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。     

Intrachromosomal mapping of genes for dwarfing (Rht12) and vernalization response (Vrn1) in wheat by using RFLP and microsatellite markers

For intrachromosomal mapping of the dominant GA-sensitive dwarfing gene Rht12 and the vernalization response gene Vrn1 on chromosome 5 A, an F₂ population was established using a wide (synthetic) wheat cross. In addition to restriction fragment length polymorphism (RFLP) probes four microsatellite markers were incorporated. Rht12 was mapped distally to four RFLP loci (Xmwg616, Xpsr164, Xwg114, Xpsr1201) and three microsatellite markers (Xgwm179, Xgwm410, Xgwm291), known to be located on the segment of chromosome SAL which was ancestrally translocated and is homoeologous to Triticeae 4 L. The map position of Rht12 suggests that it is homoeologous to the dominant GA-sensitive dwarfing gene Ddw1, present on chromosome 5RL. The vernalization response gene Vrn1 showed linkage to Xwg644, as might be expected from comparative maps.     

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.     

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.     

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.     

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.     

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.     

The association of genes for purple coleoptile with chromosomes of the wheat variety Mironovskaya 808

Summary The association of genes for purple pigment in the coleoptile with the chromosomes of the winter wheat variety Mironovskaya 808 was investigated using monosomic F₂ analysis. The segregation ratio for F₂ hybrids of Chinese Spring monosomics x Mironovskya 808 seems to indicate that the purple colour of the coleoptile is determined by two dominant genes, Rc₃ and Rc₄, which are located on the chromosomes 7D and 6B respectively, and which reinforce each other. Apart from these two genes, suppressors found on the chromosomes 2A, 2B, 2D, 4B and 6A also play a role in the intensity of the purple colour.With the aid of a Chinese Spring telocentric chromosome marker it was observed that the Rc₃ gene is located on the chromosome arm 7DS, at a distance of 16±4.23 crossover units from the centromere.     

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.     

A statistical assessment of genotypic sensitivity of groundnut (Arachis hypogae L.) to drought in line source sprinkler experiments

Summary Crosses between common bean genotypes from different centres of origin can result in F₁ plants with dwarfing that is severe enough to prevent formation of F₂ seed. This hybrid dwarfness is controlled by two complementary genes, Dl-1 and Dl-2. The dominant alleles at both loci must be present to cause dwarfing, and an increased dose of either allele increases the severity of the dwarfing. Since dwarfing restricts crossing among gene pools, it holds interests for bean breeding and studies of bean evolution and physiology. Results of grafting studies are described which confirm the previously reported dependance of dwarfing on independent root and shoot effects, and further show that grafting may be used to characterize genotypes for presence of the Dl-1 and Dl-2 alleles and to produce normal shoots in plants which otherwise would have been dwarfed. In the first experiment, two carriers each of Dl-1 and Dl-2 and a non-carrier were grafted reciprocally; of the 25 combinations, only the four combinations of Dl-2-carrying shoots with Dl-1-carrying roots produced dwarfing. In a second experiment, 11 carriers of Dl-1 and 9 of Dl-2 were grafted to Dl-1 roots and Dl-2 shoots. Although most graft combinations where Dl-1 roots were combined with Dl-2 shoots resulted in dwarfed plants, 10 genotypes showed mixed responses or otherwise did not respond as expected. Further experiments confirmed the occurrence of materials giving mixed responses or showing dwarfing irrespective of the graft combination. While grafting has utility as a screening tool, results should be interpreted with caution. There remains a need to determine whether the unexplained results reflect effects of additional alleles or genes affecting the dwarfing response. Grafting was used successfully to obtain normal F₁ shoots, and ultimately, to produce F₂ seed after two cycles of backcrossing (BC₂) for four crosses which otherwise would have given dwarfed plants. Segregation of dwarfed and normal plants in progeny rows of F₂ plants derived from the BC₂ populations showed 1:3 ratios, this being consistent with the reported inheritance.     

Genetic mapping of quantitative trait loci in rye (Secale cereale L.)

Using the marker information of 275 F₂ plants quantitative traits determining morphological and yield characters were studied analyzing F₃progenies grown in four different experiments at three sites. The map constructed contains 113 markers including the major dwarfing gene Ddw1 with an average distance of about 10 cM between adjacent markers. Of the 21 QTLs detected ten were found to map on chromosome 5RL in the region of Ddw1. Beside the expected effects on plant height and peduncle length that are most probably due to the presence of the major dwarfing gene, additional effects on yield characters and flowering time were discovered in that region which may be caused by pleiotropic effects of Ddw1. An additional supposed gene cluster consisting of four QTLs controlling flowering time and yield components was discovered in the centromere region of chromosome 2R. Further loci are distributed on chromosomes 1R (1), 4R (1) 6R (3) and 7R (1). The map positions of the quantitative trait loci detected in rye are discussed in relation to major genes or QTLs determining agronomically important traits in other cereals. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Genetical analysis of resistance to Mycosphaerella pinodes in pea seedlings

Summary In studies of the inheritance of resistance, pea seedlings of seven lines in which stems and leaves were both resistant to Mycosphaerella pinodes were crossed with a line in which they were both susceptible. With seven of the crosses resistance was dominant to susceptibility. When F₂ progenies of five crosses were inoculated on either stems or leaves independently, phenotypes segregated in a ratio of 3 resistant: 1 susceptible indicating that a single dominant gene controlled resistance. F₂ progenies of one other cross gave ratios with a better fit to 9 resistant: 7 susceptible indicating that two co-dominant genes controlled resistance. The F₂ progeny of another cross segregated in complex ratios indicating multigene resistance.When resistant lines JI 97 and JI 1089 were crossed with a susceptible line and leaves and stems of each F₂ plant were inoculated, resistance phenotypes segregated independently demonstrating that leaf and stem resistance were controlled by different genes. In two experiments where the F₂ progeny of the cross JI 97×JI 1089 were tested for stem and leaf resistance separately, both characters segregated in a ratio of 15 resistant:1 susceptible indicating that these two resistant lines contain two non-allelic genes for stem resistance (designated Rmp1 and Rmp2) and two for leaf resistance (designated Rmp3 and Rmp4). Evidence that the gene for leaf resistance in JI 1089 is located in linkage group 4 of Pisum sativum is presented.     

Variation and inheritance of leaf angle, and its association with spot blotch (Bipolaris sorokiniana) severity in wheat (Triticum aestivum)

One thousand four hundred and seven spring wheat germplasm lines belonging to Indian and CIMMYT wheat programs were evaluated for their leaf angle and resistance to spot blotch caused by Bipolaris sorokiniana during three consecutive crop seasons, 1994–95, 1995–96 and 1996–97.Disease severity was recorded at six different growth stages beginning from tillering to late milk stage. Three crosses (M 3109 × Sonalika, HP 1808 × K 9006 and HD 2662 × K 9006) were made between genotypes with erect and drooping leaves. M 3109 was resistant, Sonalika susceptible while the other three lines possessed moderate resistance to spot blotch. Individually threshed F₂ plants were used to advance the generations. Leaf angle and spot blotch resistance were recorded in parents, F₁, F₃, F₄and F₅ generations. Leaf erectness in F₁ generation showed partial dominance. Evaluation of F₃, F₄ and F₅ progenies(120–150) revealed that leaf angle was under the control of few genes that appeared to be close to three. Germplasm lines with erect and semi-erect leaves displayed lower spot blotch severity than those having drooping and semi-drooping leaves. Lines homozygous for erect leaf posture in F₃,F₄ and F₅ generations showed significantly lower mean AUDPC than those with drooping leaves. A positive correlation (0.58) between leaf angle and AUDPC further indicated a positive influence of leaf erectness on severity to spot blotch disease. This revised version was published online in August 2006 with corrections to the Cover Date.     

Development of Novel Brassica napus lines with canola quality and higher levels of oleic and linoleic acids derived from intergeneric hybrids between B. napus and Orychophragmus violaceus

Parents and their F₁ and F₂ progenies were evaluated for density of type IV glandular trichomes and resistance to Tetranychus urticae Koch during three stages of the plant growth. Results indicated significant differences between density of type IV glandular trichomes on F₁ and F₂ populations at different sampling times. As age of plant increased, density of type IV glandular trichomes and resistance to two-spotted spider mite increased. Density of type IV glandular trichomes on F₂ individuals showed a broad range of variability (0–42 trichomes/mm²). The estimated heritability was very high (>85%) for damage score and density of type IV glandular trichomes at different sampling times. Acylsugars content in Lycopersicon pennellii ‘‘LA2963’’ was more than two-folds high than those found in L. esculentum ‘‘Nandi’’, F₁ and F₂ populations, indicating that recessive gene(s) are responsible for the high acylsugar contents in L. pennellii ‘‘LA2963’’.     

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.     

Wheat Rht‐B1 Dwarfs Exhibit Better Photosynthetic Response to Water Deficit at Seedling Stage Compared to the Wild Type

Wheat reduced height (Rht) genes encode modified DELLA proteins, which are gibberellin insensitive, accumulate under stress, restrain growth and affect plant stress response. The seedling reaction to soil water deficit regarding leaf gas exchange and chlorophyll fluorescence was compared in near‐isogenic lines carrying the alleles Rht‐B1a (tall), Rht‐B1b (semi‐dwarfing) and Rht‐B1c (dwarfing) and was related to leaf water content and anatomy. Under drought, Rht‐B1c line was characterized by less decreased CO₂ assimilation, delayed non‐stomatal limitation of photosynthesis and higher instantaneous water use efficiency. The functional state of its photosynthetic apparatus was better preserved as evidenced by the less decreased actual quantum yield (Φ_PSII) and potential maximum quantum yield (F_v/F_m) of PSII, and the less increased quantum yield of non‐regulated energy dissipation (Φ_NO). Rht‐B1b line also tended to perform better than Rht‐B1a, but differences were less pronounced. Although the leaves of both dwarf lines were smaller, thicker and more pubescent, their water content was not higher in comparison with the tall line. Nevertheless, in Rht‐B1c, leaf thickness was less decreased and mesophyll cells were less shrunk under drought. The more effective performance of the photosynthetic machinery of dwarf lines under water deficit could be explained by a combination of morpho‐anatomical and metabolic characteristics.     

Inheritance of internode length,plant form,and annual habit in a cross of cabbage and broccoli (Brassica oleracea var. capitata L. and var. italica Plenck.)

Summary When an inbred line of cabbage, Brassica oleracea L. var. capitata L., was crossed with an inbred line of broccoli B. oleraceae var. italica, the F₁ progeny were vigorous late annuals. All F₁ × broccoli backcross plants and 92% of the 3260 F₂ plants were annuals, while 40% of the F₁ × cabbage backcross plants were biennials. Annual habit is thus dominant and controlled by more than a single gene. Number of days to bud appearance in annuals varied continuously, and was primarily additive in inheritance. F₁ data suggested partial dominance for lateness but this was not supported by the F₂. Internode length was also continuous in distribution and primarily additive in inheritance, but with some dominance for short internodes in the F₁. Cabbage head forming ability was recessive and multigenic, with 2% of the F₂ plants forming heads, of which none were of commercial type and about half bolted as annuals. There was a significant chi square association between biennial habit and tendency for cabbage head formation. Clasping habit of terminal leaves was recessive to open leaves, multigenic, and associated with both cabbage heading and biennial habit.Technical Paper 4836, Oregon Agricultural Experiment Station; from an M.S. thesis by the senior author.     

Improvement of large-seeded common bean cultivars under sustainable cropping systems in Spain

Approaches are needed to broaden the genetic base and improve earliness and yield potential of large-seeded beans under sustainable cropping systems. The objective of this research was to develop adapted dwarf bean populations having a commercial seed quality and yield suitable for the production in the South of Europe. The original base populations were produced from crosses between genotypes within each Mesoamerica, Nueva Granada and Peru races, and between Peru and Nueva Granada, and Mesoamerica and Nueva Granada races. Visual mass selection for plant performance was practised in the F₂ and F₃ generations. In the F₄ and F₅, single plants were harvested under two cropping systems (sole cropping and intercropping with maize). From F₄, selection was based on precocity, combined with seed yield and seed commercial type. The F_4:7 selected lines from each original population were compared with their parents and five checks at four environments and two cropping systems. Differences among environments, populations, parents and checks were observed for all traits. Under intercropping with maize, there was a 50% reduction in seed yield. Yield of parents and checks belonging to Andean South American races, intraracial (Nueva Granada × Nueva Granada) and interracial (Nueva Granada × Peru) populations, was higher than that of those of Middle American origin. Intraracial crosses within large-seeded Andean South American (Peru race) and Middle American gene pools (Mesoamerica race) did not produce lines yielding more than the highest yielding parent. Only two large-seeded lines selected from crosses between small- and large-seeded gene pools out-yielded the best parent and check cultivar.