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 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.     

An assessment of the potential of 4DS.4DL-4s l L translocation lines as a means of eliminating tall off types in semi-dwarf wheat varieties

Summary Wheat varieties tend to be chromosomally unstable producing on average 2–3% of plants with abnormal chromosome numbers. A number of semi dwarf wheat varieties, carrying the gibberellic acid insensitive dwarfing genes Rht1 or Rht2, have been seen to produce distinct tall off types due to reduction in dosage of the chromosome carrying the dwarfing gene. The UK variety Brigand, carrying Rht2 on chromosome 4D, produced very distinct tall off types when this chromosome was reduced in dosage. The frequency of tall off types was sufficiently high to cause the variety to fail United Kingdom statutory uniformity tests. An attempt to prevent the loss of chromosome 4D was made by constructing translocation chromosomes involving the short arm of chromosome 4D, which carries Rht2, and the long arm of chromosome 4S ^l from Aegilops sharonensis, which carries a gene(s) conferring preferential transmission. The work in this paper describes the field evaluation of two lines carrying 4DS.4DL-4S ^l L translocations, and demonstrates their success in preventing spontaneously occurring monosomy of chromosome 4D in semi-dwarf wheats.     

Allelic variation, sequence determination and microsatellite screening at the XGWM261 locus in Chinese hexaploid wheat (Triticum aestivum) varieties

Wheat microsatellite XGWM261, due to its closely linked to the dwarfing gene Rht8, has been adopted as the diagnostic molecular marker of Rht8. Screening 408 Chinese and 98 exotic varieties showed 13 allele variants in locus of XGWM261, with 6 alleles only to be found in Chinese varieties and 2 only in exotic varieties, respectively. Sequencing results of the 13 alleles revealed their absolute fragment sizes with 216, 212, 210, 206, 204, 202, 200, 196, 194, 192, 190, 174, and 164 bp, respectively. Allelic distribution analysis showed that the 204, 192, 174, and 164 bp alleles were prevailing in Chinese varieties, and the diagnostic 192 bp allele to Rht8 had a very high percentage in the Yellow and Huai River Valleys Facultative Wheat Zone than in the Northern Winter Wheat Zone in China. The GT → AC substitution at position 35 was found in 216, 200, and 174 bp alleles. Moreover, the AG insertion immediately at the end of CT-repeat region was also found in 216, 200, 174, and 164 bp alleles.     

Distribution of the Rht-B1b, Rht-D1b and Rht8 reduced height genes in autumn-sown Chinese wheats detected by molecular markers

Reduction of plant height has played a significant role in improving wheat production and knowledge of dwarfing genes in Chinese wheat will be very important for developing high yielding cultivars. Molecular markers were used to detect the presence of genes Rht-B1b (Rht1), Rht-D1b (Rht2) and Rht8 in 220 wheat genotypes from autumn-sown wheat regions in China. They include landmark landraces, leading cultivars and core parents involved in wheat breeding from the 1950s to the present. Results indicated that Rht-D1b and Rht8 dominate with frequencies of 45.5% and 46.8%, respectively, followed by Rht-B1b with a frequency of 24.5%. The frequencies of Rht-B1b and Rht-D1b increased, from 8.6 to 32.2% and 36.2 to 53.4%, respectively, whereas the frequency of Rht8 has remained constant over time, when compared with cultivars released before and after 1990. This indicates that both the Rht-B1b and Rht-D1b were successfully used in wheat production in Chinese environments. Our study shows that Rht-B1b and Rht-D1b can be used in the post-anthesis heat stressed environments. Rht-B1b in Chinese wheats is derived from two sources, viz., Norin 10 and the Italian introduction St2422/464 (Rht-B1b and Rht8). The identity of Rht-B1b in these two sources still needs to be confirmed. Suwon 86 carrying both Rht-B1b and Rht-D1b, and Chinese cultivars, Huixianhong and Yaobaomai, are the primary sources of Rht-D1b in Chinese wheats. It is likely that Rht-D1b in Youbaomai derives from an unknown introduction. Italian introductions such as Funo and Abbondanza, and Lovrin 10 with the 1B/1R translocation, and Chinese landraces are the three major sources of Rht8. This information will be very valuable for wheat breeding in China, and internationally.     

Reduced height (Rht) and photoperiod insensitivity (Ppd) allele associations with establishment and early growth of wheat in contrasting production systems

Near isogenic lines (NILs) varying for genes for reduced height (Rht) and photoperiod insensitivity (Ppd-D1a) in a cv. Mercia background (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c + Ppd-D1a, Rht-D1c, Rht12) were compared at one field site but within contrasting (‘organic’ vs. ‘conventional’) rotational and agronomic contexts, in each of 3 years. In the final year, further NILs (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht-B1b + Rht-D1b, Rht-D1b + Rht-B1c) in both Maris Huntsman and Maris Widgeon backgrounds were added together with 64 lines of a doubled haploid (DH) population [Savannah (Rht-D1b) × Renesansa (Rht-8c + Ppd-D1a)]. Assessments included laboratory tests of germination and coleoptile length, and various field measurements of crop growth between emergence and pre jointing [plant population, tillering, leaf length, ground cover (GC), interception of photosynthetically active radiation (PAR), crop dry matter (DM) and nitrogen accumulation (N), far red: red reflectance ratio (FR:R), crop height, and weed dry matter]. All of the dwarfing alleles except Rht12 in the Mercia background and Rht8c in the DHs were associated with reduced coleoptile length. Most of the dwarfing alleles (depending on background) reduced seed viability. Severe dwarfing alleles (Rht-B1c, Rht-D1c and Rht12) were routinely associated with fewer plant numbers and reduced early crop growth (GC, PAR, DM, N, FR:R), and in 1 year, increased weed DM. In the Mercia background and the DHs the semi-dwarfing allele Rht-D1b was also sometimes associated with reductions in early crop growth; no such negative effects were associated with the marker for Rht8c. When significant interactions between cropping system and genotype did occur it was because differences between lines were more exaggerated in the organic system than in the conventional system. Ppd-D1a was associated positively with plant numbers surviving the winter and early crop growth (GC, FR:R, DM, N, PAR, height), and was the most significant locus in a QTL analysis. We conclude that, within these environmental and system contexts, genes moderating development are likely to be more important in influencing early resource capture than using Rht8c as an alternative semi-dwarfing gene to Rht-D1b.     

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.     

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.     

NORIN 10 semi-dwarfism in tetraploid wheat and associated effects on yield

Summary A genetic study of a range of NORIN 10 based semi-dwarf durum wheats showed that only Gai/Rht 1, located on chromosome 4A, was present. No varieties carrying a second Gai/Rht allele were identified and deliberate attempts to introduce Gai/Rht2 into tetraploid wheats have so far been unsuccessful.In a spaced plant trial of homozygous random F₃ lines from two tall x semi-dwarf crosses, the semi-dwarfs has lower ear yields, due mainly to reduced kernel weight, but had higher tiller numbers than the tall genotypes. Although there was no difference in overall plant yield between talls and semi-dwarfs, an analysis of character associations within the semi-dwarf F₃'s showed positive height-yield and height-kernel weight correlations indicating that selection for tall dwarfs may be a useful breeding strategy in tetraploid wheats.     

The gibberellic acid insensitive dwarfing gene from the wheat variety Saitama 27

Summary Genetic analysis of a new weak source of GA-insensitivity derived from Saitama 27 showed that it was controlled by a single gene located on chromosome 4A and allelic to Rht1 and Rht3. the gene was shown to be present in many Italian and Yugoslavian varieties suggesting a selective advantage in some breeding programmes. Varietal comparisons indicated that the gene reduces height by around 11 per cent. This was combined with an increase in spikelet fertility and in the total number of grain developing in the ear. In this initial experiment a reduction in grain size eliminated the advantages of increased grain number. Following development of isogenic lines in a number of varietal backgrounds the true potential of the gene will be determined.     

The effect of major dwarfing genes on yield potential in spring wheats

Summary A composite convergent cross of 16 spring wheat parents produced a set of unselected progeny lines among which the major dwarfing genes, Rht1, Rht2 and Rht3, were distributed against a common random genetic background. Random subsets of these lines were grown under irrigation and optimal conditions in 4 experiments with replicated bordered plots in southern New South Wales in order to measure the dwarfing gene effect on yield potential. The dwarfing gene composition of each line was determined by test crossing and seedling responsiveness to gibberellic acid.Lodging was negligible in the two experiments in 1982. While present in the two in 1983, it was not strongly associated with yield. Grain yield levels were appropriately high (mean 5.9 t/ha). In all but 1 experiment the Rht1+Rht2 dwarf genotypes gave highest yields while the Rht3 group yielded on average 3% lower, Rht2 9% lower, Rht1 11% lower, and the non-dwarf or tall group yielded 24% lower. These yield differences were positively associated with harvest index, kernels per m² and kernels per spike, but negatively associated with mature plant height. Even within major dwarfing gene classes, grain yield was significantly and negatively associated with height.     

Resistance of cowpea and cereals to the parasitic angiospermStriga

Summary Striga species are parasitic angiosperms that attack many crops grown by subsistence farmers in sub-Saharan Africa and India. Control of the parasite is difficult and genetically resistant crops are the most feasible and appropriate solution. In cowpea, complete resistance toStriga gesnerioides has been identified. Breeding for resistance in sorghum has identified varieties with good resistance toS. asiatica in Africa and India. One variety was also resistant toS. hermonthica in W. Africa. No such resistance toStriga has been found in maize or millets.Resistant varieties have usually been sought by screening germplasm in fields naturally infested withStriga. However, laboratory techniques have also been developed, including anin vitro growth system used to screen cowpeas for resistance toS. gesnerioides. Two new sources of resistance in cowpea have been identified using the system. The technique has also been used to investigate the mechanisms of resistance in this crop. Two mechanisms have been characterised, both were expressed after penetration of cowpea roots by the parasite.The resistance of some sorghum varieties toStriga is controlled by recessive genes. In cowpea, resistance toStriga is controlled by single dominant genes. The genes for resistance are currently being transferred to cowpea varieties which are high yielding or adapted to local agronomic conditions. OneStriga resistant cowpea variety, Suvita-2, is already being grown widely by farmers in Mali. Reports of breakdown of resistance in cowpea toStriga have not yet been confirmed, but a wider genetic base to the resistance is essential to ensure durability ofStriga resistance.Abbreviations ICRISAT International Crops Research Institute for the Semi-Arid Tropics - IITA International Institute of Tropical Agriculture - LARS Long Ashton Research Station - SAFGRAD Semi-Arid Food Grain Research and Development     

Inheritance and transfer of multiple-flower character from Capsicum chinense into Capsicum annuum

Summary Capsicum annuum L. produces a single flower and thus a single fruit per branching node. In contrast, Capsicum chinense Jacq. yields two or more flowers per node. If genes for multiple-flowers per node could be transferred from C. chinese to C. annuum, it might be possible to breed C. annuum varieties with a more concentrated fruit set and potentially higher yield. Using progeny from an interspecific cross between C. annuum cv. NM 6-4 and C. chinense CA4, it has been determined that a minimum of five independently segregating chromosomal regions control the difference in flowering behavior between these two accessions. One of the segments is located on a translocated chromosome. Epistatic interactions among independent chromosomal regions appear to play a major role in the determination of this trait.Although it is unlikely that the multiple-flower character can be transferred to C. annuum in its full intensity while maintaining a horticulturally-acceptable phenotype, it may be possible to breed varieties which produce an average of more than 1.5 flowers per node, a value 50% higher than existing C. annuum varieties.     

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.     

Discovery,characterization and exploitation of Mlo powdery mildew resistance in barley

Summary Mlo resistance to barley powdery mildew is a relatively new kind of resistance. It was originally described in a powdery mildew resistant barley mutant in 1942 and has been mutagen-induced repeatedly since then. About 1970 it was also recognized in barley landraces collected in Ethiopia in the 1930s. It is unique in that 1) Mlo resistance does not conform to the gene-for-gene system; 2)mlo genes originating from different mutational events map as non-complementing recessive alleles in one locus; 3) all alleles confer the same phenotype, though with small quantitative differences; 4) it is effective against all isolates of the pathogen; and 5) the resistance is caused by rapid formation of large cell wall appositions at the encounter sites preventing penetration by the fungus. Powdery mildew isolates with elevated Mlo aggressiveness have been produced on barley in the laboratory, but have not been found in nature. Mlo resistance is considered very durable. The exploitation of Mlo resistance has been hampered by pleiotropic effects of themlo genes, vix. necrotic leaf spotting and reduced grain yield, but they have been overcome by recent breeding work. During the 1980s Mlo-resistant spring barley varieties have become cultivated extensively in several European countries, in 1990 on about 700,000 ha.     

Chromosomal location of dwarfing gene Rht12 in wheat

Summary The chromosomal location of the dwarfing gene Rht12 in the mutant winter wheat Karcagi 522M7K was investigated using F₂ monosomic analysis. The segregation ratio for F₂ progenies of Chinese Spring monosomics × Karcagi 522M7K, and that of Cheyenne monosomics × Karcagi 522M7K indicated that the near complete dominant dwarfing gene Rht12 is located on chromosome 5A. The heterozygous and hemizygous states of the genes Rht12 have the same effect on plant height.     

Expression of late maturity α-amylase in wheat containing gibberellic acid insensitivity genes

Summary Two wheat cultivars, Spica and Lerma 52, which consistently produce high levels of -amylase during the later stages of grain development (late maturity -amylase), were crossed with a set of four near-isogenic lines carrying the tall (rht) allele or one of the dwarfing genes Rht1, Rht2 or Rht3 (GA-insensitive alleles). The F₁ and F₂ populations were developed and analysed for grain -amylase and plant height. The Rht3 gene exhibited the strongest influence on plant height and strongly inhibited new -amylase synthesis during the later part of grain ripening. By comparison, Rht1 and Rht2 had a less pronounced effect but still significantly reduced the expression of late maturity -amylase. These observations suggest that gibberellic acid is involved either directly or indirectly in this phenomenon. The implications of the effect of dwarfing genes on expression of late maturity -amylase are discussed in relation to cultivar improvement and to the identification and control of high -amylase germplasm.     

Exploiting the diversity of <Emphasis Type="Italic">Viviparous-1</Emphasis> gene associated with pre-harvest sprouting tolerance in European wheat varieties

Pre-harvest sprouting (PHS) reduces the quality of wheat (Triticum aestivum L.) and the economic value of the grain. The objective of this study was to evaluate the diversity of the Viviparous-1B (Vp-1B) gene associated with PHS tolerance in a collection of 490 widely grown winter wheat varieties from central and northern Europe. Four alleles of Vp-1B were found in the wheat varieties tested, three of which (Vp-1Ba, Vp-1Bb and Vp-1Bc) had previously been identified in Chinese wheat varieties. The fourth was a new allele which had a 25-bp of deletion in the third intron region compared with the nucleotide sequence of Vp-1Ba, and was designated as Vp-1Bd. The frequencies of different alleles in this set of European wheat germplasm were: Vp-1Ba (54%) > Vp-1Bc (21%) > Vp-1Bd (20%) > Vp-1Ba + c (4%) > Vp-1Bb (1%), with Vp-1Bb being present only in two French varieties, ‘Altria’ and ‘Recital’. In addition, the frequencies of the alleles differed in varieties from different European countries. For example, Vp-1Ba had the highest frequency (76%) in varieties included in the UK National List (NL), but was least frequent in the Recommended List (RL) of Sweden (19%). Similarly, Vp-1Bc was present with the highest frequency (58%) in wheat varieties from Sweden, and the lowest in UK NL varieties (8%) while Vp-1Bd had the highest frequency of 32% in German varieties, and the lowest in Sweden varieties with only 8%. The Vp-1Ba allele was present in over half of the UK wheat varieties tested but the frequency was lower in RL varieties than in NL ones. Furthermore, heterogeneities were found between Vp-1Ba and Vp-1Bc in the varieties from Sweden, Netherlands, Germany and UK.     

Allelic variation at the dwarfing gene Rht8 locus and its significance in international breeding programmes

A microsatellite marker WMS261 developed at I.P.K., Gatersleben,Germany, co-segregates with the semi-dwarfing gene Rht8. Screens ofover 800 wheat varieties from 20 countries show 90% carry WMS-261alleles with 165, 174 or 192 base pairs (bp). The 192bp allele diagnosticof Rht8 occurs in most screened Southern European varieties. Anallele with 165 bp occurs in the majority of CIMMYT Mexican varieties andin most varieties bred in countries that utilize CIMMYT germplasm.Agronomic studies of single chromosome recombinant line mappingpopulations segregating for WMS 261 alleles with 165, 174 or 192 bp,show compared to the 174bp allele the 192bp allele reduces height byaround 8 cm and the 165 bp allele increases height by around 3cm. Theresults indicate the importance and widespread utilization of Rht8 asa dwarfing gene in southern Europe and suggest an adaptative significanceto the height promoting 165 bp allele in CIMMYT material.     

Search for novel genes for resistance to powdery mildew (Sphaerotheca fuliginae) in cucumber (Cucumis sativus)

Summary The current powdery mildew (Sphaerotheca fuligninea) resistant cucumber varieties suffer from leaf chlorosis during autumn, winter and early spring cultivation in the Netherlands. Therefore screening was carried out for novel powdery mildew resistance genes. From 177 accessions, derived from different sources, 108 accessions proved to be partially resistant to S. fuliginea. Crosses were made with 53 resistant accessions to distinguish the presence of novel genes. It is likely that the accessions C. sativus 2145, C. sativus LV 41, PI 188807, Vladivostokij, White and Yellow 1 have one or more recessive powdery mildew resistance genes, different from powdery mildew resistance genes of the line NPI, which was used for variety breeding. Powdery mildew resistance tests with S. fuliginea give similar results in different regions of the world.Abbreviations pmr powdery mildew resistance