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.     

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.     

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.     

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.     

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.     

Effects of dwarfing genes on yield and yield components under irrigated and rainfed conditions in wheat (Triticum aestivum L.)

Summary Near-isogenic tall (no dwarfing gene), semidwarf (Rht1 or Rht2) and dwarf (Rht1 + Rht2 or Rht3) spring wheat lines were evaluated for yield and yield components under irrigated and rainfed conditions. Under irrigated conditions, the dwarf and the semidwarf lines exhibited a significant yield advantage over the tall lines. Under rainfed conditions, the semidwarf lines outyielded the tall as well as the dwarf lines. Percent yield reduction in response to drought stress was highest with the dwarfs and lowest with the tall lines. Dry matter production of the tall lines and that of the semidwarf lines did not differ significantly and both produced significantly more dry matter than the dwarf lines under irrigated as well as rainfed conditions. Plant height and kernel weight decreased with increasing degree of dwarfness while number of kernels per spikelet, harvest index and days to heading increased under both moisture regimes. The dwarfing genes did not have any significant influence on number of tillers/m² and spikelets per spike in either moisture regime.     

The effects of whole chromosome substitutions differing in alleles for hybrid dwarfing and photoperiodic sensitivity on tissue culture response (TCR) in wheat

Summary The wheat varieties Chinese Spring, Marquis and Thatcher and five intervarietal Chinese Spring substitution lines for chromosomes 2B (2 lines), 2D (2 lines) and 4A^*, differing from the recipient variety in alleles for hybrid dwarfing genes and/or the photoperiodic response genes were analysed for tissue culture response (TCR). It could be demonstrated that only chromosome 2B has a major effect on TCR. Allelic variation at the hybrid dwarfing loci seems to have no effect on tissue culture performance, even in the combination D1D2D3 which gives the grass dwarf phenotype. Also the allelic constitution at the Ppd loci, gave no indication for a direct major effect of those alleles, however there seems to exist genetical factors for TCR on the homoeologous group 2 chromosomes which may be closely linked to the Ppd loci.     

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.     

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.     

Distribution of microsatellite alleles linked to Rht8 dwarfing gene in wheat

A wheat microsatellite locus, Xgwm 261, whose 192-bp allele closelylinked to the dwarfing gene Rht8, on chromosome 2D, was used toscreen 71 wheat cultivars from 13 countries to assess the variation at thislocus. Screening of this wheat collection showed that a 165-bp allele anda 174-bp allele were the most frequent. None of the New Zealand cultivarspossessed a 192-bp allele specific to Rht8, while only one cultivarfrom the US produced this important allele. The frequency of a 192-bpallele among these wheat cultivars was 5.63%. The highest allelefrequency was observed for a 174-bp fragment (52.11%) followed by a165-bp fragment (26.76%). The only durum wheat `Cham 1', did notshow any amplification due to the absence of D genome. Four new novelalleles, 180-bp, 198-bp, 200-bp and 204-bp present in the US and NewZealand wheat cultivars are reported.     

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.     

Production of intergeneric hybrid plants between <Emphasis Type="Italic">Sandersonia aurantiaca</Emphasis> and <Emphasis Type="Italic">Gloriosa rothschildiana</Emphasis> via ovule culture (Colchicaceae)

Intergeneric hybrid plants between Colchicaceous ornamental plants, Sandersonia aurantiaca and Gloriosa rothschildiana, have successfully been produced via ovule culture. After 5 days of reciprocal cross-pollination, a few pollen tubes were observed in the ovary. Although seeds were obtained in both reciprocal cross-combinations, they did not germinate under ex vitro conditions. Ovules with placental tissues isolated 14 days after cross-pollination of S. aurantiaca × G. rothschildiana were cultured on a medium containing 0.01 mg l^–1 each of -naphthaleneacetic acid (NAA) and 6-benzyladenine (BA), on which 41.5% of ovules swollen and produced callus-like structures within 10 weeks. When such swollen ovules were transferred to a medium containing 0.1 mg l^–1 each of NAA and BA, 7.5% of the initially cultured ovules produced rhizome-like structures within 6 weeks. Among the rhizome-like structures, those derived from two independent ovules (3.7% of the initially cultured ovules) produced multiple shoots following transfer to a medium containing 0.25 mg l^–1 NAA and 2.5 mg l^–1 BA. Multiple shoot-derived plantlets were established on a plant growth regulator-free medium, and they were successfully transplanted to pots. Early verification of their hybridity was accomplished by flow cytometry (FCM) analysis, chromosome observation and rDNA analysis.     

四倍体小麦地方品种矮蓝麦矮秆性状的遗传分析

四倍体圆锥小麦(Triticum turgidum L.ssp.turgidum)地方品种矮蓝麦是我国重要的小麦矮秆基因资源,经鉴定其矮秆特性对外源赤霉酸敏感。2012年配制矮蓝麦与2个高秆圆锥小麦的正反交组合,2012—2013年在四川绵阳分别种植F1、F2代和F2:3家系,对株高的遗传分析表明,矮蓝麦的矮秆性状受1对隐性基因控制。利用BSA法构建高秆和矮秆池筛选多态性SSR标记,并对矮蓝麦/青稞麦F2分离群体进行连锁分析,将目标基因定位于7AS染色体上,与标记GWM471的遗传距离为2.5 c M。矮蓝麦与矮秆番麦正反交的F1和F2群体表现非常相似的株高变异特征,初步推测矮蓝麦的矮秆基因是Rht22;进一步用高通量SNP和DAr T标记对两品种进行全基因组扫描,发现二者的遗传相似性高达98.7%~99.3%。因此认为,历史上矮蓝麦和矮秆番麦可能是同一品种,是通过人为交流而传播到不同地方。矮蓝麦携带的矮秆基因在人工合成六倍体小麦遗传背景中降低株高能力中等或较弱,在育种中需要聚合其他矮秆基因而被利用。     

Genetic and cytogenetic studies with a male sterile mutant of common wheat (Triticum aestivum)

Summary Genetic and cytogenetic studies were done on a male sterile mutant of the wheat variety Probus. Association of the 4A chromosome carrying the ms gene was studied in the F₁ of the male sterile Probus with Chinese Spring ditelo 4AS, with Transec and with line T4AS-DRS respectively. The presumption that the genetic male sterility of the mutant was due to a terminal deletion of the short arm of chromosome 4A could be confirmed.Linkage studies showed that the ms gene was at 17 map units from the dwarfing gene (Rht3) of Minister dwarf. This allows selection of short male sterile plants at the seedling stage.     

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.     

Isolation of a Spontaneous Chromosome Translocation in Common Wheat

The genes Ms2 for male sterility and Rht10 for dominant dwarfing located on the short arm of chromosome 4D in common wheat arc closely linked. Male sterile, dwarf F₁ plants from the cross of male sterile‘Chinese Spring’× dwarf‘Ai-bian’were backcrossed with the variety‘Chinese Spring, From this offspring a spontaneous chromosome translocation was isolated resulting in a recombinant male sterile and dwarf genotype.     

Molecular mapping of an aluminum tolerance locus on chromosome 4D of Chinese Spring wheat

Summary The tolerance of aluminum (Al) of disomic substitution lines having the chromosomes of the D genome of Triticum aestivum L. cv. Chinese Spring individually substituted for their homoeologues in T. turgidum L. cv. Langdon was investigated by the hematoxylin method. The disomic substitution lines involving chromosome 4D were more Al tolerant than Langdon. The tolerance was found to be controlled by a single dominant gene, designated Alt2, that is in the proximal region of the long arm of chromosome 4D. The locus was mapped relative to molecular markers utilizing a population of recombinant chromosomes from homoeologous recombination between Chinese Spring chromosome 4D and T. turgidum chromosome 4B. Comparison of the location of Alt2 in this map with a consensus map of chromosomes 4B and 4D based on homologous recombination indicated that Alt2 is in a vicinity of a 4 cM interval delineated by markers Xpsr914 and Xpsr1051. The Alt2 locus is distal to marker Xpsr39 and proximal to XksuC2. The Altw locus is also proximal to the Knal locus on chromosome 4D that controls K⁺/Na⁺ selectivity and salt tolerance. In two lines, Alt 2 and Knal were transferred on a single 4D segment into the long arm of T. turgidum chromosome 4B.     

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.     

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.     

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.