Studies on the sensitivity of monogerm sugar beet germination to water

Excessive water in the substratum inhibited germination of monogerm sugar beet seed but it was not lethal, and other crop seeds tested were not as sensitive as beet. The inhibition was not evident in a pure oxygen atmosphere and was lessened when the fruit cap was removed. Although bacterial populations on fruits absorbed appreciable quantities of oxygen, differences between populations in optimum and excessive water were insufficient to account for the inhibition. Anatomical observations revealed the presence of a pore through the sclerified fruit wall at the point of attachment to the mother plant. Evidence indicated that the pore was the route for oxygen and water uptake to the seed, and theoretical calculations of gas flux suggested that oxygen supply would be insufficient to sustain germination when the pore was filled with water.     

Nuclear and cytoplasmic genetic diversity in weed beet and sugar beet accessions compared to wild relatives: new insights into the genetic relationships within the <Emphasis Type="Italic">Beta vulgaris</Emphasis> complex species

Hybridization between cultivated species and their wild relatives is now widely considered to be common. In the Beta vulgaris complex, the sugar beet seed multiplication areas have been the scene of inadvertent pollination of sugar beet seed bearers by wild ruderal pollen donors, generating a weedy form of beet which infests sugar beet fields in European countries. Up to now, investigations of evolutionary dynamics of genetic diversity within the B. vulgaris complex were addressed using few genetical markers and few accessions. In this study, we tackled this issue using a panel of complementary markers: five nuclear microsatellite loci, four mitochondrial minisatellite loci and one chloroplastic PCR-RFLP marker. We sampled 1,640 individuals that illustrate the actual distribution of inland ruderal beets of South Western France, weed beets and wild sea beets of northern France as well as the diversity of 35 contemporary European diploid cultivars. Nuclear genetic diversity in weed beets appeared to be as high as those of ruderal beets and sea beets, whereas the narrowness of cultivar accessions was confirmed. This genetic bottleneck in cultivars is even more important in the cytoplasmic genome as only one haplotype was found among all sugar beet cultivars. The large majority of weed beet populations also presented this unique cytoplasmic haplotype, as expected owing to their maternal cultivated origin. Nonetheless, various cytoplasmic haplotypes were found within three populations of weed beets, implying wild-to-weed seed flows. Finally, our findings gave new insights into the genetical relationships between the components of the B. vulgaris complex: (1) we found a very strong genetic divergence between wild sea beet and other relatives, which was unexpected given the recent evolutionary history and the full cross-compatibility of all taxa and (2) we definitely confirmed that the classification into cultivated, wild, ruderal and weed forms according to their geographical location, phenotype or their domesticated status is clearly in accordance with genetic clustering despite the very recent domestication process of sugar beet. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Dark-Germination of Reciprocal Hybrid Seed from Light-Requiring and -Indifferent Nicotiana tabacum

Seed resulting from self-fertilization of Nicotiana tabacum plants was tested for germination in uninterrupted darkness at monthly intervals beginning at time of harvest. All seed lots were light-requiring immediately after harvest. Some lots continued to be light-requiring, but others gradually became light-indifferent during storage under laboratory conditions. One light-requiring and one light-indifferent selection were self-pollinated, and reciprocally crossed. The seed resulting from self-fertilization was light-requiring or light-indifferent, respectively, according to the parentage. Seed resulting from the reciprocal crosses differed in dark-germinability. Further, dark-germination of seed from each of the reciprocal crosses differed from that of either parent. Both parents contributed toward light sensitivity of the seed; however, contribution of the maternal parent was greater than that of the pollen parent.     

Long distance pollen-mediated gene flow at a landscape level: the weed beet as a case study

Gene flow is a crucial parameter that can affect the organization of genetic diversity in plant species. It has important implications in terms of conservation of genetic resources and of gene exchanges between crop to wild relatives and within crop species complex. In the Beta vulgaris complex, hybridization between crop and wild beets in seed production areas is well documented and the role of the ensuing hybrids, weed beets, as bridges towards wild forms in sugar beet production areas have been shown. Indeed, in contrast to cultivated beets that are bi-annual, weed beets can bolt, flower and reproduce in the same crop season. Nonetheless, the extent of pollen gene dispersal through weedy lineages remains unknown. In this study, the focus is directed towards weed-to-weed gene flow, and we report the results of a pollen-dispersal analysis within an agricultural landscape composed of five sugar beet fields with different levels of infestation by weed beets. Our results, based on paternity analysis of 3240 progenies from 135 maternal plants using 10 microsatellite loci, clearly demonstrate that even if weedy plants are mostly pollinated by individuals from the same field, some mating events occur between weed beets situated several kilometres apart (up to 9.6 km), with rates of interfield-detected paternities ranging from 11.3% to 17.5%. Moreover, we show that pollen flow appears to be more restricted when individuals are aggregated as most mating events occurred only for short-distance classes. The best-fit dispersal curves were fat-tailed geometric functions for populations exhibiting low densities of weed beets and thin-tailed Weibull function for fields with weed beet high densities. Thus, weed beet populations characterized by low density with geographically isolated individuals may be difficult to detect but are likely to act as pollen traps for pollen emitted by close and remote fields. Hence, it appears evident that interfield pollen-mediated gene flow between weed beets is almost unavoidable and could contribute to the diffusion of (trans)genes in the agricultural landscape.     

Effect of virus yellows on yield of some monogerm cultivars of sugar beet

The yield of plants of monogerm cultivars of sugar beet artificially infected with both beet yellows and beet mild yellowing viruses was, on average, depressed 2–7% for every 100 ‘infected plant weeks’, equivalent to c. £25/ha at 1976 prices. The cv. Vytomo, previously recommended to growers as being tolerant of infection by virus yellows, had a high sugar content and abundant foliage but in field trials its actual yield of sugar was no greater when infected, and lower when virus-free, than that of some other monogerm cultivars.     

RFLP markers for sugar beet breeding: chromosomal linkage maps and location of major genes for rhizomania resistance, monogermy and hypocotyl colour

An RFLP linkage map for the nine chromosomes of sugar beet (Beta vulgaris L. ssp. vulgaris var. altissima Doell) was constructed by using a segregating population from a cross between two plants which were heterozygous for several agronomically interesting characters. One hundred and eleven RFLP loci have been mapped to nine linkage groups using 92 genomic markers. The current RFLP map covers a total length of 540 cM. Evidence for the existence of a major gene for rhizomania resistance (Rr1) is given, together with its map position on linkage group IV in the interval between loci GS44 and GS28a. The presence of an RFLP fragment at the GS3d locus is, until now, the best molecular marker for rhizomania-resistant genotypes in segregating populations of sugar beet; GS3d is linked to Rr1 with 6.7 cM. The gene MM, controlling the polygerm/monogerm seed type, has been mapped on linkage group IX in a distal position at 4.2 cM from the locus GS7. The gene R controlling the hypocotyl colour maps to linkage group VII and does not recombine with the RFLP locus GS42. The inheritance of a group of RFLP loci revealed the possible presence of a translocation in the population used to establish the map. The data presented are discussed in relation to the possibility of using RFLP markers in sugar beet breeding.     

Arable weeds identified as new sources of beet mosaic potyvirus in Greece

Field and laboratory studies were carried out between 1992 and 1994 to assess the potential for arable weeds to act as reservoirs of beet mosaic potyvirus (BtMV) in fields where sugar beet is cultivated for seed. 933 weed samples were collected and tested by biological methods, while a representative number of weeds were tested by serological methods. The results showed that 97 samples, representing 14 weed species from eight families, were infected with BtMV. The following weeds were found to be naturally infected with BtMV for the first time: Bilderdykia convolvulus, Nonea pulla, Cerastium glomeratum, Galium aparine, Conyza (Erigeron) spp., Fumaria officinalis, Heliotropium europaeum and Rumex spp. The most common weeds found to be infected with BtMV in high percentage were members of the Papaveraceae.     

Analysis of gene inheritance and expression in hybrids between transgenic sugar beet and wild beets

Reciprocal gene exchange between cultivated sugar beet and wild beets in seed production areas is probably the reason for the occurence of weed beets in sugar beet production fields. Therefore, when releasing transgenic sugar beet plants into the environment, gene transfer to wild beets ( Beta vulgaris ssp. maritima ) has to be considered. In this study the transfer of BNYVV- (beet necrotic yellow vein virus) resistance and herbicide-tolerance genes from two transgenic sugar beet lines that were released in field experiments in 1993 and 1994 in Germany to different wild beet accessions was investigated. In order to evaluate the consequences of outcrossing, manual pollinations of emasculated wild beet plants with homozygous transgenic sugar beet plants were performed. In the resulting hybrids the transgenes were stably inherited according to Mendelian law. Gene expression in leaves and roots of the hybrids was in the same range as in the original transgenic sugar beet plants. Moreover, it was found that in one of the wild beet accessions, transfer and expression of the BNYVV resistance gene did considerably increase the level of virus resistance.     

Crop-weed interactions in the Beta vulgaris complex at a local scale: allelic diversity and gene flow within sugar beet fields

Crop-wild hybrids and weed beets are the main source of agronomic concern for sugar beet production all over Europe. In order to understand the dynamics of crop-wild interactions and the evolution of weediness in Beta vulgaris, we investigated genetic features of bolting individuals occurring at a local scale, i.e. within two sugar beet fields of the French northern area of sugar beet production. By analysing ploidy level, mitochondrial DNA and microsatellite polymorphism, the genetic diversity and the genetic relationships among three different classes of individuals (variety, in-row and out-row weed-beets) from a given field were examined. Such genetic analyses provide a unique opportunity to obtain evidence for the weeds origin and the evolutionary hypotheses previously stated. All the individuals shared in common the Svulg mitochondrial haplotype, and thus a common maternal origin. Conversely, the large genetic diversity at microsatellite loci highlighted the large diversity of the pollinator plants (cultivated and wild plants) during the-seed production process, as well as during the further evolution of weed beets in the sugar production area. Received: 23 April 2001 / Accepted: 15 June 2001     

Bird damage to sugar beet

Pest damage to sugar beet, including that by birds, has been recorded since 1957. During that time damage by rooks has decreased almost to nil, but some other bird damage has greatly increased, most probably as a result of changing agronomic practices, especially the extensive use of herbicides, the introduction of monogerm seed, and the increasing practice of ‘planting-to-a-stand’. The most severe bird damage in the spring is grazing by several species, and in early summer localized felling of plants by pheasants. Observations in the mid–1960's of causes of seedling and plant losses suggested that birds were then of minor importance; the British Sugar Corporation currently consider that birds are the most serious pest of sugar beet. The distribution of the reported damage does not seem to follow any national pattern. In small-plot field trials possible repellent materials such as anthraquinone, methiocarb or thiram, applied to seed or foliage, did not decrease the extent of grazing.     

Enzyme and storage protein electrophoresis in varietal identification of sugar beet

Summary Different methods of classification, based on total protein patterns as well as on specific isoenzyme patterns, were compared in order to establish an identification system for sugar beet varieties and lines. Single seed patterns and bulk extractions of total and fractionated proteins were compared on SDS-PAGE. Due to important intra-populational variation contrasting with similarity at the varietal level no method proved to be sufficiently discriminatory. Twelve sugar beet lines have been genotypically fingerprinted on the basis of five allozyme systems. The allele frequencies of each variety have been measured by using 60–100 individuals. From the data, genetic distance coefficients have been calculated in order to group the different entries by cluster analysis. In addition, a comparison has been made between two seed lots independently obtained from the same parental lines, to test the stability over years. Seeds from the same parental lines, but produced in different localities (Denmark, Italia and USA), were compared to test the influence of the environment on the classification. It has been concluded that isozymes could provide a useful tool for cultivar distinction. The variability at the level of allele frequencies within localities was small. The stability of different generations of the strains was relatively constant. Different strains originating from the same seed firm were less distinct than strains originating from different seed firms.     

Violation of the law of uniformity of the first generation of hybrids

Variability was revealed in the isozyme spectra in the seed progeny from crosses of two beet plants homozygous for contrasting alleles of marker enzyme loci.     

Pollen dispersal in sugar beet production fields

Pollen-mediated gene flow has important implications for biodiversity conservation and for breeders and farmers’ activities. In sugar beet production fields, a few sugar beet bolters can produce pollen as well as be fertilized by wild and weed beet. Since the crop, the wild beets, and the weed beets are the same species and intercross freely, the question of pollen flow is an important issue to determine the potential dispersal of transgenes from field to field and to wild habitats. We report here an experiment to describe pollen dispersal from a small herbicide-resistant sugar beet source towards male sterile target plants located along radiating lines up to 1,200 m away. Individual dispersal functions were inferred from statistical analyses and compared. Pollen limitation, as expected in root-production fields, was confirmed at all the distances from the pollen source. The number of resistant seeds produced by bait plants best fitted a fat-tailed probability distribution curve of pollen grains (power–law) dependent on the distance from the pollen source. A literature survey confirmed that power–law function could fit in most cases. The b coefficient was lower than 2. The number of fertilized flowers by background (herbicide-susceptible) pollen grains was uniform across the whole field. Airborne pollen had a fertilization impact equivalent to that of one adjacent bolter. The individual dispersal function from different pollen sources can be integrated to provide the pollen cloud composition for a given target plant, thus allowing modeling of gene flow in a field, inter-fields in a small region, and also in seed-production area. Long-distance pollen flow is not negligible and could play an important role in rapid transgene dispersal from crop to wild and weed beets in the landscape. The removing of any bolting, herbicide-resistant sugar beet should be compulsory to prevent the occurrence of herbicide-resistant weed beet, thus preventing gene flow to wild populations and preserving the sustainable utility of the resistant varieties. Whether such a goal is attainable remains an open question and certainly would be worth a large scale experimental study.     

The origin and evolution of weed beets: consequences for the breeding and release of herbicide-resistant transgenic sugar beets

Populations of weed beets have expanded into European sugar beet production areas since the 1970s, thereby forming a serious new weed problem for this crop. We sampled seeds in different French populations and studied mitochondrial DNA, chloroplast DNA and life-cycle variability. Given the maternal inheritance of the mitochondrial and chloroplastic genomes and the nuclear determinism of the annual habit, we were able to determine the maternal origin and evolution of these weed beet populations. Our study shows that they carry the dominant allele B for annual habit at high frequency. The main cytoplasmic DNA type found in northern weed beet populations is the cytoplasmic male-sterile type characteristic of sugar beets. We were able to determine that these populations arise from seeds originating from the accidental pollinations of cultivated beets by adventitious beets in the seed production area, which have been transported to the regions where sugar beets are cultivated. These seeds are supposedly the origin of the weed forms and a frequently disturbed cultivated environment has selected for annual habit and early flowering genotypes. We discuss the consequences of the weed beet populations for the breeding, seed production and release of herbicide-resistant transgenic sugar beets.     

Effects of beet cryptic virus infection on sugar beet in field trials

The effects of beet cryptic virus (BCV) infection on sugar beet crops were investigated in field trials in 1990. Two sugar beet breeding stock lines were screened for infection by BCV. Seed lots containing different proportions of seed infected with BCV1 & 2 were obtained by crossing the stock lines and used in field trials at five different sites. Five characteristics of the infected plants were assessed. BCV infection appeared to have no significant effects on the sugar beet crop at four locations which suffered from drought stress but significant effects were found at one site where the crop was grown on grade 1 land with good moisture retention properties. Root yield and sugar yield were reduced by up to 17% and 20%, respectively, by BCV infection.     

Identification of the predominant nonrestoring allele for Owen-type cytoplasmic male sterility in sugar beet (Beta vulgaris L.): development of molecular markers for the maintainer genotype

Hybrid seed production in sugar beet relies on cytoplasmic male sterility (CMS). As time-consuming and laborious test crosses with a CMS tester are necessary to identify maintainer lines, development of a marker-assisted selection method for the rf gene (the nonrestoring allele of restorer-of-fertility locus) is highly desirable for sugar-beet breeding. To develop such a method, we investigated genetic variation at the Rf1 locus, one of two Rf loci known in sugar beet. After HindIII-digestion, genomic DNAs from beet plants known to have a restoring Rf1 allele yielded a range of hybridization patterns on agarose gels, indicating that Rf1 is a multi-allelic locus. However, the hybridization patterns of 22 of 23 maintainer lines were indistinguishable. The nucleotide sequences of the rf1 coding regions of these 22 maintainer lines were found to be identical, confirming that the lines had the same rf1 allele. Two PCR markers were developed that targeted a downstream intergenic sequence and an intron of Rf1. The electrophoretic patterns of both markers indicated multiple Rf1 alleles, one of which, named the dd(L) type, was associated with the maintainer genotype. To test the validity of marker-assisted selection, 147 sugar beet plants were genotyped using these markers. Additionally, the 147 sugar beet plants were crossed with CMS plants to determine whether they possessed the maintainer genotype. Analysis of 5038 F1 offspring showed that 53 % of the dd(L) plants, but none of the plants with other alleles, had the maintainer genotype. Thus, selection for the dd(L) type considerably enriched the proportion of plants with the maintainer genotype.     

Effects of increasing weed-beet density on sugar-beet yield and quality

Weed beets are an increasing problem in many sugar-beet crops in many countries. At present about one sugar-beet field in four in England is infested with weed-beet seed. Control in other crops can be achieved using selective herbicides but in sugar beet the weed beets, many of which are of annual habit, are not easily controlled and often compete with the crop. Experiments were done to quantify the yield loss caused by weed beet in sugar-beet crops. Transects were laid out across three fields in 1985 and 1986 and plots located thereon to include the range of weed-beet densities found in the field. Weed beet did not affect the concentration of sugar (sucrose), potassium, sodium, α amino nitrogen or invert sugar in the crop beets. Root and sugar yields were progressively reduced by increasing densities of weed beet. A rectangular hyperbola described the data slightly better than an asymptotic model. There was no indication of a threshold density of weed beet below which there was no yield loss, which averaged 11.7% for each weed beet plant/m². This corresponds to an average 0.6% sugar yield loss for each 1% of bolted weed beet in the root crop up to 100%, which is similar to the reported losses resulting from bolters in the root crop.     

Detection of Acidovorax valerianellae, the causing agent of bacterial leaf spots in corn salad [Valerianella locusta (L.) Laterr.], in corn salad seeds

Aim: The black leaf spot disease on corn salad caused by the bacterium Acidovorax valerianellae has been observed in Europe for several years and causes economic losses in corn salad cropping. Contaminated seeds or infested soil are considered as the major infection sources. The use of healthy seed material is the only way to prevent disease outbreaks. Therefore, a sensitive diagnostic method for seed testing should be developed. Methods and Results: Using a triple antibody sandwich ELISA with a high‐specific monoclonal antibody, a quick and reliable detection method for contamination of seed lots with the pathogen was developed. This method allowed to detect contaminated seed lots as well as contamination with A. valerianellae in single seeds. Furthermore, the occurrence and distribution of the pathogen could be shown in symptomatic corn salad leaves and in naturally infested seeds by transmission electron microscopy and immunogold labelling for the first time. Conclusion: Our results confirm the seed transmission of this corn salad disease. Pathogen load and distribution vary between positively tested seed lots. Significance and Impact of the study: With this method, not only routine testing of seed material to eliminate contaminated seed lots from production is possible but also the control of sanitation procedures to reduce contamination.     

Weed seed resources for birds in fields with contrasting conventional and genetically modified herbicide-tolerant crops

The UK Farm Scale Evaluations (FSEs) have shown that the use of broad spectrum herbicides on genetically modified herbicide-tolerant (GMHT) crops can have dramatic effects on weed seed production compared to management of conventional varieties. Here, we use FSE data and information on bird diets to determine how GMHT cropping might change the food resources available to farmland birds. More than 60 fields of each of four crops, spring- and winter-sown oilseed rape, beet and maize, were split, one half being sown with a conventional variety, the other with a GMHT variety. Seed rain from weeds known to be important in the diets of 17 granivorous farmland bird species was measured under the two treatments. In beet and spring oilseed rape, rain of weed seeds important in the diets of 16 bird species was significantly reduced in GMHT compared to conventional halves; for no species did it increase. In winter oilseed rape, rain of weed seeds important in the diets of 10 species was significantly reduced in GMHT halves; for only one species did it increase significantly. By contrast, in maize, rain of weed seeds important in the diets of seven species was significantly greater in GMHT halves; for no species was it reduced. Treatment effects for the total weed seed energy available to each bird species were very similar to those for seed rain alone. Measuring the effects on individual bird species was outside the scope of this study. Despite this, these results suggest that should beet, spring and winter rape crops in the UK be largely replaced by GMHT varieties and managed as in the FSEs, this would markedly reduce important food resources for farmland birds, many of which declined during the last quarter of the twentieth century. By contrast, GMHT maize would be beneficial to farmland birds.