under environmental conditions of middle Europe.     

A classification of the NORIN 10 and Tom Thumb dwarfing genes in British,Mexican, Indian and other hexaploid bread wheat varieties

Summary The two semi-dwarfing genes Rht1 and Rht2 from Norin 10 have now been incorporated in successful varieties in use in most major wheat growing areas. The more potent dwarfing gene, Rht3, from Tom Thumb has been used in a limited way.These genes may be identified and classified by assessing the associated character of GA-insensitivity in the progeny from test crosses.This paper describes these classifications in the CIMMYT, Mexican, PBI, Cambridge and Indian breeding programmes and for a number of other international varieties.     

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.     

Genetical Studies of Gibberellic Acid Insensitivity in Rye (Secale cereale L.)

Genetic analysis of three semi-dwarf genotypes of rye (Secale cereale L.)—‘Moskowskij Karlik’, ‘Gülzow kurz’ and ‘R 18’, which were shown to be insensitive to applied gibberellic acid (GA₃), has been carried out by using a seedling test. It could be demonstrated that all of the three genotypes are carrying recessive alleles for GA-insensitivity. Whereas the alleles of ‘Moskowskij Karlik’ and ‘R 18’ seem to have the same locus on chromosome 5R, the GA-insensitivity of ‘Gülzow kurz’ is governed by a different gene, most probably located on chromosome 7R. The relationship between the genes (alleles) for GA-insensitivity and semi-dwarfness, including the symbolization of the Gai-genes as well as their utilization in rye breeding is discussed.     

Aluminum tolerance in triticale, wheat, and rye

Acid soils containing high levels of aluminum (Al) are known to severely limit plant growth on over 1.6 billion hectares worldwide. In the United States, a gradual decline in the pH of many soils both in the Great Plains as well as the Southeast, has caused many soils to become high in levels of free Al. This worldwide condition encouraged the analysis of wheat (Triticum aestivum L. em Thell.), triticale (X Triticosecale Wittmack), and rye (Secale cereale L.) germplasm from one of the major acid soil regions of the world (Brazil) in order to evaluate and compare the genetic potential of Al genes for cereal improvement. The objectives were to compare Al-tolerance levels in wheats, triticales, and ryes by measuring root elongation responses in Al-containing hydroponic nutrient solutions. Root elongation was impaired for all species grown in 1 mg/L concentrations of Al. Rye had the longest root regrowth and Al-sensitive wheats had the shortest root regrowth. The triticales containing a 2D(2R) substitution developed in the mid-1970s had the poorest root regrowth of all triticale types. The newly developed advanced triticale lines (AABBRR) yet to be released for commercial production showed the highest degree of Al tolerance of all the triticale types and approached or exceeded the levels observed in rye. This indicated that progress is being made in improving Al-tolerance of triticale in Brazil. Of all the old and new wheat varieties showing the highest degree of Al-tolerance, none of them were better than ‘BH 1146’ a variety that is at least 50 years old. This indicated that over the past 50 years, although Brazilian wheat breeders have made yield improvements in wheat production, they have not improved Al-tolerance. Rye showed a higher degree of Al-tolerance than the other cereals when tested in 1 mg/L of Al, but as expected, some variation was noted. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

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.     

Mutations blocking sensitivity to gibberellic acid promote ethylene-induced male sterility in wheat

Summary Ethylene is known to perturb normal reproductive development in wheat, particularly the development of functional pollen. Two experiments were carried out to test the hypothesis that increasing insensitivity to gibberellic acid (GA), conferred by the Rht genes, would be associated with increased male sterility in ethrel or Cerone^®-treated wheat. Wild type (WT=rht1/rht1, rht2/rht2), single dwarf (SD=Rht1/Rht1, rht2/rht2 or SD=rht1/rht1, Rht2/Rht2), and double dwarf (DD=Rht1/Rht1, Rht2/Rht2) near-isogenic lines in six genetic backgrounds were treated with ethrel or Cerone^® at the late tetrad to early uninucleate stage of pollen grain development. Ethrel induced pollen abortion in all genotypes but was highest for DD (41% above background) followed by SD (20%), and then WT genotypes (10%). Spikelet fertility decreased as the number of Rht alleles increased in response to ethrel or Cerone^® treatments. Expressed as a percent of controls, spikelet fertility was 56% for WT, 42% for SD, and 29% for DD. The consistent linear relationship between the number of Rht alleles and sensitivity of ethylene-induced male sterility suggests that GA and its recognition may exert a stabilizing effect in pollen development in the presence of stress or an ethylene shock.Paper No. 764 of the Department of Plant Breeding and Biometry, Cornell Agricultural Experiment Station.     

Identification, characterisation, and chromosome locations of rye and wheat specific ISSR and SCAR markers useful for breeding purposes

Rye (Secale cereale L. and S. strictum) offers potential to increase the genetic variability and to introduce desirable characters for wheat improvements. Cytogenetic techniques have been used to screen wheat lines containing rye chromatin. These techniques are not adequate since they are highly technical and time consuming. They are not suitable for breeding programs that require rapid screening of large numbers of genotypes. The main objective of this study was to develop and characterize ISSR and SCAR markers that can distinguish wheat from rye genome. Total DNA from wheat, rye, and triticale accessions from different provenances were amplified with ISSR primers in PCR assays. Three wheat-diagnostic sequences were identified. In addition three rye-diagnostic ISSR markers of which, one marker specifically diagnostic for Secale strictum were characterized. Pairs of primers flanking these specific sequences were designed to produce SCAR markers. Two SCAR markers were rye genome-specific. One SCAR was present in all the seven rye chromosome, and another was specific to rye chromosomes two, three, four, and seven. These newly developed ISSR and SCAR markers should be useful to wheat breeders screening genotypes that may contain rye chromatins.     

Intergeneric Transfer (Rye to Wheat) of a Gene(s) for Russian Wheat Aphid Resistance

An octoploid triticale was derived from the F, of a Russian wheat aphid-resistant rye, ‘Turkey 77’, and ‘Chinese Spring’ wheat. The alloploid was crossed to common wheat, and to ‘Imperial’ rye/‘Chinese Spring’ disomic addition lines. F₂, progeny from these crosses were tested for Russian wheat aphid resistance and C-banded. A resistance gene(s) was found to be associated with chromosome arm IRS of the ‘Turkey 77’ rye genome. A monotelosomic IRS (‘Turkey 77’) addition plant was then crossed with the wheat cultivar ‘Gamtoos’, which has the 1BL.1RS ‘Veery’ translocation. Unlike the IRS segment in ‘Gamtoos’, the ‘Turkey 77’-derived 1 RS telosome did not express the rust resistance genes Sr31 and Ar26, which could then be used as markers. From the F, a monotelosomic 1 RS addition plant that was also heterozygous for the 1BL. 1 RS translocation was selected and testerossed with an aphid-susceptible common wheat, ‘Inia 66’ Meiotic pairing between the rye arms resulted in the recovery of five euploid Russian-wheat-aphid-resistant plants. One recombinant also retained Sr31 and Lr26 and was selfed to produce translocation homozygotes.     

Comparison of rye,triticale, durum wheat and bread wheat genotypes for Fusarium head blight resistance and deoxynivalenol contamination

Small-grain winter cereal crops can be infected with Fusarium head blight (FHB) leading to mycotoxin contamination and reduction in grain weight and quality. Although a number of studies have investigated the genetic variation of genotypes within each small-grain cereal, a systematic comparison of the winter crops rye, triticale, durum and bread wheat for their FHB resistance, Fusarium-damaged kernels (FDK) and deoxynivalenol (DON) contamination across species is still missing. We have therefore evaluated twelve genotypes each of four crops widely varying in their FHB resistance under artificial infection with one DON-producing F. culmorum isolate at constant spore concentrations and additionally at crop-specific concentrations in two environments. Rye and triticale were the most resistant crops to FHB followed by bread and durum wheat at constant and crop-specific spore concentrations. On average, rye accumulated the lowest amount of DON (10.08 mg/kg) in the grains, followed by triticale (15.18 mg/kg) and bread wheat (16.59 mg/kg), while durum wheat had the highest amount (30.68 mg/kg). Genotypic variances within crops were significant (p ≤ .001) in most instances. These results underline the differing importance of breeding for FHB resistance in the different crops.     

Chromosomal location of aluminium tolerance genes in rye

Rye is known for its high tolerance of aluminium in the soils in comparison with wheat and other cereals. To localize the major gene/ genes controlling aluminium tolerance on the rye chromosomes, four series of wheat‐rye addition lines, two sets of triticale D(R) substitution lines and several wheat/rye translocation lines were tested in experiments on seedlings grown in nutrient solutions with various concentrations of aluminium. The results indicate that the major locus responsible for Al tolerance in rye is located on the short arm of chromosome 3R. The importance of these results for controlled introgressions into cereals is discussed.     

Identification of GA-insensitive Rht genes in Japanese modern varieties and landraces of wheat

Summary GA-insensitive Rht genes of 18 Japanese modern varieties and landraces were identified. Out of 12 modern varieties tested 6 carried only Rht1, and the other 6 carried only Rht2. No varieties carried both Rht1 and Rht2 or Rht3. The geographical distribution of the Rht genotypes in the Japanese modern varieties was clearly localized. All 6 landraces tested carried only Rht2.     

The Genetics and Breeding Potential of Rht12, a Dominant Dwarfing Gene in Wheat

Rht12, a dominant dwarfing gene of wheat, was shown to be located distally on the long arm of chromosome 5A. Lack of recombination with the awn inhibitor B1 suggested that Rht12 is cither tightly linked to this gene or is, in this material, a pleiotropic expression of the gene. Linkage to β-Amy-A1 was also very tight, indicating that Rht12 is present on the segment of chromosome SAL ancestrally translocated from 4AL. The close linkage to β-Amy-A1 also suggests that Rht12 is not a homoeoallele of the commercially important GA-insensitive dwarfing genes. Analysis of near-isogenic lines in a number of genetic backgrounds showed that Rht12 reduces height without altering ear size and significantly increases spikelet fertility. However its successful utilization in breeding programmes will require careful selection since in some backgrounds the gene reduces grain numbers and grain size. In all backgrounds, Rht12 delayed ear emergence time by around 6 days. A delay of this magnitude could, in many environments, adversely affect yield if it is not neutralized by altering the balance of other genes determining ear emergence time.     

Chromosomal Location of Genes for Gibberellic Acid Insensitivity in 'Chinese Spring' Wheat by Tetrasomic Analysis

The tetrasomics of the homoeologous groups 2, 5 and 7 of‘Chinese Spring’wheat were, together with the euploid standard, screened at the seedling stage for sensitivity to exogenously applied gibberellic acid (GA³). Whilst the seedling length of lines tetrasomic for group 2 chromosomes were taller and those for chromosomes 5A, 5D and 7D shorter in both treatments (with and without GA³) compared to the euploid control, the remaining tetrasomics — 5B, 7A and 7B — were significantly shorter than the euploids in the GA variant only. These results suggest the presence of additional genetic factors for GA insensitivity on chromosomes of the groups 5 and 7 of hexaploid wheat. This corresponds with the localization of GA insensitive dwarfing genes on the homoeologous chromosomes 5R and 7R in diploid rye.     

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.