H. H. Geiger

Improved methodologies for breeding striga-resistant sorghums.

Parasitic flowering weeds of the genus Striga (Scrophulariaceae) cause substantial losses in sorghum [Sorghum bicolor (L.) Moench] production in sub-Saharan Africa. Striga-resistant sorghum cultivars could be a major component of integrated striga management, if resistance was available in adapted, productive germplasm. In this paper we review methodologies for breeding striga-resistant sorghums. The agar-gel assay is an excellent tool to screen host genotypes in the laboratory for low production of the striga seed germination stimulant. Further laboratory assays are needed which allow the non-destructive, rapid and inexpensive evaluation of individual plants for additional resistance mechanisms. Field screening for striga resistance is hampered by high microvariability in African soils, heterogeneity of natural infestations, and concomitant large environmental effects on striga emergence. An improved field testing methodology should include one or several of the following practices: field inoculation with striga seeds; appropriate experimental design including elevated replication number; specific plot layout; use of appropriate susceptible and resistant checks; evaluation in adjacent infested and uninfested plots; and the use of selection indices derived from emerged striga counts, striga vigor, and grain yield or a host plant damage score. Due to the extreme variability of the parasite and significant genotype×environment interaction effects, multi-locational screening is recommended to obtain materials with stable performance. Additional strategies include: careful definition of the target environments; determination of the most important selection traits in each target environment; characterization of crop germplasm and improvement of available sources of resistance for better agronomic performance; transfer and pyramiding of resistance genes into adapted, farmer-selected cultivars; development of striga-resistant parent lines for hybrid or synthetic cultivars; and development of random-mating populations with multiple sources of resistance. The development of marker-assisted selection techniques for broad-based, polygenic striga resistance is underway. This approach is particularly promising because striga resistance tests are difficult, expensive, and sometimes unreliable; the parasite is quarantined; and some resistance genes are recessive. Transgenic, herbicide-tolerant sorghums could contribute to an immediate, cost-effective control of striga by herbicides, but such cultivars are not yet available. The selection of sorghum cultivars with specific adaptation to integrated striga management approaches could contribute to sustainable sorghum production in striga-infested areas of sub-Saharan Africa

Association mapping reveals gene action and interactions in the determination of flowering time in barley

The interaction between members of a gene network has an important impact on the variation of quantitative traits, and can influence the outcome of phenotype/genotype association studies. Three genes (Ppd-H1, HvCO1, HvFT1) known to play an essential role in the regulation of flowering time under long days in barley were subjected to an analysis of nucleotide diversity in a collection of 220 spring barley accessions. The coding region of Ppd-H1 was highly diverse, while both HvCO1 and HvFT1 showed a rather limited level of diversity. Within all three genes, the extent of linkage disequilibrium was variable, but on average only moderate. Ppd-H1 is strongly associated with flowering time across four environments, showing a difference of five to ten days between the most extreme haplotypes. The association between flowering time and the variation at HvFT1 and HvCO1 was strongly dependent on the haplotype present at Ppd-H1. The interaction between HvCO1 and Ppd-H1 was statistically significant, but this association disappeared when the analysis was corrected for the geographical origin of the accessions. No association existed between flowering time and allelic variation at HvFT1. In contrast to Ppd-H1, functional variation at both HvCO1 and HvFT1 is limited in cultivated barley.

Estimating the outcrossing rate of barley landraces and wild barley populations collected from ecologically different regions of Jordan

The results of previous studies conducted at the University of Hohenheim and the International Center for Agricultural Research in the Dry Areas (ICARDA) indicated that the yielding ability and stability of barley (Hordeum vulgare L.) could be improved in environments with drought stress by increasing the level of heterozygosity. This would require increasing the outbreeding rate of locally adapted breeding materials. As a first step, we estimated the outcrossing rate of 12 barley landraces (Hordeum vulgare ssp. vulgare, in short H. vulgare) and 13 sympatrically occurring populations of its wild progenitor [Hordeum vulgare ssp. spontaneum (C. Koch), in short H. spontaneum] collected from semi-arid localities in Jordan during the 1999/2000 growing season. In each H. vulgare or H. spontaneum population 28–48 spikes were sampled, and up to six offspring (seeds) per spike (called a family) were used for PCR analyses. Collection sites covered high–low transects for rainfall and altitude in order to detect possible environmental effects on the outcrossing rate. Four microsatellite markers located on different chromosomes were used to genotype the samples for estimating the outcrossing rate. Low season-specific multilocus outcrossing rates (tm) were found in both cultivated and wild barley, ranging among populations from 0–1.8% with a mean of 0.34%. Outcrossing rates based on inbreeding equilibrium (te), indicating outcrossing averaged across years, were two- to threefold higher than the season-specific estimates. Under high rainfall conditions somewhat higher—though not significantly higher—outcrossing rates were observed in H. spontaneum than in H. vulgare. The season-specific outcrossing rate in H. spontaneum was positively correlated (r=0.67, P=0.01) with average annual precipitation and negatively correlated (r=0.59, P=0.05) with monthly average temperature during flowering. The results suggest that outcrossing may vary considerably among seasons and that high precipitation and cool temperatures during flowering tend to enhance outcrossing. The rather low levels of outcrossing detected indicate that increased vigour due to heterozygosity has not been a major fitness advantage in the evolution and domestication of H. spontaneum and H. vulgare, respectively. Stable seed production to secure survival under extreme heat and drought stress may have been more important. Cleistogamy may be considered as an effective mechanism to warrant pollination even in drought-stunted plants with non-extruding spikes.

Quantitative-genetic basis of aggressiveness of 42 isolates of fusarium culmorum for winter rye head blight

Forty-two isolates of Fusarium culmorum obtained from diseased plant parts collected from fields in nine European countries and Australia were tested on a synthetic winter rye population. A spore suspension of each isolate was sprayed during midanthesis in five environments (location-year combinations) onto the heads. All isolates were pathogenic as judged by head blight rating scored on a 1 to 9 scale and grain weight relative to the noninoculated control. Isolates differed, however, in their ability to cause disease (aggressiveness). Quantitative geno-typic variation for aggressiveness occurred, while isolate-environment interaction variance, although significant, accounted for only 14% of total variance averaged over both traits. Correlations for aggressiveness across environments ranged from 0.6 to 0.8 (P = 0.01). Estimates of heritability on an entry-mean basis were high (h 2 = 0.9) for both traits, indicating that a substantial proportion of the phenotypic variation was caused by genetic effects. It is concluded that aggressiveness of F. culmorum is inherited as a complex trait.

Rye (Secale cereale L.)

Rye (Secale cereale L.) is mainly a European cereal with about 75% of the global production growing in Russia, Belarus, Poland, Germany, and Ukraine. It has the best overwintering ability, and the highest tolerance to drought, salt, or aluminium stress from all small-grain cereals. Harvest is used for bread making, feed, and in growing demands for ethanol and biomethane production as a renewable energy source. Hybrid rye is competitive to triticale and wheat also on better soils and grown in Germany on about 70% of the total rye acreage. Rye developed in the Middle East as a secondary crop, cultivated rye has its greatest diversity in landraces and populations from Central and East Europe. Their utility for breeding has considerably increased by progress in marker-based introgression of donor chromosome segments. Resistance breeding is presently focused on leaf and stem rust (Puccinia recondita, P. graminis f.sp. secalis), ergot (Claviceps purpurea), and Fusarium diseases. Leaf blotch (Rhynchosporium secalis) and soilborne viruses might gain more attention in the future. Main breeding goals are grain yield, straw shortness, lodging resistance, high kernel weight, tolerances to pre-harvest sprouting and abiotic stresses. Population varieties comprise open-pollinated and synthetic varieties. Both are derived from self-incompatible breeding populations which are steadily improved by recurrent half- or full-sib selection. Open pollinated varieties (OPVs) constitute selected fractions of those populations whereas synthetic varieties are composed of specifically selected parents from which they can identically be reconstituted. Most modern population varieties contain germplasm from two or more genetically distant gene pools. Hybrid breeding is based on self-fertile gene pools and cytoplasmic genic male sterility (CMS) is used as hybridizing mechanism. Long-lasting breeding cycles are needed for the development of seed parent lines since testcrossing is only possible after the inbred lines have been converted to CMS analogues by repeated backcrossing. Options to speed up this process are discussed. Development of restorer lines is straightforward once effective restorer genes have been introduced to the respective breeding populations. Recurrent improvement of fertility restoration is most efficiently accomplished by recombining selected inbred lines after the first or second testcrossing stage. Commercial hybrid seed production requires well-skilled farmers, careful seed processing, and deliberate logistics since rye produces huge amounts of pollen which may be transported over long distances. Even the slightest genetic contamination of the CMS pre-basis and basis seed production may render the respective seed lots worthless for subsequent multiplication. To reduce the cost of the final step of seed production, the CMS seed parent and the pollinator parent are grown as a mixture in a 95:5 ratio. Thus, only about 95% of the certified seed consists of true hybrid seed. Whereas the remainder 5% are randomly intermated plants of the pollinator. However, the latter generally are poor competitors and therefore do not impair the yielding performance of the ‘hybrid’ stand. In the last decades, population and hybrid breeding led to substantial progress in grain yield and other traits.

Comparison of phenotypic and marker-based selection for Fusarium head blight resistance and DON content in spring wheat

Fusarium head blight (FHB) is one of the most economically important wheat diseases, resulting in losses in grain yield and quality as well as contamination with deoxynivalenol (DON). Cultivar Sumai 3 from China and its descendants as well as var. Frontana from Brazil have been identified as potent sources of resistance and subsequently mapped by molecular markers. The aim of the present study was to compare phenotypic and marker-based selection in spring wheat. In a double cross, we combined two donor-quantitative trat loci (QTL) alleles from CM82036 (Sumai 3/Thornbird) located on chromosomes 3B and 5A and one donor-QTL allele from var. Frontana on chromosome 3A with two high-yielding German spring wheat varieties. This initial population was selected phenotypically by a two- (CP1) and three-step procedure (CP1+) and by independent marker-based analysis using one to three flanking markers per QTL (CM). To estimate selection gain, the two phenotypically selected variants and the marker-selected variant as well as an unselected variant (C0) were inoculated with FHB in the field at four locations in 2004. Between 26 and 135 progeny were tested from each variant. FHB severity and DON content were significantly reduced by all selection variants. The highest total selection gain was obtained by the three-step phenotypic selection for both traits, although marker-based selection for the two donor-QTL alleles from CM82036 proved to be more powerful on an annual basis. The large range of variation for FHB resistance and, to a lesser extent, DON content within the marker-based variant, however, shows that an additional phenotypic selection will enhance selection gain.

Quantitative genetic studies on breeding maize for adaptation to organic farming

Organic farming has gained in importance in Germany during recent years. Therefore an increasing demand exists for varieties with specific adaptation to this farming system. In the present study we therefore conducted comparative field experiments with modern maize breeding materials under organic versus conventional farming conditions (ORG and CON, respectively) to estimate quantitative genetic parameters needed for developing optimal breeding strategies and to investigate the perspectives of selection for specific adaptation to ORG. Starting from two broad samples of elite germplasm, consisting of 178 flint and dent lines, respectively, fractions of 11 flint and 11 dent lines were selected based on their testcross performance under ORG. A corresponding set of lines was selected under CON. Testcross performance was evaluated in three regions of Germany and selection of superior lines was practiced across two stages in 2004 and 2005, respectively. The specifically selected lines were crossed in a factorial manner for production of experimental inter-pool single-cross hybrids which were field-tested under ORG and CON in two regions in 2006. Average grain yields were about 16% lower under ORG than under CON. Variance components and entry-mean heritability coefficients under ORG largely resembled those obtained under CON. Phenotypic correlations between ORG and CON were moderate for grain yield and strong for grain dry matter content. No consistent estimates were obtained for the corresponding genotypic correlation for grain yield. At the first stage of testcross selection no evidence of specific adaptation to ORG or CON was observed whereas the factorial crosses tested in 2006 displayed distinct, yet non-significant, advantages when evaluated under the respective target farming system. A small top fraction of hybrids showed outstanding performance under both ORG and CON. The chances of detecting such broadly adapted genotypes are increased if ORG test sites are included in the regular testing system.

Interval mapping of genes for quantitative resistance of maize to Setosphaeria turcica, cause of northern leaf blight, in a tropical environment

Quantitative trait loci (QTL) involved in the resistance of maize to Setosphaeria turcica, the causal agent of northern leaf blight, were located by interval mapping analysis of 121 F2:3 lines derived from a cross between Mo17 (moderately resistant) and B52 (susceptible). A linkage map spanning 112 RFLP loci with 15 cM mean interval length was constructed, based on marker data recorded in a previous study. Field tests with artificial inoculation were conducted at three sites in tropical mid- to high-altitude regions of Kenya, East Africa. Host-plant response was measured in terms of incubation period, disease severity (five scoring dates), and the area under the disease progress curve (AUDPC). Heritability of all traits was high (around 0.75). QTL associated with the incubation period were located on chromosomes 2S and 8L. For disease severity and AUDPC, significant QTL were detected in the putative centromeric region of chromosome 1 and on 2S, 3L, 5S, 6L, 7L, 8L and 9S. On 2S the same marker interval which carried a gene enhancing latent period was also associated with reduced disease severity of juvenile plants. QTL on chromosomes 3L, 5S, 7L and 8L were significant across environments but all other QTL were affected by a large genotype x environment interaction. Partially dominant gene action for resistance as well as for susceptibility was prevailing. Single QTL explained 10 to 38% of the phenotypic variation of the traits. All but the QTL on chromosomes 1, 6 and 9 were contributed by the resistant parent Mo17. On chromosome 8L a QTL mapped to the same region as the major race-specific gene Ht2, supporting the hypothesis that some qualitative and quantitative resistance genes may be allelic.

A gene for resistance to the maize streak virus in the African CIMMYT maize inbred line CML202

Resistance to maize streak virus (MSV) is an essential trait of improved maize varieties in sub-Saharan Africa. We mapped quantitative trait loci (QTL) for resistance to MSV in a population of 196 F2:3 lines derived from a cross between the maize inbred lines CML202 (resistant) from CIMMYT-Zimbabwe and Lo951 (susceptible) from Italy. Field tests were planted at two locations in Zimbabwe, inoculated with viruliferous leaf hoppers (Cicadulina mbila), and scored twice (21 and 83 days after infesting, DAI) on a 1–5 scale. The mean final streak intensity (score 2) of the parent lines was 2.2 (CML202) and 4.8 (Lo951). Genotype × location interaction was large for score 1 but negligible for score 2. Consequently, the heritability was higher for score 2 (0.93) than for score 1 (0.62). By composite interval mapping across locations, using a linkage map with 110 RFLP loci, four significant (LOD ≥3.0) QTL were identified for score 1 on chromosomes (C) 1, 2, 3, and 4, respectively. All four were contributed by CML202. For score 2, only the QTL on C 1 was significant (LOD =37), explaining 59% of the phenotypic and 64% of the genotypic variance. The QTLs partially dominant gene action was consistent with the nearly intermediate resistance of the F1 generation (relative heterosis for resistance 12%). The presence of one major QTL is consistent with the bimodal frequency distribution of the mapping population showing a clear 3:1 segregation. This gene seems to be allelic or identical to Msv1, a major resistance gene which was previously identified in the same genomic region in Tzi4, an inbred line from IITA. Inbred CML202 had lower final disease ratings than Tzi4. The greater resistance of CML202 may be due to allelic differences at the msv1 locus or due to the minor QTL on C 2, 3, and 4 which were not detected in Tzi4.z y Trigo (International Maize and Wheat Improvement Center); IITA, International Institute of Tropical Agriculture; IRAT, Institute de Recherches Agronomiques Tropicales et des Cultures Vivrières; KARI, Kenya Agricultural Research Institute; MSV, maize streak virus; QTL, quantitative trait locus/loci

Methodical improvements in rye anther culture

SummaryThe crucial problem in anther culture of rye (Secale cereale L.) is the very low regeneration capacity. Our study was conducted to overcome this restriction. The plant material used included a doubled haploid line (DH), two single crosses between DH Unes, and a tetraploid Secale cereale L. population. The factors carbohydrate source, post-plating temperature treatment, and gelling agent were investigated. Substantial progress was achieved by substituting maltose for sucrose. Top rates of 49 % responding anthers and 20 % green plants were obtained from one of the single crosses after a post-plating cold treatment on geirrte solidified medium. We consider our results a methodical step forward in rye anther culture.