Interaction between rye B-chromosomes and wheat genetic systems controlling homoeologous pairing

The interactive effect on homoeologous pairing of rye B-chromosomes with the absence of both pairing suppressor (3A, 3D, 5B) and promotor (3B, 5A, 5D) chromosomes of common wheat (Triticum aestivum L.) is analyzed by comparison of pairing at Metaphase I of 27-, 27+2B, 28- and 28+2B-chromosome plants. These plants were obtained from crosses between the respective wheat monosomics (2n=41) and rye plants (Secale cereale L.) carrying or not carrying two B-chromosomes (2n=14 or 14+2Bs). —The effect of rye B-chromosomes on pairing depends on the function of the wheat chromosome which is absent in the appropriate hybrids, i.e., rye B-chromosomes have a suppressor effect on pairing when the pairing suppressing wheat chromosomes 3A, 3D or 5B are absent, while they behave as promotors when the pairing promoting chromosomes 3B, 5A or 5D are absent.     

Cytogenetics of Triticum x Dasypyrum hybrids and derived lines

Genomic in situ hybridization was used to study Triticum x Dasypyrum wide hybrids and derived lines. A cytogenetic investigation was carried out in progenies of (i) amphiploids derived from T. turgidum var. durum (T. durum; 2n = 14; genomes AABB) x D. villosum (2n = 14; genome VV), (ii) three-parental hybrids (T. durum x D. villosum) x T. aestivum (2n = 42, genomes A'A'B'B'D'D'), and (iii) T. aestivum aneuploid lines carrying D. villosum chromosomes or chromatin. The amphiploids derived from T. durum x D. villosum showed a stable chromosomal constitution, made up of 14 V chromosomes, 14 chromosomes carrying the wheat A genome and 14 chromosomes carrying the B genome. High karyological instability was observed in the progenies of three-parental hybrids ([T. durum x D. villosum] x T. aestivum). Plants having the expected 14 A chromosomes, 14 B chromosomes, 7 D chromosomes, and 7 V chromosomes were rather rare (4.5%). Many progeny plants (45.5%) had the hexaploid wheat genome with 42 chromosomes and lacked any detectable D. villosum chromatin. Other plants (50%) had 14 A chromosomes and 14 B chromosomes, plus variable numbers of D and V chromosomes, the former being better retained than the latter in most cases. Some T. aestivum lines carrying D. villosum chromosomes or chromatin, as the result of addition, substitution, or recombination events or even a combination of these karyological events, were found to be stable. Other lines were unstable, and these lines carried 1V, 3V, or 5V chromosomes or their portions. Substitution or recombination events where 1V chromosomes were involved could concern the homeologous counterparts in both the A and B and D genomes of wheat. No line could be recovered where the shorter arm of 3V chromosomes was present. Changes in the morphology and banding pattern of V chromosomes were observed in hybrids that did not carry the entire D. villosum complement. By comparing the results of our cytogenetic analyses with certain phenotypic characteristics of the lines studied, genes for discrete traits could be assigned to specific V chromosomes or V chromosome arms. From the frequency of V chromosomes that were involved in chromatin exchanges with or substituted for one of their homeologous counterparts in the A, B, and D wheat genomes, it was inferred that D. villosum belongs to the same phyletic lineage as T. urartu (donor of the A genome of wheat) and Aegilops speltoides (B genome), and that Ae. squarrosa (D genome) diverged earlier from D. villosum.     

Genetic analysis of grain protein-content,grain yield and thousand-kernel weight in bread wheat

Grain yield and grain protein content are two very important traits in bread wheat. They are controlled by genetic factors, but environmental conditions considerably affect their expression. The aim of this study was to determine the genetic basis of these two traits by analysis of a segregating population of 194 F(7) recombinant inbred lines derived from a cross between two wheat varieties, grown at six locations in France in 1999. A genetic map of 254 loci was constructed, covering about 75% of the bread wheat genome. QTLs were detected for grain protein-content (GPC), yield and thousand-kernel weight (TKW). 'Stable' QTLs (i.e. detected in at least four of the six locations) were identified for grain protein-content on chromosomes 2A, 3A, 4D and 7D, each explaining about 10% of the phenotypic variation of GPC. For yield, only one important QTL was found on chromosome 7D, explaining up to 15.7% of the phenotypic variation. For TKW, three QTLs were detected on chromosomes 2B, 5B and 7A for all environments. No negative relationships between QTLs for yield and GPC were observed. Factorial Regression on GxE interaction allowed determination of some genetic regions involved in the differential reaction of genotypes to specific climatic factors, such as mean temperature and the number of days with a maximum temperature above 25 degrees C during grain filling.     

IDENTIFICATION OF MONOSOMES FOR SIX CHROMOSOMES IN GOSSYPIUM HIRSUTUM

Six primary monosomes of the amphidiploid species Gossypium hirsutum have been identified as separate chromosomes of the complement. On the basis of cytogenetic tests involving translocations, a diploid species of the D genome group, and genetic markers, 4 of the 6 monosomes were designated chromosomes 1, 2, 4 and 6 of the A subgenome and 2 were designated chromosomes 17 and 18 of the D subgenome. Fourteen other monosomes were identified as duplicates of 3 of the above 6 primaries.     

用Langdon二体代换系统建立小麦染色体RAPD标记

以一套Ｌａｎｇｄｏｎ硬粒小麦二体代换系及其亲本Ｌａｎｇｄｏｎ、中国春和中国春双端体为材料,研究适于硬粒小麦和普通小麦的理想ＲＡＰＤ分析条件,进行小麦Ａ、Ｂ和Ｄ染色体组各个染色体的ＲＡＰＤ分析。结果表明,ＡｍｐｌｉＴａｑＳｔｏｆｆｅｌｆｒａｇｍｅｎｔ比ＴａｑＤＮＡＰｏｌｙｍｅｒａｓｅ优越。所用１２个随机引物中,７个引物扩增出的１３个特异产物,可确定在硬粒小麦ＬａｎｇｄｏｎＡ、Ｂ染色体组和中国春Ｄ染色体组中的１０个个别染色体上。４个标记进一步定位在相应的４个染色体臂上。结果还表明,用Ｌａｎｇｄｏｎ二体代换系统、中国春双端体为材料,容易得到重复性高、特异性强的ＲＡＰＤ标记。     

Physical molecular maps of wheat chromosomes

In bread wheat, a set of 527 simple sequence repeats (SSRs) were tried on 164 deletion lines, leading to a successful mapping of 270 SSRs on 313 loci covering all 21 chromosomes. A maximum of 119 loci (38%) were located on B subgenome, and a minimum of 90 loci (29%) mapped on D subgenome. Similarly, homoeologous group 7 carried a maximum of 61 loci (19%), and group 4 carried a minimum of 22 loci (7%). Of the cited 270 SSRs, 39 had multiple loci, but only eight of these detected homoeologous loci. Linear order of loci in physical maps largely corresponded with those in the genetic maps. Apparently, distances between each of only 26 pairs of loci significantly differed from the corresponding distances on genetic maps. Some loci, which were genetically mapped close to the centromere, were physically located distally, while other loci that were mapped distally in the genetic maps were located in the proximal bins in the physical maps. This suggested that although the linear order of the loci was largely conserved, variation does exist between genetic and physical distances.Electronic Supplementary Material Supplementary material is available for this article at .     

A Detailed RFLP Map of Cotton, Gossypium Hirsutum X Gossypium Barbadense: Chromosome Organization and Evolution in a Disomic Polyploid Genome

We employ a detailed restriction fragment length polymorphism (RFLP) map to investigate chromosome organization and evolution in cotton, a disomic polyploid. About 46.2% of nuclear DNA probes detect RFLPs distinguishing Gossypium hirsutum and Gossypium barbadense; and 705 RFLP loci are assembled into 41 linkage groups and 4675 cM. The subgenomic origin (A vs. D) of most, and chromosomal identity of 14 (of 26), linkage groups is shown. The A and D subgenomes show similar recombinational length, suggesting that repetitive DNA in the physically larger A subgenome is recombinationally inert. RFLPs are somewhat more abundant in the D subgenome. Linkage among duplicated RFLPs reveals 11 pairs of homoeologous chromosomal regions-two appear homosequential, most differ by inversions, and at least one differs by a translocation. Most homoeologies involve chromosomes from different subgenomes, putatively reflecting the n = 13 to n = 26 polyploidization event of 1.1-1.9 million years ago. Several observations suggest that another, earlier, polyploidization event spawned n = 13 cottons, at least 25 million years ago. The cotton genome contains about 400-kb DNA per cM, hence map-based gene cloning is feasible. The cotton map affords new opportunities to study chromosome evolution, and to exploit Gossypium genetic resources for improvement of the world's leading natural fiber.     

Genetic analysis of durable powdery mildew resistance in a common wheat line

Genetic studies using monosomic and hybridological analyses had confirmed that resistance of a common wheat line k-15560 to powdery mildew in seedling stage was conditioned by one dominant gene located on chromosome 7B, and resistance in adult stage was controlled by two dominant genes. Cytological analysis of meiosis in the F1 monosomic hybrids has revealed reciprocal translocation involving chromosomes 2A/7A. In the F1 monosomic hybrids genes, causing a decrease in pairing were found on chromosomes 3B and 4D, and genes enhancing pairing--on chromosomes 2A and 3A.     

易组"太谷核不育基因"(Ms2)基因定位的研究

将在远缘杂交中由普通小麦（AABBDD）4D染色体易组导入六倍体小黑麦（AABBRR）以及硬粒小麦（AABB）的太谷核不育基因Ms2（原位于普通小麦4D染色体短臂距着丝点31．2cM的显性雄性不育核基因）。重新异回普通小麦染色体组中,所获得携带易组Ms2基因的新型太谷核不育小麦其显性雄性不育特性表达正常,且雄性不育株的雌性可育机制正常,对不育株幼穗花粉母细胞减数分型期染色体构型的观察可见其为整倍体（2n=42),尚未发现回归普通小麦的易组太谷核不育与原位 的太谷核不育基因有不同的表型。采用系统的标志基因测交法对回归普通小麦的易组太谷不育基因进行测交定位,发现易组Ms2基因与普通小麦显性秆标志基因Rht3连锁,从而将其定位于普通小麦4B 色体虎Rht3基因9.7cM处,新位点被命名为Ms2（4BS）,对Ms2基因在六倍体小黑麦与原太谷核不育小麦远缘杂交中位时的走向,普通小麦4A与4B染色体的互换更名以及Ms2（4BS）新位点的开发利用进行了讨论,认为异源多倍体生物核基因的组间易位倾向于从供体染色体向进化亲缘关系较密切,且染色体序数与染色体臂相同的部分同源染色体易位;1988年第7届国际小麦遗传学会对普通小麦4A与4B染色体的互换更名是正确的;Ms2（4BS）作为一个新型的遗传标记,作为小麦族内所有携带B染色体组的物种的育种工具和在拓建各为小麦种质资源的基因库等方面均有广泛的用途。     

QTL mapping for some grain traits in bread wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Grain traits are important agronomic attributes with the market value as well as milling yield of bread wheat. In the present study, quantitative trait loci (QTL) regulating grain traits in wheat were identified. Data for grain area size (GAS), grain width (GWid), factor form density (FFD), grain length-width ratio (GLWR), thousand grain weight (TGW), grain perimeter length (GPL) and grain length (GL) were recorded on a recombinant inbred line derived from the cross of NW1014?×?HUW468 at Meerut and Varanasi locations. A linkage map of 55 simple sequence repeat markers for 8 wheat chromosomes was used for QTL analysis by Composite interval mapping. Eighteen QTLs distributed on 8 chromosomes were identified for seven grain traits. Of these, five QTLs for GLWR were found on chromosomes 1A, 6A, 2B, and 7B, three QTLs for GPL were located on chromosomes 4A, 5A and 7B and three QTLs for GAS were mapped on 5D and 7D. Two QTLs were identified on chromosomes 4A and 5A for GL and two QTLs for GWid were identified on chromosomes 7D and 6A. Similarly, two QTLs for FFD were found on chromosomes 1A and 5D. A solitary QTL for TGW was identified on chromosome 2B. For several traits, QTLs were also co-localized on chromosomes 2B, 4A, 5A, 6A, 5D, 7B and 7D. The QTLs detected in the present study may be validated for specific crosses and then used for marker-assisted selection to improve grain quality in bread wheat.     

Wheat chromosome instability in the selfed progeny of the double monosomics 1Rv-1A

Structural alterations of chromosomes are often found in wheat-rye hybrids. In the majority of cases modifications are observed for rye chromosomes, yet chromosome aberration cases are described for wheat, including the progeny of Triticum aestivum disomic and monosomic addition lines. Since wheat-rye substitution and translocation lines are the source of rye chromatin in wheat breeding programs, the information on possible chromosome changes in the genomes of introgressive forms is important. Chromosome behavior in F₁ meiosis and chromosomal composition of F₂ karyotypes for double monosomics 1Rv-1A were studied by applying C-banding, genomic in situ hybridisation (GISH) using rye genomic DNA, and sequential in situ hybridization using repetitive sequences pAs1, pSc119.2 and centromere specific pAet-06 as probes. The double monosomics 1Rv-1A were obtained by crossing of disomic substitution line with chromosome 1A replaced by Secale cereale 1Rv in the bread wheat Saratovskaya 29 (S29) background with S29. The results indicated a high frequency of bipolar chromosome 1Rv orientation, as compared to 1A, at metaphase I (MI) (58.6 and 34.7 % of meiocytes, respectively), and, at anaphase I (AI), chromatid segregation of 1Rv compared to 1A (70.53 and 32.14 % of meiocytes, respectively). In few cases desynapsis of wheat homologues was observed, at AI, the chromosomes randomly distributed between the poles or underwent chromatid segregation. At AI, the two wheat homologues separated onto sister chromatids in 10.89 % of cells.The plants F2 karyotypes were marked with aneuploidy not only of chromosomes 1A and 1Rv, but also of 1D, 2D, 3D, 3B, 3A, 4A, 6D, 6B, 6A, and 7D. Structural changes were observed for the chromosomes of the first homoeologous group (1Rv, 1A, 1D, 1B), as well as for 2B, 5D, 6B, and 7B. The chromosomes 1Rv and 6B often demonstrated aberrations. The types of aberrations were centromeric break, deletions of various sizes, and a changed repeat pSc119.2 localization pattern.     

Cotton genome mapping with new microsatellites from Acala ‘Maxxa’ BAC-ends

Fine mapping and positional cloning will eventually improve with the anchoring of additional markers derived from genomic clones such as BACs. From 2,603 new BAC-end genomic sequences from Gossypium hirsutum Acala ‘Maxxa’, 1,316 PCR primer pairs (designated as MUSB) were designed to flank microsatellite or simple sequence repeat motif sequences. Most (1164 or 88%) MUSB primer pairs successfully amplified DNA from three species of cotton with an average of three amplicons per marker and 365 markers (21%) were polymorphic between G. hirsutum and G. barbadense. An interspecific RIL population developed from the above two entries was used to map 433 marker loci and 46 linkage groups with a genetic distance of 2,126.3 cM covering approximately 45% of the cotton genome and an average distance between two loci of 4.9 cM. Based on genome-specific chromosomes identified in G. hirsutum tetraploid (A and D), 56.9% of the coverage was located on the A subgenome while 39.7% was assigned to the D subgenome in the genetic map, suggesting that the A subgenome may be more polymorphic and recombinationally active than originally thought. The linkage groups were assigned to 23 of the 26 chromosomes. This is the first genetic map in which the linkage groups A01 and A02/D03 have been assigned to specific chromosomes. In addition the MUSB-derived markers from BAC-end sequences markers allows fine genetic and QTL mapping of important traits and for the first time provides reconciliation of the genetic and physical maps. Limited QTL analyses suggested that loci on chromosomes 2, 3, 12, 15 and 18 may affect variation in fiber quality traits. The original BAC clones containing the newly mapped MUSB that tag the QTLs provide critical DNA regions for the discovery of gene sequences involved in biological processes such as fiber development and pest resistance in cotton. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Genetic mapping and QTL analysis of fiber-related traits in cotton (<Emphasis Type="Italic">Gossypium</Emphasis>)

Cotton, the leading natural fiber crop, is largely produced by two primary cultivated allotetraploid species known as Upland or American cotton (Gossypium hirsutum L.) and Pima or Egyptian cotton (G. barbadense L.). The allotetraploid species diverged from each other and from their diploid progenitors (A or D genome) through selection and domestication after polyploidization. To analyze cotton AD genomes and dissect agronomic traits, we have developed a genetic map in an F₂ population derived from interspecific hybrids between G. hirsutum L. cv. Acala-44 and G. barbadense L. cv. Pima S-7. A total of 392 genetic loci, including 333 amplified fragment length polymorphisms (AFLPs), 47 simple sequence repeats (SSRs), and 12 restriction fragment length polymorphisms (RFLPs), were mapped in 42 linkage groups, which span 3,287 cM and cover approximately 70% of the genome. Using chromosomal aneuploid interspecific hybrids and a set of 29 RFLP and SSR framework markers, we assigned 19 linkage groups involving 223 loci to 12 chromosomes. Comparing four pairs of homoeologous chromosomes, we found that with one exception linkage distances in the A-subgenome chromosomes were larger than those in their D-subgenome homoeologues, reflecting higher recombination frequencies and/or larger chromosomes in the A subgenome. Segregation distortion was observed in 30 out of 392 loci mapped in cotton. Moreover, approximately 29% of the RFLPs behaved as dominant loci, which may result from rapid genomic changes. The cotton genetic map was used for quantitative trait loci (QTL) analysis using composite interval mapping and permutation tests. We detected seven QTLs for six fiber-related traits; five of these were distributed among A-subgenome chromosomes, the genome donor of fiber traits. The detection of QTLs in both the A subgenome in this study and the D subgenome in a previous study suggests that fiber-related traits are controlled by the genes in homoeologous genomes, which are subjected to selection and domestication. Some chromosomes contain clusters of QTLs and presumably contribute to the large amount of phenotypic variation that is present for fiber-related traits.Communicated by J. Dvorak     

Genome‐specific introgression between wheat and its wild relative Aegilops triuncialis

Introgression of sequences from crop species in wild relatives is of fundamental and practical concern. Here, we address gene flow between cultivated wheat and its widespread polyploid relative, Aegilops triuncialis, using 12 EST‐SSR markers mapped on wheat chromosomes. The presence of wheat diagnostic alleles in natural populations of the barbed goatgrass growing in proximity to cultivated fields highlights that substantial gene flow occurred when both species coexisted. Furthermore, loci from the A subgenome of wheat were significantly less introgressed than sequences from other subgenomes, indicating differential introgression into Ae. triuncialis. Gene flow between such species sharing nonhomeologous chromosomes addresses the evolutionary outcomes of hybridization and may be important for efficient gene containment.     

Chromosomal assignment of genes controlling salt-soluble proteins (albumins and globulins) in wheat and related species

Summary Salt-soluble proteins from the endosperms of wheat, barley, and rye have been separated by nonequilibrium electrofocusing x electrophoresis. Genes encoding 14 of the 25 components observed in wheat have been unambiguously assigned to 10 different chromosomes (1B, 3B, 3D, 4A, 4D, 5B, 6B, 6D, 7B, 7D) by analysis of the compensated nulli-tetrasomic series. Five more wheat proteins seem to be controlled by group 2 chromosomes. Analysis of wheat-barley and wheat-rye addition lines has led to the location of genes for 6 out of 20 barley proteins in 4 different chromosomes (1H, 3H, 4H, 6H; 1H is homoeologous to group 7 chromosomes of wheat) and of genes for 5 out of 20 rye proteins in two different chromosomes (2R, 4R). The relationship between the proteins reported here and previously characterized ones is discussed.     

The genetic control of the α-amylase isozymes of the durum wheat (<Emphasis Type="Italic">Triticum durum</Emphasis> Desf.)

The hybridoiogical analysis was provided on several durum wheat genotypes with utilizing three F₂ populations developed from the crossing between parental forms that differed in the invariable malt-zone triplet on elecirophoretic spectrum of α-amylase. Three components of this zone are controlled by three genes with an independent way of inheritance: one of them is located on the 6B or 5B chromosome, and two genes are located on the chromosomes of A subgenome. The article is published in the original.     

Molecular cloning of three homoeologous cDNAs encoding orthologs of the maize KNOTTED1 homeobox protein from young spikes of hexaploid wheat

The plant knotted1 (kn1)-like homeobox genes are known to play important roles in the maintenance of shoot apical meristem (SAM), determination of cell fate and differentiation of vegetative tissues. To study structural diversity of the three homologous loci encoding a KN1-like homeobox protein in the hexaploid wheat genome, we isolated clones from a cDNA library of young spikes of Japanese common wheat cultivar 'Norin 26'. Three different but highly homologous cDNAs were isolated and their sequences were determined. The mean homology of the deduced amino acid sequences was 96% as compared to the barley ortholog KNOX3. The wheat kn1-like homeobox proteins named WKNOX1 are encoded by a single set of homologous genes on the homologous group 4 chromosomes in the three component genomes of common wheat, i.e. 4A, 4B and 4D. The nucleotide sequence data and the Southern blot pattern suggested that the three homologous loci of wknox1 genes are highly conserved through polyploid evolution of wheat. They were expressed in SAM-containing shoots and young spikes but not in developed leaves, glumes and lemmas and callus tissues. The ectopic expression of the wknox1 was observed in lemma of wheat Hooded (Hd) mutants. The result suggested that the Hd gene is a dominant allele of the wknox1 locus on chromosome 4A.     

Putative interchromosomal rearrangements in the hexaploid wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) genotype ‘Chinese Spring’ revealed by gene locations on homoeologous chromosomes

Background

Chromosomal rearrangements are a major driving force in shaping genome during evolution. Previous studies show that translocated genes could undergo elevated rates of evolution and recombination frequencies around these genes can be altered. Based on the recently released genome sequences of Triticum urartu, Aegilops tauschii, Brachypodium distachyon and bread wheat, an analysis of interchromosomal translocations in the hexaploid wheat genotype ‘Chinese Spring’ (‘CS’) was conducted based on chromosome shotgun sequences from individual chromosome arms of this genotype.

Results

A total of 720 genes representing putative interchromosomal rearrangements was identified. They were distributed across the 42 chromosome arms. About 59% of these translocated genes were those involved in the well-characterized translocations involving chromosomes 4A, 5A and 7B. The other 41% of the genes represent a large numbers of putative interchromosomal rearrangements which have not yet been described. The number of the putative translocation events in the D subgenome was about half of those presented in either the A or B subgenomes, which agreed well with that the times of interaction between the A and B subgenomes almost doubled that between either of them and the D subgenome.

Conclusions

The possible existence of a large number of interchromosomal rearrangements detected in this study provide further evidence that caution should be taken when using synteny in ordering sequence contigs or in cloning genes in hexaploid wheat. The identification of these putative translocations in ‘CS’ also provide a base for a systematic evaluation of their presence or absence in the full spectrum of bread wheat and its close relatives, which could have significant implications in a wide array of fields ranging from studies of systematics and evolution to practical breeding.

     

Whole Genome Association Mapping of Plant Height in Winter Wheat (Triticum aestivum L.)

The genetic architecture of plant height was investigated in a set of 358 recent European winter wheat varieties plus 14 spring wheat varieties based on field data in eight environments. Genotyping of diagnostic markers revealed the Rht-D1b mutant allele in 58% of the investigated varieties, while the Rht-B1b mutant was only present in 7% of the varieties. Rht-D1 was significantly associated with plant height by using a mixed linear model and employing a kinship matrix to correct for population stratification. Further genotyping data included 732 microsatellite markers, resulting in 770 loci, of which 635 markers were placed on the ITMI map plus a set of 7769 mapped SNP markers genotyped with the 90 k iSELECT chip. When Bonferroni correction was applied, a total of 153 significant marker-trait associations (MTAs) were observed for plant height and the SSR markers (−log₁₀ (P-value) ≥4.82) and 280 (−log₁₀ (P-value) ≥5.89) for the SNPs. Linear regression between the most effective markers and the BLUEs for plant height indicated additive effects for the MTAs of different chromosomal regions. Analysis of syntenic regions in the rice genome revealed closely linked rice genes related to gibberellin acid (GA) metabolism and perception, i.e. GA20 and GA2 oxidases orthologous to wheat chromosomes 1A, 2A, 3A, 3B, 5B, 5D and 7B, ent-kaurenoic acid oxidase orthologous to wheat chromosome 7A, ent-kaurene synthase on wheat chromosome 2B, as well as GA-receptors like DELLA genes orthologous to wheat chromosomes 4B, 4D and 7A and genes of the GID family orthologous to chromosomes 2B and 5B. The data indicated that besides the widely used GA-insensitive dwarfing genes Rht-B1 and Rht-D1 there is a wide spectrum of loci available that could be used for modulating plant height in variety development.