The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) canopy size on grass weed growth and grass weed seed return

Four experiments conducted over three seasons (2002-05) at the Crops Research Unit, University of Reading, investigated effects of canopy management of autumn sown oilseed rape (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) on competition with grass weeds. Emphasis was placed on the effect of the crop on the weeds. Rape canopy size was manipulated using sowing date, seed rate and the application of autumn fertilizer. Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds. were sown as indicative grass weeds. The effects of sowing date, seed rate and autumn nitrogen on crop competitive ability were correlated with rape biomass and fractional interception of photosynthetically active radiation (PAR) by the rape floral layer, to the extent that by spring there was good evidence of crop: weed replacement. An increase in seed rate up to the highest plant densities tested increased both rape biomass and competitiveness, e.g. in 2002/3, L. multiflorum head density was reduced from 539 to 245 heads/m(2) and spikelet density from 13 170 to 5960 spikelets/m(2) when rape plant density was increased from 16 to 81 plants/m(2). Spikelets/head of Lolium spp. was little affected by rape seed rate, but the length of heads of A. myosuroides was reduced by 9 % when plant density was increased from 29-51 plants/m(2). Autumn nitrogen increased rape biomass and reduced L. multiflorum head density (415 and 336 heads/m(2) without and with autumn nitrogen, respectively) and spikelet density (9990 and 8220 spikelets/m(2) without and with autumn nitrogen, respectively). The number of spikelets/head was not significantly affected by autumn nitrogen. Early sowing could increase biomass and competitiveness, but poor crop establishment sometimes overrode the effect. Where crop and weed establishment was similar for both sowing dates, a 2-week delay (i.e. early September to mid-September) increased L. multiflorum head density from 226 to 633 heads/m(2) and spikelet density from 5780 to 15 060 spikelets/m(2).

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) cultivar and grass genotype on the competitive balance between crop and grass weeds

Three experiments conducted over two years (2002-04) at the Crops Research Unit, University of Reading, investigated competition between autumn sown oilseed rape cultivars (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) and Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds., sown as indicative grass weeds. Rape cultivar (cv.) had a substantial effect on grass weed seed return. Over the six cultivars tested, L. multiflorum spikelet production ranged from just under 400 spikelets/m(2) in the presence of cv. Winner to nearly 5800 in competition with cv. Lutin. Cultivar competitiveness was associated with high biomass, large dense floral layers and early stem extension. There was some evidence of differential competitive tolerance between rape cultivars. The results suggested that rape cultivars could be screened for competitiveness by measuring floral layer interception of photosynthetic active radiation. L. x boucheanum cultivars varied in ability to compete with rape. In the absence of inter-specific competition, spikelet density was similar for Aberecho and Polly (circa 31000 spikelets/m(2)) but when grown with rape Polly outyielded Aberecho (i.e. 12 090 and 7990 spikelets/m(2) respectively).

The influence of winter oilseed rape (Brassica napus ssp. oleifera var. biennis) cultivar and grass genotype on the competitive balance between crop and grass weeds

Four experiments conducted over three seasons (2002–05) at the Crops Research Unit, University of Reading, investigated effects of canopy management of autumn sown oilseed rape (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) on competition with grass weeds. Emphasis was placed on the effect of the crop on the weeds. Rape canopy size was manipulated using sowing date, seed rate and the application of autumn fertilizer. Lolium multiflorum Lam., L.rboucheanum Kunth and Alopecurus myosuroides Huds. were sown as indicative grass weeds. The effects of sowing date, seed rate and autumn nitrogen on crop competitive ability were correlated with rape biomass and fractional interception of photosynthetically active radiation (PAR) by the rape floral layer, to the extent that by spring there was good evidence of crop: weed replacement. An increase in seed rate up to the highest plant densities tested increased both rape biomass and competitiveness, e.g. in 2002/3, L. multiflorum head density was reduced from 539 to 245 heads/m2 and spikelet density from 13 170 to 5960 spikelets/m2 when rape plant density was increased from 16 to 81 plants/m2. Spikelets/head of Lolium spp. was little affected by rape seed rate, but the length of heads of A. myosuroides was reduced by 9%when plant density was increased from 29–51 plants/m2. Autumn nitrogen increased rape biomass and reduced L. multiflorum head density (415 and 336 heads/m2 without and with autumn nitrogen, respectively) and spikelet density (9990 and 8220 spikelets/m2 without and with autumn nitrogen, respectively). The number of spikelets/head was not significantly affected by autumn nitrogen. Early sowing could increase biomass and competitiveness, but poor crop establishment sometimes overrode the effect. Where crop and weed establishment was similar for both sowing dates, a 2-week delay (i.e. early September to mid-September) increased L. multiflorum head density from 226 to 633 heads/m2 and spikelet density from 5780 to 15 060 spikelets/m2.

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) canopy size on grass weed growth and grass weed seed return

Four experiments conducted over three seasons (2002-05) at the Crops Research Unit, University of Reading, investigated effects of canopy management of autumn sown oilseed rape (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) on competition with grass weeds. Emphasis was placed on the effect of the crop on the weeds. Rape canopy size was manipulated using sowing date, seed rate and the application of autumn fertilizer. Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds. were sown as indicative grass weeds. The effects of sowing date, seed rate and autumn nitrogen on crop competitive ability were correlated with rape biomass and fractional interception of photosynthetically active radiation (PAR) by the rape floral layer, to the extent that by spring there was good evidence of crop: weed replacement. An increase in seed rate up to the highest plant densities tested increased both rape biomass and competitiveness, e.g. in 2002/3, L. multiflorum head density was reduced from 539 to 245 heads/m(2) and spikelet density from 13 170 to 5960 spikelets/m(2) when rape plant density was increased from 16 to 81 plants/m(2). Spikelets/head of Lolium spp. was little affected by rape seed rate, but the length of heads of A. myosuroides was reduced by 9 % when plant density was increased from 29-51 plants/m(2). Autumn nitrogen increased rape biomass and reduced L. multiflorum head density (415 and 336 heads/m(2) without and with autumn nitrogen, respectively) and spikelet density (9990 and 8220 spikelets/m(2) without and with autumn nitrogen, respectively). The number of spikelets/head was not significantly affected by autumn nitrogen. Early sowing could increase biomass and competitiveness, but poor crop establishment sometimes overrode the effect. Where crop and weed establishment was similar for both sowing dates, a 2-week delay (i.e. early September to mid-September) increased L. multiflorum head density from 226 to 633 heads/m(2) and spikelet density from 5780 to 15 060 spikelets/m(2).

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) cultivar and grass genotype on the competitive balance between crop and grass weeds

Three experiments conducted over two years (2002-04) at the Crops Research Unit, University of Reading, investigated competition between autumn sown oilseed rape cultivars (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) and Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds., sown as indicative grass weeds. Rape cultivar (cv.) had a substantial effect on grass weed seed return. Over the six cultivars tested, L. multiflorum spikelet production ranged from just under 400 spikelets/m(2) in the presence of cv. Winner to nearly 5800 in competition with cv. Lutin. Cultivar competitiveness was associated with high biomass, large dense floral layers and early stem extension. There was some evidence of differential competitive tolerance between rape cultivars. The results suggested that rape cultivars could be screened for competitiveness by measuring floral layer interception of photosynthetic active radiation. L. x boucheanum cultivars varied in ability to compete with rape. In the absence of inter-specific competition, spikelet density was similar for Aberecho and Polly (circa 31000 spikelets/m(2)) but when grown with rape Polly outyielded Aberecho (i.e. 12 090 and 7990 spikelets/m(2) respectively).

Distribution of calcium (Ca) and magnesium (Mg) in the leaves of Brassica rapa under varying exogenous Ca and Mg supply

Background and Aims Leafy vegetable Brassica crops are an important source of dietary calcium (Ca) and magnesium (Mg) and represent potential targets for increasing leaf Ca and Mg concentrations through agronomy or breeding. Although the internal distribution of Ca and Mg within leaves affects the accumulation of these elements, such data are not available for Brassica. The aim of this study was to characterize the internal distribution of Ca and Mg in the leaves of a vegetable Brassica and to determine the effects of altered exogenous Ca and Mg supply on this distribution. Methods Brassica rapa ssp. trilocularis ‘R-o-18’ was grown at four different Ca:Mg treatments for 21 d in a controlled environment. Concentrations of Ca and Mg were determined in fully expanded leaves using inductively coupled plasma-mass spectrometry (ICP-MS). Internal distributions of Ca and Mg were determined in transverse leaf sections at the base and apex of leaves using energy-dispersive X-ray spectroscopy (EDS) with cryo-scanning electron microscopy (cryo-SEM). Key Results Leaf Ca and Mg concentrations were greatest in palisade and spongy mesophyll cells, respectively, although this was dependent on exogenous supply. Calcium accumulation in palisade mesophyll cells was enhanced slightly under high Mg supply; in contrast, Mg accumulation in spongy mesophyll cells was not affected by Ca supply. Conclusions The results are consistent with Arabidopsis thaliana and other Brassicaceae, providing phenotypic evidence that conserved mechanisms regulate leaf Ca and Mg distribution at a cellular scale. The future study of Arabidopsis gene orthologues in mutants of this reference B. rapa genotype will improve our understanding of Ca and Mg homeostasis in plants and may provide a model-to-crop translation pathway for targeted breeding.

Use of airborne remote sensing to detect riverside Brassica rapa to aid in risk assessment of transgenic crops

High resolution descriptions of plant distribution have utility for many ecological applications but are especially useful for predictive modelling of gene flow from transgenic crops. Difficulty lies in the extrapolation errors that occur when limited ground survey data are scaled up to the landscape or national level. This problem is epitomized by the wide confidence limits generated in a previous attempt to describe the national abundance of riverside Brassica rapa (a wild relative of cultivated rapeseed) across the United Kingdom. Here, we assess the value of airborne remote sensing to locate B. rapa over large areas and so reduce the need for extrapolation. We describe results from flights over the river Nene in England acquired using Airborne Thematic Mapper (ATM) and Compact Airborne Spectrographic Imager (CASI) imagery, together with ground truth data. It proved possible to detect 97% of flowering B. rapa on the basis of spectral profiles. This included all stands of plants that occupied >2m square (>5 plants), which were detected using single-pixel classification. It also included very small populations (<5 flowering plants, 1-2m square) that generated mixed pixels, which were detected using spectral unmixing. The high detection accuracy for flowering B. rapa was coupled with a rather large false positive rate (43%). The latter could be reduced by using the image detections to target fieldwork to confirm species identity, or by acquiring additional remote sensing data such as laser altimetry or multitemporal imagery.

Use of airborne remote sensing to detect riverside Brassica rapa to aid in risk assessment of transgenic crops

High resolution descriptions of plant distribution have utility for many ecological applications but are especially useful for predictive modeling of gene flow from transgenic crops. Difficulty lies in the extrapolation errors that occur when limited ground survey data are scaled up to the landscape or national level. This problem is epitomized by the wide confidence limits generated in a previous attempt to describe the national abundance of riverside Brassica rapa (a wild relative of cultivated rapeseed) across the United Kingdom. Here, we assess the value of airborne remote sensing to locate B. rapa over large areas and so reduce the need for extrapolation. We describe results from flights over the river Nene in England acquired using Airborne Thematic Mapper (ATM) and Compact Airborne Spectrographic Imager (CASI) imagery, together with ground truth data. It proved possible to detect 97% of flowering B. rapa on the basis of spectral profiles. This included all stands of plants that occupied >2m square (>5 plants), which were detected using single-pixel classification. It also included very small populations (<5 flowering plants, 1-2m square) that generated mixed pixels, which were detected using spectral unmixing. The high detection accuracy for flowering B. rapa was coupled with a rather large false positive rate (43%). The latter could be reduced by using the image detections to target fieldwork to confirm species identity, or by acquiring additional remote sensing data such as laser altimetry or multitemporal imagery.

Fitness of hybrids between rapeseed (Brassica napus) and wild Brassica rapa in natural habitats

Fitness of hybrids between genetically modified (GM) crops and wild relatives influences the likelihood of ecological harm. We measured fitness components in spontaneous (non-GM) rapeseed x Brassica rapa hybrids in natural populations. The F-1 hybrids yielded 46.9% seed output of B. rapa, were 16.9% as effective as males on B. rapa and exhibited increased self-pollination. Assuming 100% GM rapeseed cultivation, we conservatively predict < 7000 second-generation transgenic hybrids annually in the United Kingdom (i.e. similar to 20% of F-1 hybrids). Conversely, whilst reduced hybrid fitness improves feasibility of bio-containment, stage projection matrices suggests broad scope for some transgenes to offset this effect by enhancing fitness.

Regulatory hotspots are associated with plant gene expression under varying soil phosphorus supply in Brassica rapa

Gene expression is a quantitative trait that can be mapped genetically in structured populations to identify expression quantitative trait loci (eQTL). Genes and regulatory networks underlying complex traits can subsequently be inferred. Using a recently released genome sequence, we have defined cis- and trans-eQTL and their environmental response to low phosphorus (P) availability within a complex plant genome and found hotspots of trans-eQTL within the genome. Interval mapping, using P supply as a covariate, revealed 18,876 eQTL. trans-eQTL hotspots occurred on chromosomes A06 and A01 within Brassica rapa; these were enriched with P metabolism-related Gene Ontology terms (A06) as well as chloroplast-and photosynthesis-related terms (A01). We have also attributed heritability components to measures of gene expression across environments, allowing the identification of novel gene expression markers and gene expression changes associated with low P availability. Informative gene expression markers were used to map eQTL and P use efficiency-related QTL. Genes responsive to P supply had large environmental and heritable variance components. Regulatory loci and genes associated with P use efficiency identified through eQTL analysis are potential targets for further characterization and may have potential for crop improvement.

Spontaneous gene flow from rapeseed (Brassica napus) to wild Brassica oleracea

Research on the environmental risks of gene flow from genetically modified ( GM) crops to wild relatives has traditionally emphasized recipients yielding most hybrids. For GM rapeseed ( Brassica napus), interest has centred on the 'frequently hybridizing' Brassica rapa over relatives such as Brassica oleracea, where spontaneous hybrids are unreported in the wild. In two sites, where rapeseed and wild B. oleracea grow together, we used flow cytometry and crop-specific microsatellite markers to identify one triploid F-1 hybrid, together with nine diploid and two near triploid introgressants. Given the newly discovered capacity for spontaneous introgression into B. oleracea, we then surveyed associated flora and fauna to evaluate the capacity of both recipients to harm cohabitant species with acknowledged conservational importance. Only B. oleracea occupies rich communities containing species afforded legislative protection; these include one rare micromoth species that feeds on B. oleracea and warrants further assessment. We conclude that increased attention should now focus on B. oleracea and similar species that yield few crop-hybrids, but possess scope to affect rare or endangered associates.

A Brassica exon array for whole-transcript gene expression profiling

Affymetrix GeneChip (R) arrays are used widely to study transcriptional changes in response to developmental and environmental stimuli. GeneChip (R) arrays comprise multiple 25-mer oligonucleotide probes per gene and retain certain advantages over direct sequencing. For plants, there are several public GeneChip (R) arrays whose probes are localised primarily in 39 exons. Plant whole-transcript (WT) GeneChip (R) arrays are not yet publicly available, although WT resolution is needed to study complex crop genomes such as Brassica, which are typified by segmental duplications containing paralogous genes and/or allopolyploidy. Available sequence data were sampled from the Brassica A and C genomes, and 142,997 gene models identified. The assembled gene models were then used to establish a comprehensive public WT exon array for transcriptomics studies. The Affymetrix GeneChip (R) Brassica Exon 1.0 ST Array is a 5 mu M feature size array, containing 2.4 million 25-base oligonucleotide probes representing 135,201 gene models, with 15 probes per gene distributed among exons. Discrimination of the gene models was based on an E-value cut-off of 1E(-5), with <= 98 sequence identity. The 135 k Brassica Exon Array was validated by quantifying transcriptome differences between leaf and root tissue from a reference Brassica rapa line (R-o-18), and categorisation by Gene Ontologies (GO) based on gene orthology with Arabidopsis thaliana. Technical validation involved comparison of the exon array with a 60-mer array platform using the same starting RNA samples. The 135 k Brassica Exon Array is a robust platform. All data relating to the array design and probe identities are available in the public domain and are curated within the BrassEnsembl genome viewer at http://www.brassica.info/BrassEnsembl/index.html.

Genetical and comparative genomics of Brassica under altered Ca supply identifies Arabidopsis Ca-transporter orthologs

Although Ca transport in plants is highly complex, the overexpression of vacuolar Ca2+ transporters in crops is a promising new technology to improve dietary Ca supplies through biofortification. Here, we sought to identify novel targets for increasing plant Ca accumulation using genetical and comparative genomics. Expression quantitative trait locus (eQTL) mapping to 1895 cis- and 8015 trans-loci were identified in shoots of an inbred mapping population of Brassica rapa (IMB211 × R500); 23 cis- and 948 trans-eQTLs responded specifically to altered Ca supply. eQTLs were screened for functional significance using a large database of shoot Ca concentration phenotypes of Arabidopsis thaliana. From 31 Arabidopsis gene identifiers tagged to robust shoot Ca concentration phenotypes, 21 mapped to 27 B. rapa eQTLs, including orthologs of the Ca2+ transporters At-CAX1 and At-ACA8. Two of three independent missense mutants of BraA.cax1a, isolated previously by targeting induced local lesions in genomes, have allele-specific shoot Ca concentration phenotypes compared with their segregating wild types. BraA.CAX1a is a promising target for altering the Ca composition of Brassica, consistent with prior knowledge from Arabidopsis. We conclude that multiple-environment eQTL analysis of complex crop genomes combined with comparative genomics is a powerful technique for novel gene identification/prioritization.

The role of satellite image-processing for national-scale estimates of gene flow from genetically modified crops: rapeseed in the UK as a model.

1. There is concern over the possibility of unwanted environmental change following transgene movement from genetically modified (GM) rapeseed Brassica napus to its wild and weedy relatives. 2. The aim of this research was to develop a remote sensing-assisted methodology to help quantify gene flow from crops to their wild relatives over wide areas. Emphasis was placed on locating sites of sympatry, where the frequency of gene flow is likely to be highest, and on measuring the size of rapeseed fields to allow spatially explicit modelling of wind-mediated pollen-dispersal patterns. 3. Remote sensing was used as a tool to locate rapeseed fields, and a variety of image-processing techniques was adopted to facilitate the compilation of a spatially explicit profile of sympatry between the crop and Brassica rapa. 4. Classified satellite images containing rapeseed fields were first used to infer the spatial relationship between donor rapeseed fields and recipient riverside B. rapa populations. Such images also have utility for improving the efficiency of ground surveys by identifying probable sites of sympatry. The same data were then also used for the calculation of mean field size. 5. This paper forms a companion paper to Wilkinson et al. (2003), in which these elements were combined to produce a spatially explicit profile of hybrid formation over the UK. The current paper demonstrates the value of remote sensing and image processing for large-scale studies of gene flow, and describes a generic method that could be applied to a variety of crops in many countries. 6. Synthesis and applications. The decision to approve or prevent the release of a GM cultivar is made at a national rather than regional level. It is highly desirable that data relating to the decision-making process are collected at the same scale, rather than relying on extrapolation from smaller experiments designed at the plot, field or even regional scale. It would be extremely difficult and labour intensive to attempt to carry out such large-scale investigations without the use of remote-sensing technology. This study used rapeseed in the UK as a model to demonstrate the value of remote sensing in assembling empirical information at a national level.

The occurrence and economic impact of plasmodiophora brassicae and clubroot disease

The significance of Plasmodiophora brassicae Woronin and clubroot disease which it incites in members of the family Brassicaceae is reviewed as the focus for this special edition of the Journal of Plant Growth Regulation. This is a monographic treatment of recent research into the pathogen and disease; previous similar treatments are now well over half a century old. Vernacular nomenclature of the disease indicates that it had a well-established importance in agriculture and horticulture from at least the Middle Ages onward in Europe and probably earlier. Subsequently, the pathogen probably spread worldwide as a result of transfer on and in fodder taken by colonists as livestock feed. It is a moot point, however, whether there was much earlier spread by P. brassicae into China and subsequently Japan as Brassica rapa (Chinese cabbage and many variants) colonized those lands in archaeological time. Symptoms, worldwide distribution, and economic impact are briefly described here to provide a basis for understanding subsequent papers. Clubroot disease devastates both infected field and protected vegetable and agricultural Brassica crops. Particular importance is placed on recent reports of crop losses in tropical countries, albeit where the crops are grown in cooler altitudes, and in the Canadian prairie land canola crops. The latter is of enormous importance because this crop is the single most important and essential source of vegetable oils used in human foodstuffs and in industrial lubricants where mineral oils are inappropriate.