Response and effect traits of arable weeds in agro‐ecosystems: a review of current knowledge

Integrating principles of ecological intensification into weed management strategies requires an understanding of the many relationships among weeds, crops and other organisms of agro‐ecosystems in a changing context. Extensively used during the last two decades in weed science, trait‐based approaches have provided general insights into weed community response to agricultural practices, and recently to understanding the effect of weeds on agro‐ecosystem functioning. In this review, we provide a holistic synthesis of the current knowledge on weed response and effect functional traits. Based on the literature and recent advances in weed science, we review current knowledge on (i) weed functional groups and ecological strategies, (ii) weed functional response traits to cropping systems and (iii) weed functional effect traits affecting agro‐ecosystem functioning. For each functional trait, we explicitly present the assumptions and evidence on the linkage between trait values and ecological functions, in response to either management practices, for example tillage, sowing and herbicides, or biotic interactions, for example crop–weed competition and pollination. Finally, we address and discuss major research avenues that may significantly improve the use of traits and the knowledge of functional diversity in weed science for the future, especially to design and implement more environmentally sustainable weed management strategies.     

A new hypothesis for the functional role of diversity in mediating resource pools and weed–crop competition in agroecosystems

We develop a new conceptual model we call the Resource Pool Diversity Hypothesis (RPDH) aimed at explaining how soil resource pool diversity may mediate competition for soil resources between weeds and crops. The primary tenets of the RPDH are that (i) in plant communities, the intensity of inter-specific competition can depend upon the degree to which niche differentiation and resource partitioning occur among species, (ii) agricultural systems are unique in that management practices, such as crop rotation, source of fertility and weed management, result in inputs to the soil and (iii) these inputs directly or indirectly become soil resource pools from which crops and weeds may partition resources. The RPDH leads to the novel prediction that along a gradient of increasing cropping system diversity, yield loss due to weed–crop competition (i.e. the impact on yield per unit weed density) for soil resources should decrease. Similarly, the degree to which crops and weeds overlap in soil resource niche breadth (which is determined by species-specific functional traits for resource acquisition), will determine the extent to which weed–crop competition weakens as resource pool diversity increases. While there have been no direct tests of the RPDH, we highlight evidence from the agricultural literature that provides strong support for components of the hypothesis. Validation of the RPDH would have important implications across a broad range of cropping systems for the development of management strategies that aim to reduce yield loss impact per unit weed plant density and the fundamental principles of integrated weed management, such as the concepts of weed thresholds and critical periods.     

The Potential Benefits of Weeds with Reference to Small Holder Agriculture in Africa

Weed control is one of the most important crop protection activities undertaken in both intensive and low-input farming systems. However, even under intensive systems, crop protection which is less dependent on pesticides may require that weeds be managed to obtain a balance between crop and non-crop vegetation to encourage an increase in natural enemies of crop pests. In the low-input farming systems which sustain much of the rural population of Africa, weed control is usually done by hand and clean weeding is often beyond the labour resources of the farming family. The vegetational diversity of peasant agriculture in Africa to which weeds make their contribution, helps to decrease the risk of disease and pest epidemics. In addition to the pest control benefits of a diverse agroecosystem, weeds contribute to the resource base of the rural community, providing a source of secondary foods, medicines and insecticides. Weed control within an integrated crop protection system appropriate to the needs of the resource-poor farmer, requires that weeds are managed in such a way that their biodiversity is maintained and the more useful species retained within the field or field margin. Those weeds with high food potential or which have pesticidal or medicinal properties might be deliberately encouraged within the crop or field margins. Certain weed species may harbour important pests or diseases of local crops and therefore should be selectively removed. The paper reviews and discusses the literature on the beneficial and deleterious effects of weeds and argues for a weed management strategy which balances the effects of weed competition on crop production with the ethnobotanical and pest control attributes of individual weed species and weed communities.     

Cropping systems modify soil biota effects on wheat (Triticum aestivum) growth and competitive ability

Plants alter soil biota which subsequently modifies plant growth, plant–plant interactions and plant community dynamics. While much research has been conducted on the magnitude and importance of soil biota effects (SBEs) in natural systems, little is known in agro‐ecosystems. We investigated whether agricultural management systems could affect SBEs impacts on crop growth and crop–weed competition. Utilising soil collected from eight paired farms, we evaluated the extent to which SBEs differed between conventional and organic farming systems. Soils were conditioned by growing two common annual weeds: Amaranthus retroflexus (redroot pigweed) or Avena fatua (wild oat). Soil biota effects were measured in wheat (Triticum aestivum) growth and crop–weed competition, with SBEs calculated as the natural log of plant biomass in pots inoculated with living soil divided by the plant biomass in pots inoculated with sterilised soil. SBEs were generally more positive when soil inoculum was collected from organic farms compared with conventional farms, suggesting that cropping systems modify the relative abundance of mutualistic and pathogenic organisms responsible for the observed SBEs. Also, as feedbacks became more positive, crop–weed competition decreased and facilitation increased. In annual cropping systems, SBEs can alter plant growth and crop–weed competition. By identifying the management practices that promote positive SBEs, producers can minimise the impacts of crop–weed competition and decrease their reliance on off‐farm chemical and mechanical inputs to control weeds, enhancing agroecosystem sustainability.     

Integration of soil, crop and weed management in low-external-input farming systems

Greater adoption and refinement of low-external-input (LEI) farming systems have been proposed as ways to ameliorate economic, environmental and health problems associated with conventional farming systems. Organic soil amendments and crop diversification are basic components of LEI systems. Weed scientists can improve the use of these practices for weed management by improving knowledge of four relevant ecological mechanisms. First, multispecies crop rotations, intercrops and cover crops may reduce opportunities for weed growth and regeneration through resource competition and niche disruption. Secondly, weed species appear to be more susceptible to phytotoxic effects of crop residues and other organic soil amendments than crop species, possibly because of differences in seed mass. Thirdly, delayed patterns of N availability in LEI systems may favour large-seeded crops over small-seeded weeds. Finally, additions of organic materials can change the incidence and severity of soil-borne diseases affecting weeds and crops. Our research on LEI sweetcorn and potato production systems in central and northern Maine (USA) suggests that these mechanisms can reduce weed density and growth while maintaining crop yields. Low-external-input farming systems will advance most quickly through the application of interdisciplinary research focused on these and other ecological mechanisms.     

The implications of spatially variable pre‐emergence herbicide efficacy for weed management

BACKGROUND

The efficacy of pre‐emergence herbicides within fields is spatially variable as a consequence of soil heterogeneity. We quantified the effect of soil organic matter on the efficacy of two pre‐emergence herbicides, flufenacet and pendimethalin, against Alopecurus myosuroides and investigated the implications of variation in organic matter for weed management using a crop–weed competition model.

RESULTS

Soil organic matter played a critical role in determining the level of control achieved. The high organic matter soil had more surviving weeds with higher biomass than the low organic matter soil. In the absence of competition, surviving plants recovered to produce the same amount of seed as if no herbicide had been applied. The competition model predicted that weeds surviving pre‐emergence herbicides could compensate for sublethal effects even when competing with the crop. The ED50 (median effective dose) was higher for weed seed production than seedling mortality or biomass. This difference was greatest on high organic matter soil.

CONCLUSION

These results show that the application rate of herbicides should be adjusted to account for within‐field variation in soil organic matter. The results from the modelling emphasised the importance of crop competition in limiting the capacity of weeds surviving pre‐emergence herbicides to compensate and replenish the seedbank. © 2017 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.     

A functional group approach to the management of UK arable weeds to support biological diversity

Weeds have an important role in maintaining farmland biodiversity. This needs to be balanced with their potential negative impact on crop yield and quality. Mechanistic models of crop–weed competition are an important tool in striking this balance. A range of common UK annual weeds were screened for the eco‐physiological traits required by the models. Using multivariate techniques, a number of functional groups with a similar pattern of productivity and competition were identified, based on trade‐offs between traits. A scheme was developed to assign species outside of the data set to one of the groups, based on life cycle, seed mass, maximum height and time of first flowering. As well as having a similar competitive ability, species within a group also appeared to have a similar ecosystem function, in terms of supporting higher trophic groups. Two beneficial groups of species were identified that combined a relatively low competitive ability with a high importance for invertebrates and birds. The identification of functional groups in the UK arable flora is a useful tool for assessing a weed community in the context of reconciling biodiversity provision with crop production. Preserving beneficial plant functional types within the crop would complement non‐cropped wildlife refuges, such as field margins.     

Strategic tillage in Australian conservation agricultural systems to address soil constraints: How does it impact weed management?

In the conservation agricultural systems practised in Australia, cultivation is not commonly utilised for the purpose of weed control. However, occasional use of tillage (strategic tillage) is implemented every few years for soil amelioration, to address constraints such as acidity, water repellence or soil compaction. Depending on the tillage method, the soil amelioration process buries or disturbs the topsoil. The act of amelioration also changes the soil physical and chemical properties and affects crop growth. While these strategic tillage practices are not usually applied for weed control, they are likely to have an impact on weed seedbank burial, which will in turn affect seed dormancy and seedbank depletion. Strategic tillage impacts on seed burial and soil characteristics will also affect weed emergence, plant survival, competitive ability of weeds against the crop and efficiency of soil applied pre-emergent herbicides. If growers understand the impacts of soil amelioration on weed demography, they can more effectively plan management strategies to apply following the strategic tillage practice. Weed seed burial resulting from a full soil inversion is understood, but for many soil tillage implements, more data is needed on the extent of soil mixing, burial of topsoil and the weed seedbank, physical control of existing weeds and stimulation of emergence following the tillage event. Within the agronomic system, there is no research on optimal timing for a tillage event within the year. There are multiple studies to indicate that strategic tillage can reduce weed density, but in most studies, the weed density increases in subsequent years. This indicates that more research is required on the interaction of amelioration and weed ecology, and optimal weed management strategies following a strategic tillage event to maintain weeds at low densities. However, this review also highlights that, where the impacts of soil amelioration are understood, existing data on weed ecology can be applied to potentially determine impacts of amelioration on weed growth.     

United States Department of Agriculture-Agricultural Research Service research on pest biology: weeds

Over 125 permanent full-time scientists conduct research within the USDA Agricultural Research Service (ARS) on issues related to weeds. The research emphasis of most of these scientists involves ecology and management or biological control of weeds. Many scientists perform research on weed biology as components of their primary projects on weed control and integrated crop and soil management. Describing all ARS projects involved with weed biology is impossible, and consequently only research that falls within the following arbitrarily chosen topics is highlighted in this article: dormancy mechanisms; cell division; diversity of rangeland weeds; soil resources and rangeland weeds; poisonous rangeland plants; horticultural weeds; weed traits limiting chemical control; aquatic and semi-aquatic weeds; weed/transgenic wheat hybrids; seedbanks, seedling emergence and seedling populations; and weed seed production. Within these topics, and others not highlighted, the desire of ARS is that good information on weed biology currently translates or eventually will translate into practical advice for those who must manage weeds.     

Improved management of Avena ludoviciana and Phalaris paradoxa with more densely sown wheat and less herbicide

Summary The effectiveness of crop competition for better weed control and reducing herbicide rates was determined for Avena ludoviciana and Phalaris paradoxa . Four experiments, previously broadcast with seeds of the two weeds in separate plots, were sown with three wheat densities, and emerged weeds were treated with four herbicide doses (0–100% of recommended rate). The measured crop and weed traits were first analysed across experiments for treatment effects. Grain yield and weed seed production data were then analysed using cubic smoothing splines to model the response surfaces. Although herbicide rate for both weeds and crop density for P. paradoxa had significant linear effects on yield, there was a significant non-linearity of the response surface. Similarly, herbicide rate and crop density had significant linear effects on weed seed production, and there was significant non-linearity of the response surface that differed for the weed species. Maximum crop yield and reduction in seed production of P. paradoxa was achieved with approximately 80 wheat plants m⁻² and weeds treated with 100% herbicide rate. For A. ludoviciana , this was 130 wheat plants m⁻² applied with 75% herbicide rate. Alternatively, these benefits were achieved by increasing crop density to 150 plants m⁻² applied with 50% herbicide rate. At high crop density, application of the 100% herbicide rate tended to reduce yield, particularly with the A. ludoviciana herbicide, and this impacted adversely on the suppression of weed seed production. Thus, more competitive wheat crops have the potential for improving weed control and reducing herbicide rates.     

Herbicide‐resistant weed management: focus on glyphosate

This review focuses on proactive and reactive management of glyphosate‐resistant (GR) weeds. Glyphosate resistance in weeds has evolved under recurrent glyphosate usage, with little or no diversity in weed management practices. The main herbicide strategy for proactively or reactively managing GR weeds is to supplement glyphosate with herbicides of alternative modes of action and with soil‐residual activity. These herbicides can be applied in sequences or mixtures. Proactive or reactive GR weed management can be aided by crop cultivars with alternative single or stacked herbicide‐resistance traits, which will become increasingly available to growers in the future. Many growers with GR weeds continue to use glyphosate because of its economical broad‐spectrum weed control. Government farm policies, pesticide regulatory policies and industry actions should encourage growers to adopt a more proactive approach to GR weed management by providing the best information and training on management practices, information on the benefits of proactive management and voluntary incentives, as appropriate. Results from recent surveys in the United States indicate that such a change in grower attitudes may be occurring because of enhanced awareness of the benefits of proactive management and the relative cost of the reactive management of GR weeds. Copyright © 2011 Society of Chemical Industry     

Competition of sorghum cultivars and densities with Japanese millet (Echinochloa esculenta)

Field studies were conducted at two locations in southern Queensland, Australia during the 2003–2004 and 2004–2005 growing seasons to determine the differential competitiveness of sorghum (Sorghum bicolor L. Moench) cultivars and crop densities against weeds and the sorghum yield loss due to weeds. Weed competition was investigated by growing sorghum in the presence or absence of a model grass weed, Japanese millet (Echinochloa esculenta). The correlation analyses showed that the early growth traits (height, shoot biomass, and daily growth rate of the shoot biomass) of sorghum adversely affected the height, biomass, and seed production of millet, as measured at maturity. “MR Goldrush” and “Bonus MR” were the most competitive cultivars, resulting in reduced weed biomass, weed density, and weed seed production. The density of sorghum also had a significant effect on the crop's ability to compete with millet. When compared to the density of 4.5 plants per m², sorghum that was planted at 7.5 plants per m² suppressed the density, biomass, and seed production of millet by 22%, 27% and 38%, respectively. Millet caused a significant yield loss in comparison with the weed‐free plots. The combined weed‐suppressive effects of the competitive cultivars, such as MR Goldrush, and high crop densities minimized the yield losses from the weeds. These results indicate that sorghum competition against grass weeds can be improved by choosing competitive cultivars and by using a high crop density of >7.5 plants per m². These non‐chemical options should be included in an integrated weed management program for better weed management, particularly where the control options are limited by the evolution of herbicide resistance.     

Integrated pest management and weed management in the United States and Canada

There is interest in more diverse weed management tactics because of evolved herbicide resistance in important weeds in many US and Canadian crop systems. While herbicide resistance in weeds is not new, the issue has become critical because of the adoption of simple, convenient and inexpensive crop systems based on genetically engineered glyphosate‐tolerant crop cultivars. Importantly, genetic engineering has not been a factor in rice and wheat, two globally important food crops. There are many tactics that help to mitigate herbicide resistance in weeds and should be widely adopted. Evolved herbicide resistance in key weeds has influenced a limited number of growers to include a more diverse suite of tactics to supplement existing herbicidal tactics. Most growers still emphasize herbicides, often to the exclusion of alternative tactics. Application of integrated pest management for weeds is better characterized as integrated weed management, and more typically integrated herbicide management. However, adoption of diverse weed management tactics is limited. Modifying herbicide use will not solve herbicide resistance in weeds, and the relief provided by different herbicide use practices is generally short‐lived at best. More diversity of tactics for weed management must be incorporated in crop systems. © 2014 Society of Chemical Industry     

Weeds as hosts for new crop pests: the case of Protortonia navesi (Hemiptera: Monophlebidae) on cassava in Brazil

Weeds may serve as reservoirs for new and invasive insect pests. The cassava root scale ( Protortonia navesi ) is a recent pest in the Brazilian 'Cerrado' that causes qualitative and quantitative damage by sucking plant sap. Recently, field surveys revealed that many common weeds in this region act as host for P. navesi in cassava fields. In a discrete survey, 15 weed species were identified that were hosting P. navesi . Among these, 13 species occurred during the cropping season and five were observed 4 months after cassava harvest. Eight months after harvest, only cassava volunteer plants were found to be hosting P. navesi . This survey provided a real example of the problem created when weeds host new crop pests. The management of weeds (including crop volunteers) needs to be considered as part of generalised pest management and pest invasion prevention schemes. Effective weed management can be a means of limiting the survival of new pests and the re-infestation of susceptible crop species in subsequent years. Simple studies like this point to a practical need to create greater collaborations between pest management researchers working within discrete pest categories.     

Suppression of weeds and weed seeds in the soil by stubbles and no-tillage in an arid maize-winter wheat-common vetch rotation on the Loess Plateau of China

Reduced tillage provides ecological and economic benefits to arable land on the Loess Plateau of China, where soil erosion has long been a serious problem and soil water availability is largely restricted. However, high abundances of weeds in reduced tillage systems cause significant yield losses. In this study, we explored the effects of no-tillage and stubble retention on the number and density of weeds and weed seeds in a 12-year maize-winter wheat-common vetch rotation on the Loess Plateau. Four treatments including conventional tillage, no-tillage, conventional tillage+stubble retention and no-tillage+stubble retention were designed and applied. We found that no-tillage increased the number of weed species and weed density in most of the crops, while stubble retention decreased weed density in maize and tended to suppress weeds in both no-tillage treatments(no-tillage and no-tillage+stubble retention). No-tillage led to an increase in the number of weed species in the weed seedbank and tended to increase seed density during the spring growth of winter wheat, but it decreased seed density during post-vetch fallow. Stubble retention tended to reduce seed density during the spring growth of winter wheat and post-vetch fallow. We concluded that no-tillage can promote weeds in the experimental crop rotation, while stubble retention suppresses weeds in untilled fields. The combined effects of stubble retention and no-tillage on weed suppression varied among the three crops. Based on these results, we recommend stubble retention in untilled legume-crop rotations on the Loess Plateau to improve the control of weeds.     

Agronomic evaluation of precise mechanical hoeing and chemical weed control in sugar beet

Most herbicide applications to sugar beet (Beta vulgaris L.) are made to the whole crop area, but there is the opportunity to restrict applications to the crop row, decreasing the usage of herbicide by up to 70%. However, this would require greater use of mechanical weed control between rows. Experiments were performed in two seasons to evaluate the weed control performance of a novel, vision‐guided, inter‐row hoe in sugar beet crops grown on a peat fen soil. Hoe lateral placement was within ±30 mm. A precise hoeing and band spraying treatment was compared with overall herbicide use, and with treatments in which the herbicide applications were replaced by hand weeding to minimize competition between crop and weeds. Two hoe passes were made in each season, at crop growth stages of two and 10–12 true leaves in the first season and four and eight true leaves in the second season. Plant population density was not affected by treatment, indicating that none of the treatments caused crop plant loss. Use of the guided hoe controlled weeds better than overall spraying. Crop yields were not significantly different between treatments, indicating that weed control prevented competition with the crop in all treatments.     

Advancing cover cropping in temperate integrated weed management

The effects of cover crops on weeds and the underlying mechanisms of competition, physical control and allelopathy are not fully understood. Current knowledge reveals great potential for using cover crops as a preventive method in integrated weed management. Cover crops are able to suppress 70–95% of weeds and volunteer crops in the fall‐to‐spring period between two main crops. In addition, cover crop residues can reduce weed emergence during early development of the following cash crop by presenting a physical barrier and releasing allelopathic compounds into the soil solution. Therefore, cover crops can partly replace the weed suppressive function of stubble‐tillage operations and non‐selective chemical weed control in the fall‐to‐spring season. This review describes methods to quantify the competitive and allelopathic effects of cover crops. Insight obtained through such analysis is useful for mixing competitive and allelopathic cover crop species with maximal total weed suppression ability. It seems that cover crops produce and release more allelochemicals when plants are exposed to stress or physical damage. Avena strigose, for example, showed stronger weed suppression under dry conditions than during a moist autumn. These findings raise the question of whether allelopathy can be induced artificially. © 2019 Society of Chemical Industry     

Using phenology prediction in weed management: a review

The success of weed management based on ecological principles and weed biology will depend on a better understanding of the effect of environment on lift history strategies, growth, and competition of weeds; and crops, and particularly upon the ability to predict weed and crop phenology, This paper reviews the importance of phenotypic plasticity to weed and crop competition and other biological interactions. We also discuss the utility of phenological predictions in weed management and review current weed phenology models that are based on thermal time. By understanding the variables that drive plant phenotypic responses, new approaches and more long-term solutions for weed problems can be developed.     

Weed management in organic agriculture: are we addressing the right issues?

Summary Despite the serious threat which weeds offer to organic crop production, relatively little attention has so far been paid to research on weed management in organic agriculture, an issue that is often approached from a reductionist perspective. This paper aims to outline why and how this problem should instead be tackled from a system perspective. Compared with conventional agriculture, in organic agriculture the effects of cultural practices (e.g. fertilization and direct weed control) on crop:weed interactions usually manifest themselves more slowly. It follows that weed management should be tackled in an extended time domain and needs deep integration with the other cultural practices, aiming to optimize the whole cropping system rather than weed control per se . In this respect, cover crop management is an important issue because of its implications for soil, nutrient, pest and weed management. It is stressed that direct (physical) weed control can only be successful where preventive and cultural weed management is applied to reduce weed emergence (e.g. through appropriate choice of crop sequence, tillage, smother/cover crops) and improve crop competitive ability (e.g. through appropriate choice of crop genotype, sowing/planting pattern and fertilization strategy). Two examples of system-oriented weed management systems designed for organic agriculture are illustrated as well as future perspectives and problems.     

Modelling the effects of weeds on crop production

In most quantitative studies on interplant competition, static regression models are used to describe experimental data. However, the generality of these models is limited. More mechanistic models for interplant competition, which simulate growth and production of species in mixtures on the basis of the underlying physiological processes, have been developed in the past decade. Recently, simulation models for competition between species for light and water were improved and a detailed version was developed for sugarbeet and fat hen (Chenopodium album L.). The model was validated with data sets of five field experiments, in which the effect of fat hen on sugarbeet production was analysed. About 98% of the variation in yield loss between the experiments (which ranged from –6 to 96%) could be explained with the model. Further analysis with the model showed that the period between crop and weed emergence was the main factor causing differences in yield loss between the experiments. Sensitivity analysis showed a strong interaction between the effect of the variables weed density and the period between crop and weed emergence on yield reduction. Different quantitative approaches to crop-weed competition are discussed in view of their practical applicability. Simulations of experiments, where both the weed density and the period between crop and weed emergence were varied over a wide range, showed a close relation between relative leaf cover of the weeds shortly after crop emergence and yield loss. This relation indicates that relative leaf cover of the weeds accounts for both the effect of weed density and the period between crop and weed emergence. This relation has the potential to be developed into a powerful tool for weed-control advisory systems.