Life cycle adaptation to latitudinal and seasonal variation in photoperiod and temperature is a major determinant of evolutionary success in flowering plants. Whereas the life cycle of the dicotyledonous model species Arabidopsis thaliana is controlled by two epistatic genes, FLOWERING LOCUS C and FRIGIDA, three unrelated loci (VERNALIZATION) determine the spring and winter habits of monocotyledonous plants such as temperate cereals. In the core eudicot species Beta vulgaris, whose lineage diverged from that leading to Arabidopsis shortly after the monocot-dicot split 140 million years ago, the bolting locus B is a master switch distinguishing annuals from biennials. Here, we isolated B and show that the pseudo-response regulator gene BOLTING TIME CONTROL 1 (BvBTC1), through regulation of the FLOWERING LOCUS T genes, is absolutely necessary for flowering and mediates the response to both long days and vernalization. Our results suggest that domestication of beets involved the selection of a rare partial loss-of-function BvBTC1 allele that imparts reduced sensitivity to photoperiod that is restored by vernalization, thus conferring bienniality, and illustrate how evolutionary plasticity at a key regulatory point can enable new life cycle strategies.
In two Spanish barley landraces with outstanding resistance to scald, the Rrs1 Rh4 locus was fine mapped including all known markers used in previous studies and closely linked markers were developed. Scald, caused by Rhynchosporium commune, is one of the most prevalent barley diseases worldwide. A search for new resistance sources revealed that Spanish landrace-derived lines SBCC145 and SBCC154 showed outstanding resistance to scald. They were crossed to susceptible cultivar Beatrix to create large DH-mapping populations of 522 and 416 DH lines that were scored for disease resistance in the greenhouse using two R. commune isolates. To ascertain the pattern of resistance, parents and reference barley lines with known scald resistance were phenotyped with a panel of differential R. commune isolates. Subpopulations were genotyped with the Illumina GoldenGate 1,536 SNP Assay and a large QTL in the centromeric region of chromosome 3H, known to harbour several scald resistance genes and/or alleles, was found in both populations. Five SNP markers closest to the QTL were converted into CAPS markers. These CAPS markers, together with informative SSR markers used in other scald studies, confirmed the presence of the Rrs1 locus. The panel of differential scald isolates indicated that the allele carried by both donors was Rrs1 Rh4 . The genetic distance between Rrs1 and its flanking markers was 1.2 cM (11_0010) proximally and 0.9 cM (11_0823) distally, which corresponds to a distance of just below 9 Mbp. The number and nature of scald resistance genes on chromosome 3H are discussed. The effective Rrs1 allele found and the closely linked markers developed are already useful tools for molecular breeding programs and provide a good step towards the identification of candidate genes.
Association analyses of resistance to Rhynchosporium commune in a collection of European spring barley germplasm detected 17 significant resistance quantitative trait loci. The most significant association was confirmed as Rrs1. Rhynchosporium commune is a fungal pathogen of barley which causes a highly destructive and economically important disease known as rhynchosporium. Genome-wide association mapping was used to investigate the genetic control of host resistance to R. commune in a collection of predominantly European spring barley accessions. Multi-year disease nursery field trials revealed 8 significant resistance quantitative trait loci (QTL), whilst a separate association mapping analysis using historical data from UK national and recommended list trials identified 9 significant associations. The most significant association identified in both current and historical data sources, collocated with the known position of the major resistance gene Rrs1. Seedling assays with R. commune single-spore isolates expressing the corresponding avirulence protein NIP1 confirmed that this locus is Rrs1. These results highlight the significant and continuing contribution of Rrs1 to host resistance in current elite spring barley germplasm. Varietal height was shown to be negatively correlated with disease severity, and a resistance QTL was identified that co-localised with the semi-dwarfing gene sdw1, previously shown to contribute to disease escape. The remaining QTL represent novel resistances that are present within European spring barley accessions. Associated markers to Rrs1 and other resistance loci, identified in this study, represent a set of tools that can be exploited by breeders for the sustainable deployment of varietal resistance in new cultivars.
Major resistance gene to rhynchosporium, Rrs18, maps close to the telomere on the short arm of chromosome 6H in barley.
The transition from vegetative growth to reproductive development is a complex process that requires an integrated response to multiple environmental cues and endogenous signals. In Arabidopsis thaliana, which has a facultative requirement for vernalization and long days, the genes of the autonomous pathway function as floral promoters by repressing the central repressor and vernalization-regulatory gene FLC. Environmental regulation by seasonal changes in daylength is under control of the photoperiod pathway and its key gene CO. The root and leaf crop species Beta vulgaris in the caryophyllid clade of core eudicots, which is only very distantly related to Arabidopsis, is an obligate long-day plant and includes forms with or without vernalization requirement. FLC and CO homologues with related functions in beet have been identified, but the presence of autonomous pathway genes which function in parallel to the vernalization and photoperiod pathways has not yet been reported. Here, this begins to be addressed by the identification and genetic mapping of full-length homologues of the RNA-regulatory gene FLK and the chromatin-regulatory genes FVE, LD, and LDL1. When overexpressed in A. thaliana, BvFLK accelerates bolting in the Col-0 background and fully complements the late-bolting phenotype of an flk mutant through repression of FLC. In contrast, complementation analysis of BvFVE1 and the presence of a putative paralogue in beet suggest evolutionary divergence of FVE homologues. It is further shown that BvFVE1, unlike FVE in Arabidopsis, is under circadian clock control. Together, the data provide first evidence for evolutionary conservation of components of the autonomous pathway in B. vulgaris, while also suggesting divergence or subfunctionalization of one gene. The results are likely to be of broader relevance because B. vulgaris expands the spectrum of evolutionarily diverse species which are subject to differential developmental and/or environmental regulation of floral transition.
Beta vulgaris is a facultative perennial species which exhibits large intraspecific variation in vernalization requirement and includes cultivated biennial forms such as the sugar beet. Vernalization requirement is under the genetic control of the bolting locus B on chromosome II. Previously, ethyl methanesulfonate (EMS) mutagenesis of an annual accession had yielded several mutants which require vernalization to bolt and behave as biennials. Here, five F2 populations derived from crosses between biennial mutants and annual beets were tested for co-segregation of bolting phenotypes with genotypic markers located at the B locus. One mutant appears to be mutated at the B locus, suggesting that an EMS-induced mutation of B can be sufficient to abolish annual bolting. Co-segregation analysis in four populations indicates that the genetic control of bolting also involves previously unknown major loci not linked to B, one of which also affects bolting time and was genetically mapped to chromosome IX.
Since April 2000 a two-step anaerobic plant with two subsequent 500 m3 reactors has been producing biogas from fodder beet silage (pH 3.4-4.1) as the sole substrate. The plant is located at Kirchlengern near Bielefeld, Germany. Initially the reactors were inoculated with swine manure at 37 degrees C. After a start-up phase the process was sustained at pH 7.5-8.0 by feeding the silage as sole substrate with an HRT of about 55 d twice a day. Parallel to the biogas plant at Kirchlengern four one-step laboratory reactors were continuously driven at temperatures of 37 degrees C, 45 degrees C, 60 degrees C and 65 degrees C. They were fed with the same silage, but only once per day (one impulse). The organic loading rate (OLR) was adjusted to 3.9 g volatile solids (VS)/(l*d) with a concomitant hydraulic retention time (HRT) of 27 d. There was no problem with starting the reactors, but after 86 days the volumetric gas production of the 65 degrees C reactor ceased and a high amount of approximately 65 mM propionate could be determined. By decreasing the temperature down to 60 degrees C a stable reactor performance was recovered for a period of at least 600 further days. Interestingly microscopic analyses revealed that the morphology of methanogenic bacteria in the 60 degrees C was quite different from the 37 and 45 degrees C reactor exhibiting only rodlike methanogens whereas at 37 degrees C coccoid morphotypes besides rodlike methanogens were dominant. In a 55 degrees C reactor (separate experiment) a mixture of coccoid and rodlike methanogens established. During impulse feeding with 3.9 g (VS)/(l*d) it was observed that the quickest recovery of gas production, the pH, CH4 and CO2 content as well as the redox value could be observed at 37 degrees C or at 45 degrees C. Recovery of 75% gas volume (related to the value before or after impulse feeding) was obtained after 5.5 and 7.5 h of feeding time point whereas the 60 degrees C reactor needed 16 h. Slight significant differences were seen in the spectrum of volatile fatty acids (VFA) reaching at 37 degrees or 45 degrees C its maximum with 10-30 mM total VFA at 2-3 h after feeding. After this the VFA level declined to nearly zero (except for the 60 degrees C reactor). Therefore the 37 degrees C reactor was favoured. A double experiment with a second 37 degrees C reactor was started by a somewhat different inoculation procedure from the remaining 3 reactors, but revealed similar results. An impulse feeding experiment with a very high OLR of 16.5 g (VS)/(l*d) lasting 1 week offered a stable reactor performance with a peak GPR of up to 24 l/(l reactor *d) and an HRT of 5.45d. Therefore a long term operation with an HRT of only 7.5 days and an OLR of 12 g (VS)/(l*d) should be possible. By increasing the temperature no significantly different specific gas production rates and methane yields could be observed, e.g. it gave 600-7001 biogas from 1 kg VS. The corresponding methane content ranged between 62-64%. With a methane content of 63 +/- 1% a yield of 40.1 +/- 2 m3 methane/to...
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