QTL alleles from a winter feed type can improve malting quality in barley

Mapping quantitative trait loci (QTLs) responsible for malting quality traits in barley populations has been the main genetic approach to malting quality breeding. A ‘winter × spring’ doubled haploid barley population ‘Nure’ × ‘Tremois’, where such traits were segregating, has been recently developed. Our objective was to map QTLs for malting quality from 2 years of trials in two contrasting locations. QTLs were found on six chromosomes, with a main cluster on chromosome 1H. For wort viscosity and malt extract, favourable alleles at two loci on chromosome 5H were carried by the winter feeding parent ‘Nure’. Doubled‐haploids with higher quality than the spring malting cultivar ‘Tremois’ showed either a facultative or a winter growth habit and a level of frost tolerance comparable with that of the winter tolerant parent ‘Nure’. Markers and QTLs of quality traits were further validated on a separate set of DH lines, coming from the same cross, by means of marker‐assisted selection. This showed that, at least in the present cross combination, positive contributions to malting quality can be found in winter feed barley.     

Evaluation of Czech spring malting barleys with respect to the β-amylase allele incidence

Strong enzymatic activities in the germinating barley grain, together with protein and starch content, are crucial for high extraction values in the resulting malt and, therefore, barley malting quality. The efficient characterization of registered barley cultivars and genetic resources with respect to one of the relevant thermostability enzymes (β‐amylase) is an essential requirement. The template‐directed dye‐terminator incorporation (TDI) assay: based on flourescence resonance energy transfer (TDI‐FRET) ( Chen et al. 1995 ) was used to detect single nucleotide polymorphisms (SNPs) in the β‐amylase coding sequence resulting in low (Sd2L), intermediate (Sd1) and high (Sd2H and Sd3) thermostability enzyme across 84 Czech barley cultivars and genetic resources used over a period of time in the Czech Republic. The incidence of different alleles has changed during the last 100 years. Also the new resources with high thermostabile β‐amylase were identified. They can be used effectively to breed for malting quality improvement.     

Evaluation of methods for the assessment of malting quality in barley breeding

Summary Methods for assessing malting quality in barley breeding were evaluated for their precision, including variation between different batches, analysts and barley samples. The barley characters measured were grain moisture, ground grain moisture (two methods), grain weight, steep moisture, malt moisture (two methods), malt yield, malt nitrogen, malt soluble nitrogen, Kolbach index, malt diastatic power, malt -amylase, malt -glucanase, wort refractive index, hot water extract, extract yield, wort reducing sugars, wort total carbohydrates, wort -amino nitrogen and wort total nitrogen. The value of these measurements in barley breeding is discussed.     

Androgenic capability among genotypes of winter and spring barley

Variable efficiency of androgenesis remains a serious problem in many species of cereals. It is still unclear what makes certain genotypes more amenable to androgenesis than others. This study was undertaken to quantify the previously suspected advantage of winter barley genotypes over spring ones with regard to regeneration efficiency in anther culture. The material consisted of 40 barley hybrids originating from Polish breeding companies. The number of androgenic structures per 100 anthers did not differ significantly between analysed groups (119 vs. 152 non‐significant), but the average regeneration of green plants per 100 anthers was five times higher in winter genotypes (6.4 vs. 1.3). The incidence of albinism was lower for the winter than for the spring materials (70% vs. 90%), while the rate of spontaneous chromosome doubling was similar in both groups (58% vs. 56%). The results strongly support the notion that winter genotypes are more amenable to androgenesis and this may be a consequence of their better adaptation to stress conditions.     

Relationship between vernalization requirement and winter hardiness in doubled haploids of barley

Summary Progeny lines of chromosome-doubled haploids from crosses between one winter and four spring barley varieties were analysed for winter hardiness and vernalization requirement in the field. About one quarter or less of the offspring lines required vernalization under field conditions showing that winter versus spring habit of growth is controlled by at least two pairs of genes. The vernalization requirement could be measured quantitatively by testing a few plants in a greenhouse after artificial cold treatment during germination. The individual winter lines tested required from two to seven weeks of vernalization in order to flower as early as possible. The cold treatment had no effect on the time of flowering in the spring lines.Winter hardiness varied from zero to 100% plant survival in the field. The majority of the lines requiring vernalization survived better than 90% and the survival of the rest was at least 60%. The winter survival of the spring lines varied from zero per cent (about one third of the lines) to 100%. Hence, it is possible to obtain lines of good winter hardiness without requirements for cold treatment. Conversely, a selection for vernalization requirement has a positive effect on winter hardiness.     

Agronomical and morphological differentiation among winter and spring triticales

A collection of 299 secondary hexaploid triticale cultivars and advanced breeding lines from 18 countries, which were considered a representative sample of the existing diversity, was evaluated for morphological and agronomical characters with autumn planting at Lleida, Spain, from 1988 to 1991. The entries were classified as having winter (84) or spring (215) growth habit and among this latter group were complete (73) or substituted (147) types according to CIMMYT's terminology. Winter and spring triticales were grouped by cluster and principal component analyses. Winter triticales were taller with longer growth cycles, longer spikes, and more spikelets per spike than spring types. At early stages they also had prostrate growth. Spring-substituted types were separated from complete material. As a group, spring-substituted triticales differed more from winter types than the spring complete genotypes, which showed intermediate characteristics. Complete types of spring habit had tendency to be taller, with longer spikes, more spikelets per spike and bigger and heavier grains than substituted triticales. Greater variation in morphological and agronomical parameters was detected among winter triticales followed by the complete-spring group.     

Genetics of androgenesis in winter and spring wheat genotypes

Androgenetic response and its inheritance was studied in nine winter type and two spring type wheat genotypes by crossing them in line × tester fashion. The 18 F₁ and 11 parental genotypes were raised in completely randomized block design with three replications and subjected to anther culture. Data were recorded with respect to per cent calli induction, percent regeneration and per cent green plant development. Analysis of variance revealed considerable genetic variation for all the three component traits of androgenesis. Both additive and non-additive genetic variances were involved in the androgenetic response with the preponderance of non-additive gene action in case of calli induction and additive gene action for regeneration and green plant development. Low estimates of heritability (narrow sense) for calli induction and moderate for regeneration and green plantlet development were obtained. The genotypes, Henies VII, Pnfjoumee, Saptdhara, Sentry, Purthi of winter wheat and HPW147 of spring wheat were identified as good general combiners for almost all the androgenetic parameters and have potential use in future haploidy breeding programmes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diallel analysis of net blotch resistance in doubled haploid lines of Nordic spring barleys

A half-diallel was made between five six-rowed Nordic spring barleys to study the genetics of resistance to net blotch. Twenty-five doubled-haploid (DH) lines from each cross and the parents were sown in hill plots in Finland in 1997 and 1998. The plots were artificially inoculated with Pyrenophora teres Drechs. f. teres Smedeg. and assessed for resistance to net blotch. There were statistically significant differences in resistance of the five parents to net blotch. General combining ability (GCA) of the parents and specific combining ability (SCA) effects in the progeny were statistically significant in both years, but GCA effects predominated. Evidence for additive epistasis was minimal. Progeny of a particular cross were less resistant to net blotch than the better parent. The most resistant progeny were derived from the cross between the two most resistant parents, Pohto and WW7977, and resistance was governed by at least eleven effective factors. Narrow sense heritability estimates for resistance to net blotch were high during both years (0.84–0.99). It appears that net blotch resistance of progeny from crosses can be largely predicted from reactions of the parents. Quantitative resistance to net blotch can be further advanced by identification and incorporation of superior parents, from a screening such as reported here, into a recurrent selection breeding programme. This revised version was published online in August 2006 with corrections to the Cover Date.     

Agronomic performance of crosses between Nordic and exotic barleys

Exotic germplasm may be useful for the improvement of agronomic performance of barley breeding material. This study was conducted in order 1) to evaluate if it is possible to improve performance of Nordic barley breeding material by utilizing exotic germplasm sources (unadapted landraces and H. spontaneum), 2) to evaluate incorporation of exotic germplasm into a genetic base widely adapted to the agro-ecological conditions of the extreme Scandinavian north as well as to the base adapted to the conditions of the south of this geographical region, and 3) to evaluate whether an index composed of several traits corresponded to 'breeder's phenotypic evaluation' of the progenies. Earliness, straw length, number of ears/plant, yield/plant, seed shattering, persistence of awns and an index computed from these traits were studied in Swedish and Finnish 4-way crosses along with their adapted parents. Most of the crosses headed earlier than their adapted parents and none of the crosses outyielded the adapted parents. The most favorable frequency distributions for the index were found in the crosses with only cultivated barleys as parents. Transgressive segregates were also found in progenies with H. spontaneum parents. Incorporation of exotic barleys was most successful into the local genetic base. Comparisons between indices and breeder's phenotypic evaluation suggested reconsideration of evaluation methods early in parent development.     

Distribution and allelic expressivity of genes for hybrid necrosis in some elite winter and spring wheat ecotypes

The distribution and allelic expressivity of hybrid necrosis genes (Ne ₁ and Ne ₂) were studied in 21 winter (mostly exotic) and 43 spring type elite wheat genotypes, by crossing them with two known testers, C 306 (Ne ₁-carrier) and HD 2380 (Ne ₂-carrier).Ne ₁ gene was present in one north-west Himalayan winter wheat landrace, Shoure Local, but absent in the other winter as well as spring wheats. Ne ₂ gene was prevalent to a much lower extent in the exotic winter wheat germplasm (31.57%) as compared to the recently developed Indian and Mexican spring wheat semidwarfs (69.80%). This may suggest that breeders have tried to preclude hybrid necrosis by selecting for non-carrier genotypes in the development of exotic winter wheats in contrast to the situation in spring wheats. Based on the degree of expression of hybrid necrosis genes in the F₁ hybrids, the carrier genotypes were characterized with respect to the allelic strength of the hybrid necrosis genes. The 27 non-carrier genotypes of the two ecotypes identified in the present study have a greater potential use in future hybridization programmes so as to overcome the problem of hybrid necrosis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding and brewing quality of the Canadian malting barley variety ‘CDC Goldstar’ lacking lipoxygenase-1

Various types of malt quality profiles have been investigated to benefit the North American brewing industry. Herein, we report the development and brewing quality of the hulled, two-row malting barley (Hordeum vulgare L.) variety ‘CDC Goldstar’ lacking lipoxygenase-1 (LOX-1-less). This new variety offers a novel malt type for the improvement of beer flavor stability. The agronomic performance of ‘CDC Goldstar’ was tested in the Western Cooperative Two Row Barley Registration Trials during 2013–2014. In addition to high lodging tolerance, the new variety showed 6% higher yield than the current leading variety ‘CDC Copeland’. The malt quality of ‘CDC Goldstar’ showed higher diastatic power and lower wort β-glucan content than ‘CDC Copeland’ and controllable proteolytic modification (soluble nitrogen and Kolbach Index). Pilot- (100 L) and commercial-scale (5,000 L) brewing trials were conducted using ‘CDC PlatinumStar’, another LOX-1-less variety with a low enzymatic profile, as the control variety. Absence of the LOX-1 trait from ‘CDC Goldstar’ maintained trans-2-nonenal levels in aged beers as low as those in other LOX-1-less varieties without affecting major beer parameters, such as ester and aldehyde content or foam stability. The newly developed ‘CDC Goldstar’ malting barley provides added value for the beer industry and consumers.     

Prediction of grain protein in spring malting barley grown in northern Europe

The ability to predict grain protein concentration at harvest (CP) in Swedish malting barley (Hordeum distichum) from observations of soil conditions, sowing day, fertilisation rate, remote sensing at early stem elongation and the temperature sum during grain filling, was tested for two cultivars; Astoria (Secobra, France) and Wikingett (Svalöf-Weibulls, Sweden) in 16 fertilisation trials in southern Sweden, encompassing 3 years (2001–2003). Fertilisation was applied either as a single dose at sowing, or as both a starting application and an application at early stem elongation. The highest total application rate was 160 kg N ha⁻¹ y⁻¹. The soil was analysed for phosphorus, potassium, magnesium and calcium in the layer 0–0.3 m, and mineral N down to 0.6 m. Canopy reflectance observations at BBCH 32 was used to calculate a vegetation index (TCARI(32)) to reflect the canopy (leaf and straw) chlorophyll concentration. Harvested grain CP correlated only marginally with the observed soil variables. It was, however, for a specific cultivar, possible to make a prediction of grain CP based on day of sowing and TCARI(32) with . Part of the sowing day effect might be due to thermal stress during grain filling, as the risk for high temperatures during this phase was higher when sowing was late. This might also explain why the introduction of accumulated temperature during grain filling, into a model already including sowing day and TCARI(32) as independent variables, did not improve the predictability of grain CP.     

Influence of winter and spring wheat genetic backgrounds on haploid induction parameters and trait correlations in the wheat × maize system

The interactive influence of winter and/or spring wheat genetic background on haploid induction parameters and trait correlation was studied by hybridizing five elite and diverse genotypes each of winter and spring wheat and their F₁s (winter × winter, spring × spring, and winter × spring, generated in a diallel design excluding reciprocals) with a single genotype of maize. Data were recorded with respect to per cent seed formation, embryo formation, and regeneration. High genetic variability was present among the wheat genotypes (parents + F₁s) for the three haploid induction parameters. Significant differences were obtained within and between different groups viz., spring wheats, winter wheats, spring × spring wheats, winter × winter wheats, and winter × spring wheats with respect to the three haploid induction parameters based on ANOVA. The winter genotypes (winter parents and winter × winter wheat hybrids) responded better than the spring groups (spring wheat parents, spring × spring and winter × spring wheat hybrids) with respect to embryo formation and winter × spring wheat hybrids yielded significantly the highest numbers of regenerants. Correlation studies amongst the haploid induction parameters indicated that the genes controlling seed formation and haploid plantlet regeneration are negatively correlated when the genetic backgrounds of both ecotypes are combined in winter × spring hybrids. Haploid embryo formation had no association with seed formation and regeneration in all genetic backgrounds, suggesting independent inheritance.     

Genetic dissection of endosperm hydration in malting barley (Hordeum vulgare)

Hydration of the endosperm is a critical part of the malting process that ensures proper modification of the grain. However, little is known about the genetic controls of endosperm hydration and its relationship to agronomic and malt quality traits. The extent of endosperm hydration is estimated through hydration index (HYI). We measured HYI, agronomic, and malt quality traits on a 169-line subset of the NSGC Barley Core Panel, which includes global malt lines, some dating from the inception of European breeding programmes. Utilizing GWAS, 61 QTLs were identified for HYI, dormancy, agronomic, and malt quality traits. Of these, six were found to be related to HYI and were located on 1H, 2H, 3H, 6H, and 7H. We found HYI QTLs cosegregating with kernel size and hardness (1H and 3H), malting quality (2H and 6H), and dormancy (2H and 6H). These results indicate that endosperm hydration after steeping can be improved by selecting high HYI alleles on 2H, 6H, and 7H, positively impacting malting quality without negatively impacting kernel size or dormancy.     

Expression of dormancy in a spring wheat cross grown in field and controlled environment conditions

Pre-harvest sprouting resistance in white-seeded wheat, Triticum aestivum L. is a genetically complex trait that varies with environmental conditions. Such variation causes difficulty in phenotypic characterization of populations to study inheritance or develop suitable DNA markers. To minimize random environmental effects, we evaluated controlled environments to measure dormancy. A population of 380 doubled haploid lines, AC Karma/SC8021V2, was evaluated in the glasshouse where the developing grains would not be exposed to moisture and greater consistency in temperature could be achieved. AC Karma is sprouting susceptible and SC8021-V2 is sprouting resistant. The population plus eight checks were seeded in early spring so the plants would reach physiological maturity under long days, requiring less supplemental light, and when the external temperature would be low enough that it would not cause difficulty in cooling the glasshouse. An alpha-lattice in a randomized complete block design with three replications was used. The blocks were arranged to minimize the environmental sources of variability in the glasshouse within each block. A sub-set of this population was tested in six field environments. Dormancy was characterized by germination of seed harvested near physiological maturity, from which a germination resistance index was calculated. The dormancy expressed in the glasshouse was significantly correlated with five of six field environments and highly significant in two of these. There was significant bidirectional transgressive segregation in both field and glasshouse environments. We are currently repeating this glasshouse experiment to confirm the results.     

Grain yield and its components in spring barley under row and hill plot conditions

Twenty homozygous barley lines were grown in row and hill plots at two locations in Ontario in 1973. The hill plots were of two spacings –30×30 cm and 45×45 cm between hills. Within each spacing, three seeding rates were used i.e. 15, 25 and 35 seeds per hill. Grain yield and three of its components, namely; number of seeds per spike (NSS), number of spikes per plot (NSP) and seed weight (SW) were studied. It was concluded that both NSS and SW could be evaluated in hill plots. NSS was found to be an important component of grain yield and its relationship with grain yield was unaffected by plot types and locations. SW also remained unaffected by plot types and seeding rates but as a component of grain yield it was the least important of the three components measured. The correlation between SW and grain yield could be changed by locations. NSP was as important as NSS as a component of grain yield, and was highly correlated with grain yield. Unlike NSS, NSP could not be efficiently evaluated in hill plots since its correlation between row and hill plots was generally lower.     

Simultaneous selection of segregating two-row winter barley material in England and Italy

Summary Independent but simultaneous selection by two breeding teams of the same segregating two-row winter barley material at Cambridge, England and Fiorenzuola, Italy was carried out over the years 1978–1982 with the objective of comparing the individual selection of the two breeding teams and assessing the adaptation of the surviving selections. Visual selection in the generations F₃ to F₅ did not result in any clear cut compartmentalisation of the material and both teams were as likely to select from families previously selected only by the other team as from families they had themselves previously selected. Lines with specific adaptation to each site as well as lines well adapted to both sites were identified from yield trials carried out in F₅ and F₆. The former showed only a very small yield advantage over lines with general adaptation to both sites.     

Genetic variation of β-amylase thermostability among varieties of barley, Hordeum vulgare L., and relation to malting quality

To investigate variation in the thermostability of β-amylase among varieties of barley, Hordeum vulgare L., crude enzyme was extracted from the seeds, and the relative remaining activity was calculated after heat treatment. Our results indicated that the varieties tested were divided into three groups (types A, B and C). All the latest Japanese malting varieties showed high themostability (type A), while European, North American and Australian varieties showed intermediate (type B) or low thermostability (type C). Isoelectric focusing (IEF) of β-amylase was also investigated. Type B varieties divided into two subtypes (types B1 and B2) based on two different IEF patterns (types I and II), whereas those of types A and C showed only one IEF pattern (type II). We also examined their thermostability in the varieties based on the pedigrees of the Japanese malting barley. Our results indicated that the thermostability of β-amylase had a close relationship to fermentability for the production of beer. This fact suggests that the thermostability of β-amylase has a significant influence on the malting quality of barley.     

Agronomic performance of winter wheat grown under highly divergent soil moisture conditions in rainfed and water‐managed environments

Even in the temperate climates of Europe, increasing early season drought and rising air temperature are presenting new challenges to farmers and wheat breeders. Sixteen winter wheat (Triticum aestivum L.) genotypes consisting of three hybrids, six line cultivars and two breeding lines from Germany as well as five line cultivars from France, Austria, Slovakia, Hungary and the Ukraine (referred to as “exotic” lines) have been included in this study. The genetic materials were evaluated over three growing seasons under a range of soil moisture regimes at the three North German sites Braunschweig (irrigated and drought‐stressed), Warmse (rainfed) and Söllingen (rainfed). The average grain yields in the twelve growth environments (water regime × season combinations) ranged from 6.1 to 13.5 t ha^?1. The exotic lines showed little evidence of specific phenological adaptation to drought although they are frequently faced with water scarcity in their countries of origin. The hybrids and German lines exhibited higher regression coefficients (b_i) to environmental means than the exotic lines, indicating particular adaptation to favourable growing conditions. The phenotypical correlations of grain yield between the various environments were high, ranging for instance from 0.6 to 0.8 for the irrigated and drought‐stressed environments at Braunschweig. It is thus expected that in the foreseeable future continued selection aiming at high yield potential will suffice as a means to counter the expected increase in droughts.