A Comparative Nitrogen Balance and Productivity Analysis of Legume and Non-legume Supported Cropping Systems: The Potential Role of Biological Nitrogen Fixation.

The potential of biological nitrogen fixation (BNF) to provide sufficient N for production has encouraged re-appraisal of cropping systems that deploy legumes. It has been argued that legume-derived N can maintain productivity as an alternative to the application of mineral fertilizer, although few studies have systematically evaluated the effect of optimizing the balance between legumes and non N-fixing crops to optimize production. In addition, the shortage, or even absence in some regions, of measurements of BNF in crops and forages severely limits the ability to design and evaluate new legume-based agroecosystems. To provide an indication of the magnitude of BNF in European agriculture, a soil-surface N-balance approach was applied to historical data from 8 experimental cropping systems that compared legume and non-legume crop types (e.g., grains, forages and intercrops) across pedoclimatic regions of Europe. Mean BNF for different legume types ranged from 32 to 115 kg ha-1 annually. Output in terms of total biomass (grain, forage, etc.) was 30% greater in non-legumes, which used N to produce dry matter more efficiently than legumes, whereas output of N was greater from legumes. When examined over the crop sequence, the contribution of BNF to the N-balance increased to reach a maximum when the legume fraction was around 0.5 (legume crops were present in half the years). BNF was lower when the legume fraction increased to 0.6-0.8, not because of any feature of the legume, but because the cropping systems in this range were dominated by mixtures of legume and non-legume forages to which inorganic N as fertilizer was normally applied. Forage (e.g., grass and clover), as opposed to grain crops in this range maintained high outputs of biomass and N. In conclusion, BNF through grain and forage legumes has the potential to generate major benefit in terms of reducing or dispensing with the need for mineral N without loss of total output.

Modelling the dynamics of feral alfalfa populations and its management implications.

BACKGROUND: Feral populations of cultivated crops can pose challenges to novel trait confinement within agricultural landscapes. Simulation models can be helpful in investigating the underlying dynamics of feral populations and determining suitable management options. METHODOLOGY/PRINCIPAL FINDINGS: We developed a stage-structured matrix population model for roadside feral alfalfa populations occurring in southern Manitoba, Canada. The model accounted for the existence of density-dependence and recruitment subsidy in feral populations. We used the model to investigate the long-term dynamics of feral alfalfa populations, and to evaluate the effectiveness of simulated management strategies such as herbicide application and mowing in controlling feral alfalfa. Results suggest that alfalfa populations occurring in roadside habitats can be persistent and less likely to go extinct under current roadverge management scenarios. Management attempts focused on controlling adult plants alone can be counterproductive due to the presence of density-dependent effects. Targeted herbicide application, which can achieve complete control of seedlings, rosettes and established plants, will be an effective strategy, but the seedbank population may contribute to new recruits. In regions where roadside mowing is regularly practiced, devising a timely mowing strategy (early- to mid-August for southern Manitoba), one that can totally prevent seed production, will be a feasible option for managing feral alfalfa populations. CONCLUSIONS/SIGNIFICANCE: Feral alfalfa populations can be persistent in roadside habitats. Timely mowing or regular targeted herbicide application will be effective in managing feral alfalfa populations and limit feral-population-mediated gene flow in alfalfa. However, in the context of novel trait confinement, the extent to which feral alfalfa populations need to be managed will be dictated by the tolerance levels established by specific production systems for specific traits. The modelling framework outlined in this paper could be applied to other perennial herbaceous plants with similar life-history characteristics.

Co-adaptation of seed dormancy and flowering time in the arable weed Capsella bursa-pastoris (shepherd's purse).

BACKGROUND AND AIMS: The duration of the plant life cycle is an important attribute that determines fitness and coexistence of weeds in arable fields. It depends on the timing of two key life-history traits: time from seed dispersal to germination and time from germination to flowering. These traits are components of the time to reproduction. Dormancy results in reduced and delayed germination, thus increasing time to reproduction. Genotypes in the arable seedbank predominantly have short time to flowering. Synergy between reduced seed dormancy and reduced flowering time would create stronger contrasts between genotypes, offering greater adaptation in-field. Therefore, we studied differences in seed dormancy between in-field flowering time genotypes of shepherd's purse. METHODS: Genotypes with early, intermediate or late flowering time were grown in a glasshouse to provide seed stock for germination tests. Secondary dormancy was assessed by comparing germination before and after dark-incubation. Dormancy was characterized separately for seed myxospermy heteromorphs, observed in each genotype. Seed carbon and nitrogen content and seed mass were determined as indicators of seed filling and resource partitioning associated with dormancy. KEY RESULTS: Although no differences were observed in primary dormancy, secondary dormancy was weaker among the seeds of early-flowering genotypes. On average, myxospermous seeds showed stronger secondary dormancy than non-myxospermous seeds in all genotypes. Seed filling was similar between the genotypes, but nitrogen partitioning was higher in early-flowering genotypes and in non-myxospermous seeds. CONCLUSIONS: In shepherd's purse, early flowering and reduced seed dormancy coincide and appear to be linked. The seed heteromorphism contributes to variation in dormancy. Three functional groups of seed dormancy were identified, varying in dormancy depth and nitrate response. One of these groups (FG-III) was distinct for early-flowering genotypes. The weaker secondary dormancy of early-flowering genotypes confers a selective advantage in arable fields.

How agro-ecological research helps to address food security issues under new IPM and pesticide reduction policies for global crop production systems.

Drivers behind food security and crop protection issues are discussed in relation to food losses caused by pests. Pests globally consume food estimated to feed an additional one billion people. Key drivers include rapid human population increase, climate change, loss of beneficial on-farm biodiversity, reduction in per capita cropped land, water shortages, and EU pesticide withdrawals under policies relating to 91/414 EEC. IPM (Integrated Pest Management) will be compulsory for all EU agriculture by 2014 and is also being widely adopted globally. IPM offers a 'toolbox' of complementary crop- and region-specific crop protection solutions to address these rising pressures. IPM aims for more sustainable solutions by using complementary technologies. The applied research challenge now is to reduce selection pressure on single solution strategies, by creating additive/synergistic interactions between IPM components. IPM is compatible with organic, conventional, and GM cropping systems and is flexible, allowing regional fine-tuning. It reduces pests below economic thresholds utilizing key 'ecological services', particularly biocontrol. A recent global review demonstrates that IPM can reduce pesticide use and increase yields of most of the major crops studied. Landscape scale 'ecological engineering', together with genetic improvement of new crop varieties, will enhance the durability of pest-resistant cultivars (conventional and GM). IPM will also promote compatibility with semiochemicals, biopesticides, precision pest monitoring tools, and rapid diagnostics. These combined strategies are urgently needed and are best achieved via multi-disciplinary research, including complex spatio-temporal modelling at farm and landscape scales. Integrative and synergistic use of existing and new IPM technologies will help meet future food production needs more sustainably in developed and developing countries, in an era of reduced pesticide availability. Current IPM research gaps are identified and discussed.

The demography of feral alfalfa (Medicago sativa L.) populations occurring in roadside habitats in Southern Manitoba, Canada: implications for novel trait confinement.

PURPOSE: Feral populations of cultivated crops can act as reservoirs for novel genetically engineered (GE) traits and aid in trait movement at the landscape level. However, little information is available on the potential of cultivated crops to become feral. In this study, we investigated the ferality of alfalfa populations (non-GE version) occurring in roadside habitats. Knowledge on the nature of roadside alfalfa populations would be useful for designing efficient trait confinement protocols and coexistence strategies in alfalfa. METHODS: We investigated roadside alfalfa populations from 2006 to 2009 in three rural municipalities (Hanover, MacDonald, and Springfield) in Southern Manitoba, Canada. We studied the demography of these populations including seedbank, seedling recruitment, and fecundity and examined the impact of road verge mowing on key life stages of these populations. We also compared the growth and reproductive attributes of roadside and cultivated alfalfa populations. RESULTS: Alfalfa is reproductively successful in roadside habitats and capable of establishing self-perpetuating populations. A substantial portion of the alfalfa seeds we extracted from seedbank samples were viable but not germinable, suggesting some degree of seedbank persistence in roadside habitats. In the roadside habitat, alfalfa seedlings recruited successfully, however, seedling mortality was high when seedlings were in close proximity to well-established alfalfa plants. Mowing dramatically reduced the reproductive success of roadside alfalfa. Generally, the growth and reproduction of roadside alfalfa was comparable to cultivated alfalfa except for total fecundity. CONCLUSIONS: Considering the long lifespan (>10 years) of alfalfa and the levels of fecundity, seedbank, and seedling survival we observed, long-term persistence of roadside alfalfa populations seems reasonable. In the context of novel trait confinement, our results suggest that feral alfalfa populations required to be managed if there is a desire/need to confine novel traits in alfalfa.

Cumulative impact of GM herbicide-tolerant cropping on arable plants assessed through species-based and functional taxonomies.

BACKGROUND, AIM AND SCOPE: In a gradualist approach to the introduction of crop biotechnology, the findings of experimentation at one scale are used to predict the outcome of moving to a higher scale of deployment. Movement through scales had occurred for certain genetically modified herbicide-tolerant (GMHT) crops in the UK as far as large-scale field trials. However, the land area occupied by these trials was still <1% of the area occupied by the respective non-GM crops. Some means is needed to predict the direction and size of the effect of increasing the area of GMHT cropping on ecological variables such as the diversity among species and trophic interactions. Species-accumulation curves are examined here as a method of indicating regional-scale impacts on botanical diversity from multiple field experiments. MATERIALS AND METHODS: Data were used from experiments on the effect of (GMHT) crops and non-GM, or conventional, comparators in fields sown with four crop types (beet, maize, spring and winter oilseed rape) at a total of 250 sites in the UK between 2000 and 2003. Indices of biodiversity were measured in a split-field design comparing GMHT with the farmers' usual weed management. In the original analyses based on the means at site level, effects were detected on the mass of weeds in the three spring crops and the proportion of broadleaf and grass weeds in winter oilseed rape, but not on indices of plant species diversity. To explore the links between site means and total taxa, accumulation curves were constructed based on the number of plant species (a pool of around 250 species in total) and the number of plant functional types (24), inferred from the general life-history characteristics of a species. RESULTS: Species accumulation differed between GMHT and conventional treatments in direction and size, depending on the type of crop and its conventional management. Differences were mostly in the asymptote of the curve, indicative of the maximum number of species found in a treatment, rather than the steepness of the curve. In winter oilseed rape, 8% more species were accumulated in the GMHT treatment, mainly as a result of the encouragement of grass species by the herbicide when applied in the autumn. (Overall, GMHT winter oilseed rape had strong negative effects on both the food web and the potential weed burden by increasing the biomass of grasses and decreasing that of broadleaf weeds.) In maize, 33% more species-a substantial increase-were accumulated in the GMHT than in the conventional, consistent with the latter's highly suppressive weed management using triazine herbicides. In the spring oilseed rape and beet, fewer species (around 10%) were accumulated in the GMHT than the conventional. The GMHT treatments did not remove or add any functional (life history) types, however. Differences in species accumulation between treatments appeared to be caused by loss or gain of rarer species. The generality of this effect was confirmed by simulations of species accumulation in which the species complement at each of 50 sites was drawn from a regional pool and subjected to reducing treatment at each site. Shifts in the species-accumulation parameters, comparable to those measured, occurred only when a treatment removed the rarer species at each site. DISCUSSION: Species accumulation provided a set of simple curve-parameters that captured the net result of numerous local effects of treatments on plant species and, in some instances, the balance between grass and broadleaf types. The direction of effect was not the same in the four crops and depended on the severity of the conventional treatment and on complex interactions between season, herbicide and crop. The accumulation curves gave an indication of potential positive or negative consequences for regional species pools of replacing a conventional practice with GMHT weed management. In this and related studies, a range of indicators, through which diversity was assessed by both species and functional type, and at both site and regional scales, gave more insight into effects of GMHT treatment than provided by any one indicator. CONCLUSIONS: Species accumulation was shown to discriminate at the regional scale between agronomic treatments that had little effect on species number at the field scale. While a comprehensive assessment of GM cropping needs to include an examination of regional effects, as here, the costs of doing this in all instances would be prohibitive. Simulations of diversity-reducing treatments could provide a theoretical framework for predicting the likely regional effects from in-field plant dynamics. RECOMMENDATIONS AND PERSPECTIVES: Accumulation curves potentially offer a means of linking within-site effects to regional impacts on biodiversity resulting from any change in agricultural practice. To guide empirical measurement, there is a scope to apply a methodology such as individual-based modelling at the field scale to explore the links between agronomic treatments and the relative abundance of plant types. The framework needs to be validated in practice, using species-based and functional taxonomies, the latter defined by measured rather than inferred traits.

Heterogeneity in the distribution of genetically modified and conventional oilseed rape within fields and seed lots.

The implementation of co-existence in the commercialisation of GM crops requires GM and non-GM products to be segregated in production and supply. However, maintaining segregation in oilseed rape will be made difficult by the highly persistent nature of this species. An understanding of its population dynamics is needed to predict persistence and develop potential strategies for control, while to ensure segregation is being achieved, the production of GM oilseed rape must be accompanied by the monitoring of GM levels in crop or seed populations. Heterogeneity in the spatial distribution of oilseed rape has the potential to affect both control and monitoring and, although a universal phenomenon in arable weeds and harvested seed lots, spatial heterogeneity in oilseed rape populations remains to be demonstrated and quantified. Here we investigate the distribution of crop and volunteer populations in a commercial field before and during the cultivation of the first conventional oilseed rape (winter) crop since the cultivation of a GM glufosinate-tolerant oilseed rape crop (spring) three years previously. GM presence was detected by ELISA for the PAT protein in each of three morphologically distinguishable phenotypes: autumn germinating crop-type plants (3% GM), autumn-germinating 'regrowths' (72% GM) and spring germinating 'small-type' plants (17% GM). Statistical models (Poisson log-normal and binomial logit-normal) were used to describe the spatial distribution of these populations at multiple spatial scales in the field and of GM presence in the harvested seed lot. Heterogeneity was a consistent feature in the distribution of GM and conventional oilseed rape. Large trends across the field (50 x 400 m) and seed lot (4 x 1.5 x 1.5 m) were observed in addition to small-scale heterogeneity, less than 20 m in the field and 20 cm in the seed lot. The heterogeneity was greater for the 'regrowth' and 'small' phenotypes, which were likely to be volunteers and included most of the GM plants detected, than for the largely non-GM 'crop' phenotype. The implications of the volunteer heterogeneity for field management and GM-sampling are discussed.

Sources of uncertainty in the quantification of genetically modified oilseed rape contamination in seed lots.

Testing of seed and grain lots is essential in the enforcement of GM labelling legislation and needs reliable procedures for which associated errors have been identified and minimised. In this paper we consider the testing of oilseed rape seed lots obtained from the harvest of a non-GM crop known to be contaminated by volunteer plants from a GM herbicide tolerant variety. The objective was to identify and quantify the error associated with the testing of these lots from the initial sampling to completion of the real-time PCR assay with which the level of GM contamination was quantified. The results showed that, under the controlled conditions of a single laboratory, the error associated with the real-time PCR assay to be negligible in comparison with sampling error, which was exacerbated by heterogeneity in the distribution of GM seeds, most notably at a small scale, i.e. 25 cm3. Sampling error was reduced by one to two thirds on the application of appropriate homogenisation procedures.