Improving male fertility restoration of common wheat for Triticum timopheevii cytoplasm

The fertile pure line R3‐37 of common wheat with cytoplasm of Triticum timopheevii Zhuk. is an R‐line (restorer) that can restore the male fertility of A‐lines (male sterile lines) with T. timopheevii cytoplasm. In breeding hybrid wheat, the hybrid of the cross R3‐37/ Baimian3 was found to be completely male sterile, indicating that Baimian3 has some genes that are epistatic to the Rf genes in R3‐37. In order to elucidate the essence of this phenomenon, the male fertilities of the hybrids of 27 crosses including R3‐37 and/or Baimian3 were studied. The results show that inheritance of male fertility of the hybrid R3‐37/Baimian3 involves interactions among Rf alleles, male fertility‐inhibiting genes and genetic background. Although more than 70 different kinds of male sterile cytoplasm to common wheat have been discovered, the systems of hybrid wheat production based on male sterile cytoplasm are all the A‐line/R‐line type and all have similar problems of hybrid fertility restoration. This study confirmed that there is a new model (A‐line/R*‐line//R‐line) for producing hybrid wheat with high fertility restoration. In the new model, the completely male sterile hybrids of A‐line/R*‐line can act as common A‐line.     

Genetic analysis of fertility restoration against photoperiod-sensitive cytoplasmic male sterility in Triticum aestivum cv. Norin 61

Triticum aestivum cv. Norin 26 with the Aegilops crassa cytoplasm becomes almost completely male sterile when grown under a long-day condition (15 h of light or longer), but is highly male fertile under a short-day condition (14.5h or less). This type of male sterility is called photoperiod-sensitive cytoplasmic male sterility (PCMS). Genetic analyses were made of the fertility-restoring (Rf) genes effective against PCMS that are present in T. aestivum cv. Norin 61. Conventional and monosomic studies indicated that restoration of fertility is controlled by multiple Rf genes located on at least four chromosomes: 4 A, 1D, 3D and 5D. The genetic mechanism of fertility restoration by the genes of‘Norin 61’differs from the mechanisms reported for‘Chinese Spring’and a‘Norin 26’mutant line.     

Evaluation of three sources of pollen fertility restoration genes and natural outcrossing of cytoplasmic male-sterile wheat

Summary Fertility restoration genes in Triticum aestivum L. in Texas Restorer Composite (TRC), D6301, and four CIMMYT restorer lines were studied, and selection was made for higher fertility in TRC. Mean-while, outcrossing percentages of seed set for 27 spring habit cytoplasmic male sterile (cms) varieties were evaluated for 3 to 5 years at Davis. The winter-habit TRC material did not restore reasonably good fertility, and the response to selection for higher fertility seemed to be slow. This poor fertility could be partly due to its late winter growth habit causing flowering at a period of high temperature and low humidity at Davis. The highest F₁ fertility was 46.6% in the cross cms Ramona x TRC-6, and its F₂ segregated into the ratio of 15 fertile to 1 sterile, with fertility ranging from 3.2 to 100%. Suggested for its improvement was intensive selection in the original TRC material and in the segre-gating F₂ population, followed by intercrossing. D6301 has 2 fertility restoration genes with different strengths which restore fertility up to 45.2% when both genes are heterozygous. D6301 is quite likely heterogeneous for these genes. Four CIMMYT restorer lines, D7464, D7465, D7466, and D7467, had satisfactory F₁ fertility restoration after crossing with cms Ramona 50. In 1975, the fertilities of the F₁'s ranged from 71 to 85% and were over 90% in 1976. The F₂ population of the cross cms Ramona 50 × D7464 segregated into a ratio of 3 fertile to 1 sterile, indicating that D7464 has a single dominant gene for fertility restoration. The F₂'s of crosses cms Ramona 50 × D7465, cms Ramona 50 × D7466, and cms Ramona 50 × D7467 gave a ratio of 15 fertile to 1 sterile, indicating that two gene pairs in these three lines were responsible for the fertility restoration. The best of this group was D7467 which restored fertility fully after being crossed with cms Ramona 50 (T. timopheevi cytoplasm).The early-flowering cms male-sterile varieties had higher outcrossing rates (16 to 38%) than late varieties (6 to 30%) over a 5-year period. This was due to hot and dry weather during the late growing season as well as to the rarity of windborne pollen. In 1970, 1971, 1972, and 1976, the variation among varieties was rather great. Some of them such as Roque 66 and Bajio 67, had consistently high outcrossing rates. This outcrossing ability seemed to be inherited and probably associated with the open-flowering characteristics of each variety.     

Gametocidal properties of RH-531, RH-532, RH-2956, and RH-4667 on spring wheat (Triticum aestivum L.)

Summary Several chemicals were evaluated for their gametocidal properties to control pollination in hybrid wheat (Triticum aestivum L. em Thell.) seed production. In a preliminary study, four chemicals were applied to three cultivars of spring wheat at two application times. RH-532 was the most effective of the four gametocides, reducing fertility to a 0 10% level for all three cultivars.In a second experiment, RH-532 was applied to one cytoplasmic male-sterile and three normal spring wheats at four rates (0.56, 1.12, 2.24, and 4.48 kg/ha) and at two times of application. Alternate strips of pollinator were planted throughout the experiment. All treatments decreased fertility. plant height, and spike length of the three normal wheats. RH-532 did not inhibit spike emergence of two semidwarf cultivars or one of conventional height. Cultivars did not respond similarly to treatments.Yields of cross-pollinated seed were only 1 21% of the normal yield when fertility levels on treated plants reached 0 10%. Hybrid seed content of the harvested seed ranged from 2 55%. On the treated male-sterile line, yield was significantly reduced in comparison with the open-pollinated check. Inhibitory properties of this chemical suppressed the fertilization potential of the female as well as inducing male sterility.The differential response of genotypes to treatments indicates that cultivars should be screened for male and female sensitivity to this chemical. If differential sensitivity is found, breeding and selection to improve this trait may be possible. Different environmental conditions may alter the response of cultivars to this chemical, with less effect on the female structures.Published with the approval of the Director, North Dakota Agricultural Experiment Station, as journal article No 71a.     

Cytoplasmic male sterility in barley. III. Maintenance of sterility and restoration of fertility in the msml cytoplasm

Summary 58 varieties, strains or mutants of barley were tested in the msml cytoplasm in Finland, location ca. 61 N. Three (5%) were found to be partial restorers, the rest being maintainers of sterility. 31 Israeli strains of wild barley (ssp. spontaneum) were tested. About one third were maintainers of sterility, the majority partial restorers, and two strains were full restorers. There is probably polymorphism of the restorer gene in Israeli wild barley. Partial restoration displays an environmental response. The physiology of partial restoration is discussed.     

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).     

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.     

Inheritance of fertility restoration in sunflower (helianthus annuus L.)

Summary Fertility restoration in the cross between a cytoplasmic male sterile line, 2 cm 183, and the restorer line, BCZ 111, (both obtained from France) was dominant in F₁ and segregated in a 9:7 ratio in the F₂ generation and thus suggested the action of two independent, complementary dominant genes controlling restoration. The behaviour of F₃ families broadly confirmed the F₂ ratio. The reasons underlying this pattern of inheritance has been discussed and the genetic symbols rf ₁ rf ₁ rf₂ rf₂and Rf ₁ Rf ₁ Rf ₂ Rf ₂ have been suggested for the male sterile and the restorer parents respectively.     

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.     

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.     

Frequency and fertility restoration efficiency of Rf3 and Rf4 genes in Indian rice

Identification of new parental lines is crucial for developing ecology‐specific hybrids with ideal agronomic performance. We screened a total of 570 different ecology‐specific Indian rice varieties for the presence of fertility restorer genes, Rf3 and Rf4 using tightly linked markers DRRM Rf3‐10 and RM6100, respectively. Among these varieties, 13% carried Rf3Rf3/Rf4Rf4, 31% carried rf3rf3/rf4rf4, 6% carried Rf3Rf3/rf4rf4 and remaining 50% carried Rf4Rf4/rf3rf3 allelic combinations. A mini set of 40 varieties with variable allelic combinations of fertility restorer genes were testcrossed with WA and Kalinga‐based CMS lines. All the 80 F₁s were evaluated for spikelet fertility and fertility restoration ability. Rf3Rf3/rf4rf4 genotypes mostly behaved as partial maintainers or partial restorers. In contrast, rf3rf3/Rf4Rf4 genotypes were partial or effective restorers. However, double dominant genotypes showed better fertility restoration than the genotypes containing Rf3 or Rf4 individually. Some of the genotypes showed unexpected restoration pattern implying occurrence of other fertility restorer(s) apart from Rf3 and Rf4. The perfect restorers and maintainers identified in this study can be directly used in hybrid rice breeding.     

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.     

Inheritance of leaf angle in Triticum aestivum L.

Summary A 6×6 diallel was prepared to study the inheritance of leaf angle in T. aestivum L. Genetic analysis in terms of diallel cross parameters and graphic analysis indicated the control of additive gene effects in the expression of this character. The results of F₁ analysis were supported by the analysis of F₂ data.     

Heterotic effects of wheat-rye chromosomal translocations on agronomic traits of hybrid wheat (Triticum aestivum L.) under an adequate moisture regime

Translocated chromosomes T1BL⋅1RS and T1AL⋅1RS have been widely used in many wheat (Triticum aestivum L.) breeding programs to develop high yielding cultivars. The objective of this study was to evaluate the heterotic effects of T1BL⋅1RS + T1AL⋅1RS, T1BL⋅1RS, and T1AL⋅1RS on yield and yield components of hybrid wheat grown under adequate moisture regimes. Thirteen hybrid wheats and seven parents with different chromosome constitutions relative to T1AL⋅1RS and T1BL⋅1RS were evaluated in a randomized complete block design. Variable performance was observed among the hybrids tested. Two of the three hybrids with T1BL⋅1RS + T1AL⋅1RS, produced 25.26% and 44.64% more grain than the hybrids with only T1BL⋅1RS. This was due to increased biomass, harvest index (HI) and spike density. However, the combination of these two translocations resulted in reduced kernels/spike, spikelets/spike and spike length compared to the T1BL⋅1R Stranslocation alone. When comparing closely related parents, the parent with T1AL⋅1RS produced 23.51% more grain yield than the non-translocated parent. The presence of T1AL⋅1RS resulted in 10.37% heterotic advantage for yield due to increased biomass, KW, and spike density. When the two wheat-rye translocated chromosomes are present in the same hybrid, T1AL⋅1RS seems to have a positive effect on yield through spike density and HI, but masks the effects of T1BL⋅1RS for some agronomic traits. This revised version was published online in August 2006 with corrections to the Cover Date.     

Partial asynapsis involving specific chromosomes in intervarietal hybrids of Triticum aestivum L.

Summary Metaphase I chromosome association of the monosomic F1 and the backcross progenies made to develop a monosomic line in the Spanish common wheat Pané-247 was analyzed using a Giemsa C-banding technique. This permits the unequivocal identification of nine meiotic chromosomes (4A, 7A and the seven chromosomes of the B genome). The average frequencies of pairing per arm and of univalents for these nine pairs per arm and of univalents for these nine pairs indicate a difference between arms. The F1 showed asynapsis with univalents in 18.5 per cent of PMC's in intervarietal hybrids. This mainly involved chromosomes 4A, 1B and 6B which also have the largest amount of constitutive heterochromatin. The possible causes of reduced metaphase I association and its rapid decrease during backcrossing are discussed in relation to polymorphism between heterozygous homologous chromosomes.     

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.     

An analysis of genes for resistance against Indian stem rust races in two bread wheat cultivars

Summary The genetic constitution of two bread wheat accessions from the International Spring Wheat Rust Nurseries (E 5883 and E 6032) has been studied for reaction to four Indian races of stem rust. Analysis of E 5883 has revealed that for each of the races 15C, 21 and 40 a single dominant gene operates for resistance. The dominant gene against race 15C was identified as Sr6. The dominant genes for resistance against races 21 and 40 were found to be different from the genes described so far. Resistance against race 122 is controlled by a single recessive gene producing characteristically a 2 type of reaction. This gene was identified as Sr8.The resistance of E 6032 against each of the races 15C, 21 and 40 is controlled by two genes, one dominant and one recessive, which act independently. Dominant genes effective against 15C, 21 and 40 were conclusively identified as Sr6, Sr5 and Sr9b, respectively. From the correlated behaviour against races 15C and 40 as well as from the phenotypes of the resistance reactions rhe same recessive gene, undescribed so far, operates against the two races. The second recessive gene operating against race 21 was also observed to be different from those so far designated. E 6032 was, however, found to be susceptible to races 122.The presence of Sr6 both in E 5883 and E 6032 against race 15C was further confirmed through F₂ and F₃ segregation data.     

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.     

Boron deficiency induced male sterility in wheat (Triticum aestivum L.) and implications for plant breeding

Boron (B) deficiency causes grain set in wheat to fail. A wide range of genotypic variation in the response to low B has been observed. Genotypes were screened in low B in soil and sand culture, and classified into five groups, namely, very sensitive, sensitive, moderately sensitive, moderately tolerant and tolerant. At very low levels of B, the very sensitive to sensitive genotypes were completely male sterile and set only a few or no grain, while the tolerant genotypes set grain normally. Natural outcrossing was detected in these male sterile plants when a tolerant genotype was growing nearby. Grain set by cross fertilisation was markedly enhanced by a B application directly on the ear of the male sterile plants. Three practical implications are suggested. Firstly, genotypes that are tolerant to low B can provide a solution for grain set failure caused by B deficiency. Secondly, the potential for outcrossing in male sterile B deficient wheat has to be taken into account in the maintenance of pure lines in low B soils even though wheat is normally self pollinated. Thirdly, a simple and novel method for hybridization is suggested, in which B deficiency is used as fertility selective medium and male sterile female parents and fertile male parents are provided by genotypic variation in the response to low B. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.)