In vitro techniques for selecting wheat (Triticum aestivum L.) for Fusarium-resistance. II. Culture filtrate technique and inheritance of Fusarium-resistance in the somaclones

Summary Calli of resistant, intermediary and susceptible wheat (Triticum aestivum L.) varieties were selected using culture filtrates of Fusarium graminearum and F. culmorum and the regenerants were evaluated for resistance up to R₃. Czapek-Dox broth medium was inoculated with mycelia of Fusarium isolates and incubated for 2–6 weeks. Filtrates were added to MS callus growing medium, then 5 weeks-old calli were transferred onto this medium (MST) for 4–5 weeks. MST containing 30% filtrate was found to be suitable for selection. Resistant calli were transferred again to fresh MST for further two selection cycles. The surviving calli produced less fertile regenerated lines (R₀) than the non-selected ones. Among 18 R₁ lines tested for Fusarium-resistance in the seedling stage by artificial inoculation in the greenhouse, two (11.1%) were significantly more resistant, one (5.6%) was more susceptible than the original cultivar and the rest (83.3%) behaved similarly to the donor plants. Among unselected R₃ lines of three varieties, practically the same number of resistant plants were found as among the related selected ones. When the R₃ selfed generations obtained through double-layer and culture filtrate selection techniques were tested for Fusarium-resistance, 35.7% of the lines were found to be more resistant than the original cultivars, none was more susceptible and 64.3% had a reaction similar to that of the source materials. Thus, inheritance of the disease reaction was not stable in all cases. Success of in vitro selection for Fusarium-resistance depended also on the genotype, and toxin analysis showed that although being effective, the selective media contained deoxynivalenol only exceptionally. In selecting wheat for Fusarium-resistance in vitro, the culture filtrate technique proved better than the double-layer procedure.     

Combining abilities of in vitro traits in wheat (Triticum aestivum L.) immature embryo cultures

Summary Combining ability for six in vitro culture traits in wheat were studied in a 8×8 diallel cross (excluding reciprocals). Specific combining ability effects (sca) were significant for all six traits derived from immature embryos on two media protocols, whereas general combining ability (gca) variances were significant only for five of them. Furthermore, based on ratios obtained by comparing the ratio of K² gca to K² sca, sca was more important than gca for all six traits. Genetic correlations between shoot formation and other in vitro traits, except callus weight and root formation, were higher in magnitude than the corresponding phenotypic correlations estimates, indicating the importance of genetic effects.     

Genetic variation for nitrogen use efficiency in winter wheat (Triticum aestivum L.)

Summary The new European Common Agricultural Policy and environmental considerations are certainly to change agricultural practices toward low input cultivation systems. Nitrogen is one of the main inputs of winter wheat in northern France and it contributes highly to phreatic water pollution. A research programme has then been set up in order to study whether it is possible to breed for winter wheat cultivars using more efficiently N fertilisers. Less nitrogen would be applied, decreasing pollution risks and operational costs. It has been shown that a large variation exists for N related traits and for the resistance against N deficiency. On the one hand the cv Arche is very resistant to N deficiency, its yield on low N conditions (with no N fertiliser) is on average 89% of its yield on high N conditions (with a high N application). On the other hand, cv Récital is very susceptible to N deficiency as this same percentage is only 61%. A study on 10 hybrids showed that heterosis for grain yield was higher at low N level than at high N level. This was due to a higher number of grains per m².     

Genetic control of frost tolerance in wheat (Triticum aestivum L.)

Summary The frost tolerance of winter wheat is one component of winter hardiness. If seedlings are frost resistant, it means that they can survive the frost effect without any considerable damage. To study the genetic control of frost tolerance, an artificial freezing test was used. Frost tolerance is controlled by an additive-dominance system. The results of diallel analyses indicate the importance of both additive and non-additive gene action in the inheritance of this character. The dominant genes act in the direction of lower frost tolerance and the recessive genes in the direction of a higher level of frost tolerance. The results of monosomic and substitution analyses show that at least 10 of the 21 pairs of chromosomes are involved in the control of frost tolerance and winter hardiness. Chromosomes 5A and 5D have been implicated most frequently. The geneFr1 (Frost 1) was located on the long arm of chromosome 5A. Crosses between cultivars, chromosome manipulation and the induction of somaclonal variation may be suitable methods for broadening the gene pool for frost tolerance.     

Physiological studies on grass-dwarf selection lines in wheat (Triticum aestivum L.)

Summary Low power heating wires insulated in a flexible plastic strip were used to heat the shoot meristematic region of grass-dwarf genotypes to a precise temperature. The results indicate that the shoot apical meristem is the region requiring 26°C for the initiation of reproductive development in these genotypes. A secondary effect of the reproductive growth induced by high temperature treatments, was a reduction in the high levels of peroxidase enzymes found in vegetative grass-dwarf plants. The heating wire provides a precise method to identify Type I, II and III grass-dwarf genotypes according to their temperature requirements, of 26°C, 21°C and 16°C respectively, for reproductive growth under the same temperature (16 C) and photoperiod (12 hours).     

Variation in pigment content of flour color in bread wheat (Triticum aestivum L.)

Summary Variation in pigment content of the flour of bread wheats (Triticum aestivum L.) was studied in the progenies of F₁ and F₂ of three crosses and their reciprocals. Reciprocal differences in pigment content were observed in the F₁ and F₂ means. Low pigment content was found to be partially dominant or over dominant in the crosses studied. There was evidence of substantial mid-parent F₁ heterosis in all crosses and betterparent F₁ heterosis in three crosses. In the F₂, heritability estimates were moderate to high. The F₂ frequency distributions were not normal. Estimation of effective factor pairs indicated the presence of one or two major gene pairs involved in the expression of pigment content in the flour. Action of modifiers was also assumed in one cross and its reciprocal. A factorial approach to metrical character suggested that the F₂ segregation ratios of low pigment content to high pigment content were 3:1, 15:1, 13:3 and 9:7 for the different crosses. Utilization of the findings in a wheat breeding program is briefly discussed.     

Environmental and genetic variation for protein content in winter wheat (Triticum aestivum L.)

Summary There is a considerable amount of variation in the protein content of wheat kernels. This variation may be induced by environmental factors, but can also be attributed to genetic differences. Within a genotype the correlation between grain yield and grain protein content can be either close to zero, positive, or negative. depending on the fertility level. Between genotypes this correlation is strongly negative. It is argued that the negative inter-genotypic correlation is largely a consequence of the high harvest-index of high yielding varieties. Breeding methods to alleviate the negative yield-protein content relationship are given.     

Relationship between common wheat (Triticum aestivum L.) gluten proteins and dough rheological properties

This paper reports the correlation between the rheological properties of bread wheat dough and the types and quantities of endosperm proteins in 28 common wheat cultivars. Different methods were used to analyse the allelic composition of these cultivars and the relative quantities of the different proteins contributing to the gluten structure. Neither dough strength (W) nor tenacity/extensibility (P/L) correlated with allelic composition. Different wheats with the same allelic composition (i.e., with respect to glutenins) showed different rheological properties. The glutenins were the most influential components of W and P/L, especially the high molecular weight (HMW) glutenin subunits and in particular the type x form. These proteins seem to increase W and are the main constituents of the gluten network. The gliadins and low molecular weight (LMW) glutenin subunits appear to act as a “solvent”, and thus modify the rheological properties of the dough by either interfering with the polymerisation of the HMW glutenin subunits, or by altering the relative amounts of the different types of glutenin available. Thus, the protein subunits coded for by the alleles Glu-B1x7 and Glu-D1x5 stabilised the gluten network, whereas those coded for by Glu-B1x17 and Glu-D1x2 had the opposite effect. Dough properties therefore appear to depend on the glutenin/gliadins balance, and on the ratio of the type x and type y HMW proteins. The influence of external factors seems to depend on the allelic composition of each cultivar.     

Stability analysis over seasons and locations of multilines of wheat (Triticum aestivum L.)

Summary Three multilines each of Kalyansona and PV 18 varieties of wheat were compared for their stability of yield and agronomic characters and disease resistance against the respective recurrent parents. The experiments were conducted for four years at nine locations.The multilines had more tillers and bolder seeds than the recurrent parents. There were, however, no differences for plant height, ear length and number of spikelets per spike.The genotypes x years x locations interaction was much more important than genotypes x years or genotypes x locations interaction as well as the main effects for genotypes. It is suggested that the number of test locations should be increased while decreasing the number of years.The stability parameters indicated that the multilines had a higher mean yield in the case of Kalyansona multilines whereas the yield of multilines of PV 18 was not significantly inferior to that of the recurrent parent. The regression coefficients were very close to unity except for KSML 3 (b=1.132). The deviations from regression were much larger for the multilines than for the recurrent parents.For rust resistance the multilines were superior over the varieties Kalyansona and PV 18.This is publication No. 9 of the series Studies on multilines in wheat (Triticum aestivum L.)     

A recessive resistance gene for yellow rust (Puccinia striiformis West.) in bread wheat (Triticum aestivum L.)

Summary Three lines derived from the old dirty Dutch land variety Gelderse Ris were resistant against race 66(70)EO(16) of yellow rust. It was found that this resistance was conditioned by one recessive gene provisionally coded yrGR.     

Genes Lr48 and Lr49 for hypersensitive adult plant leaf rust resistance in wheat (Triticum aestivum L.)

The genetic bases of leaf rust resistance in wheat (Triticum aestivum L.) line CSP44, selected from the Australian cultivar Condor, and Indian cultivar VL404, were studied. The reaction patterns of CSP44 and VL404 against Indian races 12, 77, 77-1, 77-2, 77-3, 77-4, 77-5 and 108 were different from reaction patterns shown by near-isogenic lines with known adult plant resistance (APR) genes, viz. Lr12, Lr13, Lr22b and Lr34. Although the reaction patterns of CSP44 and VL404 were similar to the near-isogenic line Tc+Lr22a, tests of allelism indicated absence of Lr22a in both CSP44 and VL404. On the basis of genetic studies, their resistances in field tests against race 77-5, the most virulent race from the Indian sub-continent, were each ascribed to two genes. One of the two genes in each wheat was identified to be the non-hypersensitive APR gene Lr34. The second APR genes in CSP44 and VL404 gave hypersensitive reaction types and were recessive and dominant, respectively. The gene in CSP44 was designated Lr48and the gene in VL404, Lr49. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic basis of grain filling rate in wheat (Triticum aestivum L. emend. Thell.)

Summary Grain filling rate in wheat (Triticum aestivum L. emend. Thell.) positively influences grain yield under a wide range of conditions. The effective utilization of this trait in breeding depends on an understanding of its genetic control. A study was, therefore, conducted to determine the genetic basis of grain filling rate in six crosses of wheat. Higher order genic interactions and/or linkage were important in the genetic regulation of grain filling rate (GFR) in the majority of crosses. Additive ([d]) and dominance ([h]) gene effects were important in the control of GFR in main ears (ME) and whole plant ears (WPE). Additive and additive × additive epistatic effects were the most important in the genetic control of GFR in last ears (LE). Location effects on genetic effects for GFR were significant (P < 0.05) in all ear types of some crosses except in ME. Genotype × environment interaction effects were important (P < 0.001) in LE and WPE.It was concluded that the inheritance of GFR is complex and is dependent on ear type. Breeding procedures that facilitate the exploitation of non-additive and additive gene effects were recommended for the genetic improvement of grain filling rate of wheat.     

Evidence for a ‘uniculm effect’ in spring wheat (Triticum aestivum L.) in a mediterranean environment

Summary The concept of the uniculm habit as an important feature of a wheat ideotype for a mediterranean environment was evaluated under field conditions. A uniculm plant produces a single shoot and when sown in a stand exemplifies a non-tillering crop with a fixed density of shoots throughout the growing season. Yield and harvest index of normal tillering spring wheat was compared with that of the same crop surgically detillered throughout the growing season to a constant density of 2 shoots per plant. The use of a biculm, whilst retaining the uniculm principle of a fixed density of shoots throughout the growing season, permitted comparison on a single crop sowing at normal field density.The control plots followed the usual pattern of tillering for the region attaining a maximum of about 4.0 shoots per plant by early spring. Shoot number declined to 2.3 and 2.6 per plant by maturity in 1978 and 1979, respectively.Detillered plots outyielded the controls by 14 per cent in 1978 (2.05 v. 1.80 tonnes ha^-1) and 22 per cent in 1979 (1.84 v. 1.51 tonnes ha^-1). Harvest indices were 0.39 v. 0.35 in 1978 and 0.30 v. 0.24 in 1979 for detillered and control plots, respectively.Higher leaf area indices and better water relations after anthesis in biculm stands indicated more efficient water use when shoot population was controlled at near optimum level. Irrigation at anthesis reduced the difference in yield between detillered and control plots.Although the uniculm principle was demonstrated with biculms in practice control of shoot number will require the use of genetic uniculms.     

Flooding tolerance of spelt (Triticum spelta L.) compared to wheat (Triticum aestivum L.) – A physiological and genetic approach

In marginal, agroclimatic zones, yield is often affected by flooding, but the effect is much less for winter spelt (Triticum spelta L.) than for winter wheat (Triticum aestivum L.). This study evaluates the reaction of a wheat x spelt population (F₅ RILs of Forno x Oberkulmer) to flooding stress in the early phase of germination. Lines with greater tolerance to 48 h flooding just after imbibition showed less electrolyte leakage (r = -0.79) indicating greater membrane integrity and better survival. Five QTL explaining 40.6% of the phenotypic variance for survival to flooding were found, and localized on the chromosomes 2B, 3B,5A, and 7S. The tolerance to 48 h flooding four days after sowing was best correlated with the mean germination time (r = 0.8), indicating that the plants with a fast coleoptile growth during flooding are less susceptible to flooding. Ten QTL were found for seedling growth index after flooding explaining 35.5% of the phenotypic variance. They were localized on chromosomes 2A, 2B, 2D, 3A, 4B, 5A, 5B, 6A, and 7S. Standard varieties of spelt and wheat showed the same tolerance characteristics. The possibility to use marker assisted selection for flooding tolerance is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of desiccation and a change in temperature on germination of immature grains of wheat (Triticum aestivum L.)

A study was made of the effects of desiccation and a change in temperature on the germination of wheat grains harvested 20 days after anthesis. When the germination test was performed immediately after harvesting, germination percentages ranged from 5.2% to 10.7%. Germination percentages increased to 48.2% to 90.3% after grain had been desiccated at 20 °C and then hydrated at 10 °C. This increase occurred even if grains had been desiccated in an atmosphere of high relative humidity. The germination percentage of non-desiccated grains depended on the germination temperature. When the pericarp and testa were removed from embryos, the germination percentage of grains incubated at 20 °C and then at 10 °C was higher than that of grains incubated at 10 °C. In general, a low germination temperature is believed to be effective in breaking dormancy of wheat grains. However, a change in temperature stimulated germination to a greater extent than a constant low temperature. The germination percentage of 5-day cycle alternating temperature was greater than that with 1-day, 2-day and 10-day cycles. Although the germination of immature wheat grains was stimulated by both high and low germination temperatures, it is likely that cycles shorter and, also, longer than a critical period induce limited germination. Loss of dormancy commonly occurs when development of wheat grains proceeds at a high temperature, with imbibition at a low temperature. However, germination ability of non-desiccated immature wheat grain was enhanced by a change in temperature during germination. This revised version was published online in August 2006 with corrections to the Cover Date.     

Studies on somaclonal variants for resistance to scab in bread wheat (Triticum aestivum L.) through in vitro selection for tolerance to deoxynivalenol

This study was conducted to develop an efficient in vitro selection system for scab resistance by using in vitro screening for tolerance to deoxynivalenol (DON). Immature embryos of two wheat varieties, a scab-resistant variety Sumai 3 and a susceptible variety Mianyang 11, and their reciprocal F₁ hybrids were cultured on MS medium supplemented with 2,4-D 2 mg/l and 0.6 × 10^-4 M DON for callus induction. The responses of callus induction and plant regeneration to 0.6 × 10^-4 M DON differed significantly between resistant and susceptible varieties, according to observed scab resistance levels at the plant level in the field. The percentage of callus formation of resistant variety Sumai 3 on induction medium containing DON was higher than that of susceptible variety Mianyang 11. Regeneration of DON-tolerant calli on DON-containing differentiation medium differed significantly between Sumai 3 and Mianyang 11. Averaged across the DON-tolerant calli of two varieties and their reciprocals, regeneration of DON-tolerant calli was decreased 3-fold on DON-containing medium. By an inoculation test with conidiospores of Fusarium graminearum Schw, we obtained several resistant lines from progenies of regenerated plants from DON-tolerant calli. These somaclonal lines had lower disease scoring (reaction index, infected spikelets and disease incidence), shorter plants and better yield components than Sumai 3, a famous Chinese resistant variety. This revised version was published online in July 2006 with corrections to the Cover Date.     

Aneuploid analysis for protein content and tyrosinase activity in hexaploid wheat (Triticum aestivum L. em. Thell.)

Summary A study was conducted to locate the genes responsible for the determination of kernel protein content and tyrosinase activity in a hexaploid wheat variety UP 301 using Pb. C591 monosomic series. Genes located on chromosomes 4B, 5B, 6B, 7B, 3D and 7D of UP 301 controlled protein content of UP 301. Of these the B genome chromosomes were found to have genes for increased protein content while the D genome chromosomes were found to carry genes for low protein content. A major gene coding for tyrosinase enzyme was detected on chromosome 6B of UP 301 and a modifier on chromosome 5B. This indicated the possibility of improving these quality characters through chromosome manipulation.     

Expression of 17 rye (Secale cereale L.) traits in a range of durum wheat (Triticum durum Desf.) and bread wheat (T. aestivum L.) genetic backgrounds

Summary Expression of 17 rye traits in 24 bread wheat x rye and 8 durum wheat x rye crosses was studied, using a self-compatible, homozygous, dwarf rye. Rye showed epistasis for hairiness on the peduncle in all the crosses of Triticum aestivum and T. durum wheats with rye. Dark greenness of leaves of rye was expressed in all the durum wheat x rye and in some of the bread wheat x rye crosses. Similarly, absence of auricle pubescence, a rye trait, was expressed in most of the durum wheat x rye crosses but not in the bread wheat x rye crosses, indicating the presence of inhibitors for these traits frequently on the D genome and rarely on the A and/or B genome of wheat. Most of the wide hybrids resembled rye fully or partially for intense waxy bloom on the leaf-sheath and for the absence of basal underdeveloped spikelets. Similarly, most of the amphihaploids resembled rye for the anthocyanin in the coleoptile, stem and node. The presence of some inhibitors on A and/or B genome of wheat was indicated in some of the wheat genotypes for the expression of rye traits viz. intense waxy bloom, anthocyanin in node and absence of basal underdeveloped spikelets. Enhancement in the level of expression of the intensity and length of bristles on the mid-rib of the glume of the hybrids might be due to wheat-rye interaction. Less number of florets/spikelet as in rye showed variable expression in different wheat backgrounds. Some other rye traits like absence of auricles, terminal spikelet and glume-awn were not expressed in the wheat background. The expression of some of the rye genes might have been influenced by their interaction with Triticum cytoplasm and/or the environment.     

Chromosomal regions associated with green plant regeneration in wheat (Triticum aestivum L.) anther culture

Genetic capacity for green plant regeneration in anther culture were mapped in a population comprising 50 doubled haploid lines from a cross between two wheat varieties ‘Ciano’ and ‘Walter’ with widely different capacity for green plant regeneration. Bulked segregant analysis with AFLP markers and composite interval mapping detected four QTLs for green plant percentage on chromosomes 2AL (QGpp.kvl-2A), 2BL (QGpp.kvl-2B.1 and QGpp.kvl-2B.2) and 5BL (QGpp.kvl-5B).The three QTLs detected on chromosome 2AL and 2BL all derived their alleles favouring green plant formation from the responsive parent ‘Ciano’.The remaining QTL on chromosome 5BL had the allele favouring green plants from the low responding parent ‘Walter’. In a multiple regression analysis the four QTLs could explain a total of 80% of the genotypic variation for green plant percentage. None of the chromosomal regions with QTLs for green plant percentage showed significant influence on either embryo formation or regeneration frequencies from the anther culture. The three major QTLs located on group two chromosomes were fixed in a second DH population derived from two parents ‘Ciano’ and ‘Benoist’,both with high capacity to produce green plants. A QTL explaining31.5% of the genetic variation for green plant formation were detected on chromosome 5BL in this cross as well. This revised version was published online in July 2006 with corrections to the Cover Date.     

Major gene control of tolerance of bread wheat (Triticum aestivum L.) to high concentrations of soil boron

Summary The genetic control of tolerance of wheat to high concentrations of soil boron was studied for five genotypes. Each genotype represented one of five categories of response to high levels of boron, ranging from very sensitive to tolerant. Tolerance to boron was expressed as a partially dominant character, although the response of an F₁ hybrid, relative to the parents, varied with the level of boron applied. The F₁ hybrids responded similarly to the more tolerant parent at low B treatments and intermediate to the parents at higher treatments. Ratios consistent with monogenic segregation were observed for the F₂ and F₃ generations for the combinations (WI^*MMC) × Kenya Farmer, Warigal × (WI^*MMC) and Halberd × Warigal. The three genes, Bo1, Bo2 and Bo3, while transgressive segregation between two tolerant genotypes, G61450 and Halberd, suggested a fourth locus controlling tolerance to boron.