Dwarf alleles differentially affect barley root traits influencing nitrogen acquisition under low nutrient supply.

Sustainable food production depends critically on the development of crop genotypes that exhibit high yield under reduced nutrient inputs. Rooting traits have been widely advocated as being able to influence optimal plant performance, while breeding-based improvements in yield of spring barley suggest that this species is a good model crop. To date, however, molecular genetics knowledge has not delivered realistic plant ideotypes, while agronomic trials have been unable to identify superior traits. This study explores an intermediate experimental system in which root traits and their effect on plant performance can be quantified. As a test case, four modern semi-dwarf barley varieties, which possess either the ari-e.GP or the sdw1 dwarf allele, were compared with the long-stemmed old variety Kenia under two levels of nutrient supply. The two semi-dwarf types differed from Kenia, exhibiting smaller stem mass and total plant nitrogen (N), and improved partitioning of mass and N to grain. Amongst the semi-dwarfs, the two ari-e.GP genotypes performed better than the two sdw1 genotypes under standard and reduced nutrient supply, particularly in root mass, root investment efficiency, N acquisition, and remobilization of N and mass to grain. However, lack of between-genotype variation in yield and N use efficiency indicated limited potential for exploiting genetic variation in existing varieties to improve barley performance under reduced nutrient inputs. Experimental approaches to test the expression of desirable root and shoot traits are scrutinized, and the potential evaluated for developing a spring barley ideotype for low nutrient conditions.

Effects of genetically modified herbicide-tolerant cropping systems on weed seedbanks in two years of following crops.

The Farm Scale Evaluations (FSEs) showed that genetically modified herbicide-tolerant (GMHT) cropping systems could influence farmland biodiversity because of their effects on weed biomass and seed production. Recently published results for winter oilseed rape showed that a switch to GMHT crops significantly affected weed seedbanks for at least 2 years after the crops were sown, potentially causing longer-term effects on other taxa. Here, we seek evidence for similar medium-term effects on weed seedbanks following spring-sown GMHT crops, using newly available data from the FSEs. Weed seedbanks following GMHT maize were significantly higher than following conventional varieties for both the first and second years, while by contrast, seedbanks following GMHT spring oilseed rape were significantly lower over this period. Seedbanks following GMHT beet were smaller than following conventional crops in the first year after the crops had been sown, but this difference was much reduced by the second year for reasons that are not clear. These new data provide important empirical evidence for longer-term effects of GMHT cropping on farmland biodiversity.

Ban on triazine herbicides likely to reduce but not negate relative benefits of GMHT maize cropping.

The UK Farm-Scale Evaluations (FSE) compared the effects on biodiversity of management of genetically modified herbicide-tolerant (GMHT) spring-sown crops with conventional crop management. The FSE reported larger weed abundance under GMHT management for fodder maize, one of three crops studied. Increased seed production may be important for the long-term persistence of these arable weeds and may benefit invertebrates, small mammals and seed-eating birds. In three-quarters of FSE maize fields, growers used atrazine on the conventionally managed half, reflecting contemporary commercial practice. Withdrawal of the triazine herbicides atrazine, simazine and cyanazine from approved lists of EU chemicals could therefore reduce or even reverse the reported benefits of GMHT maize. Here we analyse effects of applications of triazine herbicides in conventional maize regimes on key indicators, using FSE data. Weed abundances were decreased greatly relative to all other regimes whenever atrazine was applied before weeds emerged. Here, we forecast weed abundances in post-triazine herbicide regimes. We predict weed abundances under future conventional herbicide management to be considerably larger than that for atrazine used before weeds emerged, but still smaller than for the four FSE sites analysed that used only non-triazine herbicides. Our overall conclusion is that the comparative benefits for arable biodiversity of GMHT maize cropping would be reduced, but not eliminated, by the withdrawal of triazines from conventional maize cropping.

Weeds in fields with contrasting conventional and genetically modified herbicide-tolerant crops. II. Effects on individual species.

We compared the effects of the management of genetically modified herbicide-tolerant (GMHT) and conventional beet, maize and spring oilseed rape on 12 weed species. We sampled the seedbank before and after cropping. During the season we counted plants and measured seed rain and biomass. Ratios of densities were used to calculate emergence, survival, reproduction and seedbank change. Treatments significantly affected the biomass of six species in beet, eight in maize and five in spring oilseed rape. The effects were generally consistent, with biomass lower in GMHT beet and spring oilseed rape and higher in GMHT maize. With few exceptions, emergence was higher in GMHT crops. Subsequent survival was significantly lowered for eight species in beet and six in spring oilseed rape in the GMHT treatments. It was increased for five species in maize and one in spring oilseed rape. Significant effects on seedbank change were found for four species. However, for many species in beet and spring oilseed rape (19 out of 24 cases), seed densities were lower in the seedbank after GMHT cropping. These differences compounded over time would result in large decreases in population densities of arable weeds. In maize, populations may increase.

Responses of plants and invertebrate trophic groups to contrasting herbicide regimes in the Farm Scale Evaluations of genetically modified herbicide-tolerant crops.

Effects of genetically modified herbicide-tolerant (GMHT) and conventional crop management on invertebrate trophic groups (herbivores, detritivores, pollinators, predators and parasitoids) were compared in beet, maize and spring oilseed rape sites throughout the UK. These trophic groups were influenced by season, crop species and GMHT management. Many groups increased twofold to fivefold in abundance between early and late summer, and differed up to 10-fold between crop species. GMHT management superimposed relatively small (less than twofold), but consistent, shifts in plant and insect abundance, the extent and direction of these effects being dependent on the relative efficacies of comparable conventional herbicide regimes. In general, the biomass of weeds was reduced under GMHT management in beet and spring oilseed rape and increased in maize compared with conventional treatments. This change in resource availability had knock-on effects on higher trophic levels except in spring oilseed rape where herbivore resource was greatest. Herbivores, pollinators and natural enemies changed in abundance in the same directions as their resources, and detritivores increased in abundance under GMHT management across all crops. The result of the later herbicide application in GMHT treatments was a shift in resource from the herbivore food web to the detritivore food web. The Farm Scale Evaluations have demonstrated over 3 years and throughout the UK that herbivores, detritivores and many of their predators and parasitoids in arable systems are sensitive to the changes in weed communities that result from the introduction of new herbicide regimes.

On the rationale and interpretation of the Farm Scale Evaluations of genetically modified herbicide-tolerant crops.

Farmland biodiversity and food webs were compared in conventional and genetically modified herbicide-tolerant (GMHT) crops of beet (Beta vulgaris L.), maize (Zea mays L.) and both spring and winter oilseed rape (Brassica napus L.). GMHT and conventional varieties were sown in a split-field experimental design, at 60-70 sites for each crop, spread over three starting years beginning in 2000. This paper provides a background to the study and the rationale for its design and interpretation. It shows how data on environment, field management and the biota are used to assess the current state of the ecosystem, to define the typical arable field and to devise criteria for selecting, sampling and auditing experimental sites in the Farm Scale Evaluations. The main functional and taxonomic groups in the habitat are ranked according to their likely sensitivity to GMHT cropping, and the most responsive target organisms are defined. The value of the seedbank as a baseline and as an indicator of historical trends is proposed. Evidence from experiments during the twentieth century is analysed to show that large changes in field management have affected sensitive groups in the biota by ca. 50% during a year or short run of years--a figure against which to assess any positive or negative effects of GMHT cropping. The analysis leads to a summary of factors that were, and were not, examined in the first 3 years of the study and points to where modelling can be used to extrapolate the effects to the landscape and the agricultural region.

Crop management and agronomic context of the Farm Scale Evaluations of genetically modified herbicide-tolerant crops.

The Farm Scale Evaluations of genetically modified herbicide-tolerant crops (GMHT) were conducted in the UK from 2000 to 2002 on beet (sugar and fodder), spring oilseed rape and forage maize. The management of the crops studied is described and compared with current conventional commercial practice. The distribution of field sites adequately represented the areas currently growing these crops, and the sample contained sites operated at a range of management intensities, including low intensity. Herbicide inputs were audited, and the active ingredients used and the rates and the timings of applications compared well with current practice for both GMHT and conventional crops. Inputs on sugar beet were lower than, and inputs on spring oilseed rape and forage maize were consistent with, national averages. Regression analysis of herbicide-application strategies and weed emergence showed that inputs applied by farmers increased with weed densities in beet and forage maize. GMHT crops generally received only one herbicide active ingredient per crop, later and fewer herbicide sprays and less active ingredient (for beet and maize) than the conventional treatments. The audit of inputs found no evidence of bias.

Weeds in fields with contrasting conventional and genetically modified herbicide-tolerant crops. I. Effects on abundance and diversity.

We compared the seedbanks, seed rains, plant densities and biomasses of weeds under two contrasting systems of management in beet, maize and spring oilseed rape. Weed seedbank and plant density were measured at the same locations in two subsequent seasons. About 60 fields were sown with each crop. Each field was split, one half being sown with a conventional variety managed according to the farmer's normal practice, the other half being sown with a genetically modified herbicide-tolerant (GMHT) variety, with weeds controlled by a broad-spectrum herbicide. In beet and rape, plant densities shortly after sowing were higher in the GMHT treatment. Following weed control in conventional beet, plant densities were approximately one-fifth of those in GMHT beet. In both beet and rape, this effect was reversed after the first application of broad-spectrum herbicide, so that late-season plant densities were lower in the GMHT treatments. Biomass and seed rain in GMHT crops were between one-third and one-sixth of those in conventional treatments. The effects of differing weed-seed returns in these two crops persisted in the seedbank: densities following the GMHT treatment were about 20% lower than those following the conventional treatment. The effect of growing maize was quite different. Weed density was higher throughout the season in the GMHT treatment. Late-season biomass was 82% higher and seed rain was 87% higher than in the conventional treatment. The difference was not subsequently detectable in the seedbank because the total seed return was low after both treatments. In all three crops, weed diversity was little affected by the treatment, except for transient effects immediately following herbicide application.

Genetic differentiation over a spatial environmental gradient in wild Rubus ideaus populations.

• The distribution is examined of molecular (random amplified polymorphic DNA) markers within and between 12 wild raspberry (Rubus ideaus) populations ranging over 600 m altitude and 40 km distance from a lowland area of extensive raspberry cultivation to remote upland sites. • Most individuals are distinct, suggesting that seedling recruitment is the main means of propagation, but mean genetic similarities within sites (> 80%) are much greater than between sites (> 50%), suggesting a hindrance to gene movement between sites. • Sites of genetically distinct populations can be grouped in lowland, valley and upland clusters; genetic similarity between sites decreased at slightly > 4% per 100 m difference in altitude from c. 80% between adjacent sites to 50% at 600 m altitude difference. • Together with physiological data collected previously, the molecular evidence confirms that wild populations are more diverse than cultivars and suggests little gene flow from cultivated to wild, a result perhaps of reproductive asynchrony and little opportunity for seedling recruitment in established populations. The cause of the genetic differentiation between sites is not known and requires further study.

Spatial trends of phenotypic diversity between colonies of wild raspberry Rubus idaeus.

• The question 'are fragmented wild populations of raspberry adaptively differentiated from each other and from cultivated forms of the same species?' is addressed here. • Plants collected from the coast, where commercial raspberries are grown, northwards by 49 km to an altitude of 600 m in Tayside, Scotland, were cultured in two common environments. Twenty phenotypic traits were recorded over 2 yr, on vegetative primocanes and then the single dominant floricane retained for year 2. A novel approach is presented for selecting traits that best discriminate between individuals using principal coordinate analysis. Phenotypic variation among accessions was then quantified using principal coordinate analysis followed by principal component analysis. • A consistent north-south trend was found. Plants from northern sites were shorter, bushier with less lateral growth and fewer flowers per lateral on the dominant fruiting cane. Plants from southern sites produced few, tall primocanes with greater cane diameters, lateral growth and flowering. The results were consistent across test environments. • The results confirm substantial, adaptive differentiation between populations and suggest a limited effect of cultivation on wild forms.

Temperature-dependent germination traits in oilseed rape associated with 5'-anchored simple sequence repeat PCR polymorphisms.

An experiment was conducted to test the hypothesis that phenotypes differing in germination rate and the presence or absence of secondary dormancy at low temperature were not genetically different. Seed of oilseed rape was germinated at 4, 10 and 19 degrees C, where selections were made in the percentile ranges 1-10 (early), 45-55 (intermediate) and 91-100 (late). Secondary dormancy occurred only in the late selections at the two lower temperatures. Thermal weighting of curves of cumulative germination on time gave circumstantial evidence that early percentiles were similar at all three temperatures and that seeds with secondary dormancy came largely from later percentiles above the 50th. To test for genetic differentiation between phenotypes, 5'-anchored simple sequence repeat primers were used to generate DNA marker profiles of seedlings raised from seed from each category. Principal coordinate analysis, and more detailed comparisons using the most discriminating markers, confirmed that the early germinators at the three temperatures were not associated with different banding profiles, but seeds entering secondary dormancy, particularly at 10 degrees C, were genetically distinct from germinators at the same temperature. Secondary dormant seeds at low temperature appear to originate mainly from the late germinating seed at higher temperature. Effects of temperature history and the requirement for alternating temperatures to break secondary dormancy were quantified. The results confirm the existence of genetically discrete sub-populations differing in ecologically significant traits.

Studies of the mechanism of action of fusicoccin, the fungal toxin that induces wilting, and its interaction with abscisic acid.

Effects of fusicoccin alone and together with abscisic acid were observed on the stomatal complex of Commelina communis. The experimental material consisted of isolated epidermal strips incubated in a medium containing the ions required for stomatal opening. Fusicoccin stimulated opening and this was accompanied by potassium entry into the guard cells, and hydrolysis of the starch in their chloroplasts. Abscisic acid alone inhibited potassium entry and starch hydrolysis, but these effects could be almost entirely overcome by fusicoccin.Attempts were made to measure the solute potential of the guard cells under the various treatments. Abscisic acid clearly increased their solute potential, but no absolute measurements could be made in the presence of fusicoccin owing to a failure of plasmolysis even with mannitol solutions of solute potential as low as -35 bars. Experiments using isotopically labelled mannitol indicated a massive uptake into the epidermis in the presence of fusicoccin.The mechanism of stimulation of stomatal opening by fusicoccin probably depends in part on a stimulation of the normal processes associated with opening in the guard cells, but may also involve release of pressure due to destruction of the surrounding cells. The effectiveness of this toxin under natural conditions may depend on its ability to counteract effects of abscisic acid, the stress hormone that induces stomatal closure.