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.     

Consequences of narrow crop row spacing and delayed Echinochloa colona and Trianthema portulacastrum emergence for weed growth and crop yield loss in maize

Echinochloa colona and Trianthema portulacastrum are weeds of maize that cause significant yield losses in the Indo‐Gangetic Plains. Field experiments were conducted in 2009 and 2010 to determine the influence of row spacing (15, 25 and 35 cm) and emergence time of E. colona and T. portulacastrum (0, 15, 25, 35, 45 and 55 days after maize emergence; DAME) on weed growth and productivity of maize. A season‐long weed‐free treatment and a weedy control were also used to estimate maize yield and weed seed production. Crop row spacing as well as weed emergence time had a significant influence on plant height, shoot biomass and seed production of both weed species and grain yield of maize in both years. Delay in emergence of weeds resulted in less plant height, shoot biomass and seed production. However, increase in productivity of maize was observed by delay in weed emergence. Likewise, growth of both weed species was less in narrow row spacing (15 cm) of maize, as compared with wider rows (25 and 35 cm). Maximum seed production of both weeds was observed in weedy control plots, where there was no competition with maize crop and weeds were in rows 35 cm apart. Nevertheless, maximum plant height, shoot biomass and seed production of both weed species were observed in 35 cm rows, when weeds emerged simultaneously with maize. Both weed species produced only 3–5 seeds per plant, when they were emerged at 55 DAME in crop rows spaced at 15 cm. Infestation of both weeds at every stage of crop led to significant crop yield loss in maize. Our results suggested that narrow row spacing and delay in weed emergence led to reduced weed growth and seed production and enhanced maize grain yield and therefore could be significant constituents of integrated weed management strategies in maize.     

Elucidating the apparent maize tolerance to weed competition in long-term organically managed systems

In a long-term cropping systems trial comparing organically and conventionally managed systems, organic maize production sustained crop yields equal to conventional methods despite higher weed levels. In 2005 and 2006, an experiment nested within the trial was conducted to gain insight into this apparent crop tolerance to weed competition. Density of mixed weed species was experimentally manipulated to achieve a broad range of weed infestation levels. Under standard management conditions, all cropping systems produced equivalent maize yields, even though weedy plant biomass in the organic treatments was between fourfold and sevenfold greater than in the conventionally managed maize. Increased yield capacity, evidenced when plots were maintained weed-free, and enhanced crop competitiveness, were the main pillars of this apparent crop tolerance to weed competition in the organic systems. Increased soil resource availability and a faster relative crop growth rate in the organic systems probably contributed to these factors, which play an important role in buffering crop fitness during years of less than ideal weed control. Simultaneously, the experiment illustrated the poor efficacy of mechanical weed management in the organic systems, which is the main reason organic maize did not out-yield conventional maize under standard management conditions.     

Does yield loss due to weed competition differ between organic and conventional cropping systems?

High weed abundance in organic crops is thought to be a key factor contributing to the greater yield loss in organic as compared with conventional cropping systems. However, even with greater weed densities than conventional systems, some organic systems have yields comparable to conventional systems, suggesting that cropping systems might differ in yield loss due to weed competition. The diversity in soil nutrient resources due to diversity in crop rotations and variable inputs might enhance crop tolerance to weed competition. We assessed the long‐term effects of contrasting levels of crop rotations (low, medium and high diversity) on weed density, weed biomass and wheat yield loss in organic and no‐till conventional cropping systems using a microplot study within a long‐term cropping systems trial at Scott, Saskatchewan, Canada. Weed density and biomass were found to be four times higher in the organic systems than in the conventional systems. Under standard weed management practices, organic had 44% lower yield than the conventional system. Lower yields in organic, even without weed competition, suggest that the lower yields are due to low soil productivity rather than weed competition. No differences in yield loss were observed among the organic and conventional systems or among the diverse crop rotations. We conclude that the organic management practices and/or increased crop rotation diversity did not enhance yield or reduce yield loss due to weed competition, due to the factors associated with lower soil fertility.     

Modelling herbicide dose and weed density effects on crop:weed competition

The effects of a range of herbicide doses on crop:weed competition were investigated by measuring crop yield and weed seed production. Weed competitivity of wheat was greater in cv. Spark than in cv. Avalon, and decreased with increasing herbicide dose, being well described by the standard dose–response curve. A combined model was then developed by incorporating the standard dose–response curve into the rectangular hyperbola competition model to describe the effects of plant density of a model weed, Brassica napus L., and a herbicide, metsulfuron‐methyl, on crop yield and weed seed production. The model developed in this study was used to describe crop yield and weed seed production, and to estimate the herbicide dose required to restrict crop yield loss caused by weeds and weed seed production to an acceptable level. At the acceptable yield loss of 5% and the weed density of 200 B. napus plants m^–2, the model recommends 0.9 g a.i. metsulfuron‐methyl ha^–1 in Avalon and 2.0 g a.i. in Spark.     

Minimal soil disturbance combined with spring cropping can halt soil seedbank accumulation of Alopecurus myosuroides

The basic mechanism of soil inversion tillage for control of annual weeds is based on the vertical translocation of weed seeds from the soil surface to deeper soil layers. Buried weed seeds either remain dormant in the soil seedbank and are exposed to biological and chemical decay mechanisms, or they germinate but the seedlings cannot reach the soil surface (fatal germination). However, depending on the seed biology of the respective target species, frequent inversion tillage can lead to a build-up of the soil seedbank. For soil seedbank depletion based on available knowledge of the biology of Alopecurus myosuroides seeds, soil inversion tillage is suggested to be reduced to every third or fourth year with reduced or even no-tillage (direct seeding) in between (rotational inversion tillage systems). Including spring crops in the crop rotation could further help dampening the population growth and hence the seed return into the seedbank. This study investigated the effect of rotational inversion tillage in combination with reduced tillage or direct seeding on the soil seedbank and population development of A. myosuroides. In a long-term field trial, set up in 2012, these tillage strategies were compared with continuous inversion tillage in a 3-year crop rotation with two consecutive years of winter wheat (Triticum aestivum) followed by spring barley (Hordeum vulgare). The results showed a significant decline in the soil seedbank following the spring crop, irrespective of the tillage system. The continuous inversion tillage system and inversion tillage before spring cropping with reduced tillage (shallow tillage with a disc harrow) before winter wheat both led to accumulation of seeds in the soil seedbank. In contrast, inversion tillage before spring cropping with direct seeding of winter wheat depleted the soil seedbank significantly after only one crop rotation. Although only covering one intensively studied field site, these findings highlight the need for diversified cropping systems and indicate potential avenues for reducing soil tillage while controlling economically important weeds.     

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.     

Does soil nitrogen affect early competitive traits of annual weeds in comparison with maize?

Soil nitrogen (N) is considered an important driver of crop‐weed interactions, yet the mechanisms involved have been only partially explored, especially with respect to early‐season growth, when competitive hierarchies are formed. This study characterises the effects of different N levels on biomass accumulation and plant morphology for maize (Zea mays), and four important weed species (Amaranthus retroflexus, Abutilon theophrasti, Setaria faberi, and Chenopodium album). Under glasshouse conditions, plants were grown in separate pots and irrigated with nutrient solution at four N concentrations (0.2, 0.5, 2, 5 μm L⁻¹) until 57 days after emergence. Except for S. faberi, which was unresponsive to N, the relative biomass growth rates (RGR) of maize and the broad‐leaved weeds were positively and similarly affected by increasing nitrogen. At all N levels, maize had a height advantage by virtue of its larger seed size, which conferred early growth benefits independent of RGR. At low N, biomass growth was instrumental to S. faberi’s improved competitive position, whereas height development per unit biomass improved the competitive position of A. theophrasti, C. album and A. retroflexus. The approach presented could be applied to other crop‐weed systems to evaluate environmental impacts on competitive outcomes.     

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.     

Weed–crop competition relationships differ between organic and conventional cropping systems

Experiments comparing conventional and organic systems often report similar yields despite substantially higher weed abundance in the organic systems. A potential explanation for this observation is that weed–crop competition relationships differ between the two types of systems. We analysed weed and crop yield data from the Rodale Institute Farming Systems Trial (FST), which provides a unique 27-year dataset of a conventional (CNV) and two organic [manure (MNR) and legume (LEG)] soyabean ( Glycine max (L.) Merr.) and maize ( Zea mays L.) cropping systems. Average soyabean yields were similar between the MNR and CNV systems and only slightly reduced in the LEG system, whereas average maize yields did not differ among systems despite the two organic systems having more than four and six times greater weed biomass in soyabean and maize respectively. Plot-level weed biomass–crop yield relationships indicated that weed–crop competition differed between the two organic and CNV systems in maize, and was strongest in the CNV system, intermediate in the LEG system and weakest in the MNR system. These results suggest that organic cropping systems may be able to tolerate a greater abundance of weeds compared to conventional systems and that fertility management within organic systems may influence weed–crop competition.     

Influence of intra-row cruciferous surrogate weed growth on crop yield in organic spring cereals

In Northern Europe, inter-row hoeing has become a popular tactic for controlling weeds in organic cereals. Hoeing is highly effective and can be implemented from crop emergence until stem elongation to maintain a nearly weed-free inter-row zone. However, hoeing has a lesser effect on weeds growing in the intra-row zone, where crop–weed proximity results in heightened competition. In the hoed cereal system, it is investigated whether tall-growing, competitive, cruciferous weeds in the intra-row zone affect crop biomass, yield and thousand kernel weight (TKW). An additive experimental design is employed to enable the fitting of rectangular hyperbolas, describing and quantifying the effects of increasing intra-row surrogate weed density on crop growth parameters. Regressions were studied under the influence of crop (spring barley and spring wheat), row spacing (narrow [12.5 or 15.0 cm] and wide [25.0 cm]) and nitrogen rate (50 and 100 kg NH₄-N/ha). Cruciferous surrogate weeds were found to impact crop yield and quality severely. For example, ten intra-row plants/m² of surrogate weed Sinapis alba reduced grains yields by 7%–14% in spring barley and by 7%–32% in spring wheat with yield losses becoming markedly greater in wheat compared to barley as weed density increases. Compared to wheat, barley limited yield and quality losses and suppressed intra-row weed growth more. Row spacing did not have a consistent effect on crop or weed parameters; in one of six experiments, the 25 cm row spacing reduced yields and increased intra-row weed biomass in wheat. Nitrogen rate did not affect crop or weed parameters. Results warrant the implementation of additional tactics to control intra-row weeds and limit crop losses.     

Competition between annual weeds and vining peas grown at a range of population densities: effects on the weeds

Linear regression of dry weight of weeds against crop density, together with the use of diversity indices and principal component analysis were used to derive information about changes in the behaviour of annual weeds over the growing season and in response to a wide range of crop densities in vining peas Pisum sativum L. Using linear regression it was possible to quantify reductions in weed dry weights per unit increase in crop plant density The ‘weed losse’ factor was acceptably consistent between experiments. Indices examining richness and evenness showed that numbers of weed species declined with increasing crop density and as the season progressed, but although species evenness became less at successive sampling dates the presence of a pea crop, whatever its density, did not radically alter the composition of the weed flora. Principal component analysis demonstrated that although there was competition within the weed flora, the crop did not replace the dominant weed species on high density plots, but reduced growth of all weed species alike.     

Weed communities in intensified cereal-based cropping systems of the northern Guinea savanna

The dynamics of weed populations were ana lysed in intensified cereal-based cropping sys tems of the northern Guinea savanna in Nigeria, A total of four common weed associations were identified through cluster analysis. Five factors describing soil fertility conditions and field his tory best differentiated the weed communities according to a discriminant model. The analysis shows that maize-based cropping systems with a high frequency of cereal cropping and a low fre quency of noncereal cropping tended to be domi nated by weeds such as Commelina spp. and Kyllinga squamulata. As soil fertility declined, Vernonia spp. and Eclipta prostrata became more important. Increased frequency of noncereal crops in mixed cropping with cereals was associ ated with reduced incidence of weeds such as Leucas martinicensis, Oldenlandia corymbosa, Spermacoce verticillata, Litdwigia hyssopifolia, Celosia laxa, and,Ipomoea spp. Further diversifi cation of cereal-based systems to obtain a re duced frequency of cereals is likely to increase the incidence of Dactvloctenium aegyptium in crop fields. The information provides guidance for technology development and transfer on weed control for intensifying systems in the northern Guinea savanna of Africa.     

Crop residue retention suppresses seedling emergence and biomass of winter and summer Australian weed species

Crop residue retention could affect the emergence and biomass of weeds in different ways. A summer and winter pot study was conducted to evaluate the effect of different amounts of sorghum and wheat residue on the emergence and biomass of 12 summer and winter Australian weeds. The equivalent amount of sorghum residue to 0, 1, 2, 4 and 6 t/ha was used in the summer study and winter weed seeds were covered with wheat residue equivalent to the amount of 0, 1, 2, 4 and 8 t/ha in the winter study. The emergence and biomass of Amaranthus retroflexus and Echinochloa colona was not affected by sorghum residue treatment. For other summer weeds, the use of the 6 t/ha sorghum residue treatment resulted in 59–94% reductions in biomass compared to no‐sorghum residue retention. Similarly, the application of 8 t/ha wheat residue in the winter study resulted in a reduced biomass of 15–100% compared to no‐crop residue treatment. The results demonstrated the high potential of using crop residues in eco‐friendly weed management strategies, such as harvest weed seed control tactics.     

Critical period for weed control in sesame production

Planning effective weed control in cropping systems requires exact appraisal of the weed intensity and duration of their competition with the crops. This 2‐year study was carried out in order to determine the critical weed control period in sesame fields. Related and relative crop yields were monitored and analyzed using a four‐parametric log‐logistic model. We recorded data from weed‐free plots and compared these with data from different periods of weed interference. In both the study years, the longer period of weed interference decreased the relative yield of sesame, whereas the yield was increased with increasing duration of the weed‐free period. A 51–78.7% decline in sesame yield was noted if the weeds were allowed to compete with the crop from planting to harvest. In the first year, the duration of the critical period for weed control (CPWC) was 177–820 growing degree days (GDD), which corresponded to 14–64 days after crop emergence (DAE), and between 170 and 837 GDD (13–64 DAE) in the second year; this was based on a 5% acceptable yield loss. The results of this study clearly elaborated that maintaining weed‐free conditions is compulsory from as early as the second week after the emergence of sesame plants, and this should be maintained at least until the ninth week to avoid sesame yield losses by more than 5%. These findings show that growers can benefit from CPWC to improve weed control in sesame production, including the efficacy of a weed control program and its cost.     

Use of perennial cover crops to suppress weeds in Nicaraguan coffee orchards

Both uncontrolled weed growth and vegetation‐free orchard floors have been shown to affect coffee (Coflea arabica L.) negatively, but using cover crops as a solution has yielded conflicting results in different studies. In this study we tested the establishment success of three cover crop species under different management intensities and planting densities, as well as their long term weed‐controlling abilities and effects on weed community composition. Monthly manual weedings during the first 12 weeks after planting resulted in more rapid and extensive cover crop development compared with less intensive management. Transplanted Commelina diffusa Burm. f. grew most rapidly and controlled weeds by limiting light availability, but disappeared during the dry season and failed to establish at all on one of the farms. Arachis pintoi established and persisted for over 2 years, providing excellent weed control by outcompeting weeds for water and/or nutrient resources. Desmodium ovalifolium Wall required the longest time to establish and controlled weeds by an undetermined competitive mechanism. The sowing method of Desmodium led to intense intraspecific competition which probably decreased its effectiveness. Both Arachis and Desmodium led to lower relative abundances of grassy weeds and more perennial forbs, but total weed biomass was so low that these differences have no practical implications.     

Modelling the effects of sub-lethal doses of herbicide and nitrogen fertilizer on crop–weed competition

The effects of sub‐lethal dose of herbicide and nitrogen fertilizer on crop–weed competition were investigated. Biomass increases of winter wheat and a model weed, Brassica napus, at no‐herbicide treatment with increasing nitrogen were successfully described by the inverse quadratic model and the linear model respectively. Increases in weed competitivity (β₀) of the rectangular hyperbola and parameter B in the dose–response curve for weed biomass, with increasing nitrogen were also successfully described by the exponential model. New models were developed by incorporating inverse quadratic and exponential models into the combined rectangular hyperbola with the standard dose–response curve for winter wheat biomass yield and the combined standard dose—response model with the rectangular hyperbola for weed biomass, to describe the complex effects of herbicide and nitrogen on crop–weed competition. The models developed were used to predict crop yield and weed biomass and to estimate the herbicide doses required to restrict crop yield loss caused by weeds and weed biomass production to an acceptable level at a range of nitrogen levels. The model for crop yield was further modified to estimate the herbicide dose and nitrogen level to achieve a target crop biomass yield. For the target crop biomass yield of 1200 g m^?2 with an infestation of 100 B. napus plants m^?2, the model recommended various options for nitrogen and herbicide combinations: 140 and 2.9, 180 and 0.9 and 360 kg ha^?1 and 1.7 g a.i. ha^?1 of nitrogen and metsulfuron‐methyl respectively.     

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.     

The impacts of traditional and novel herbicide application methods on target plants,non‐target plants and production in intensive grasslands

Herbicides contribute significantly to agricultural intensification, but some negatively impact non‐target organisms. Much research has focused on reducing herbicide use through technological improvements in application and herbicide selectivity, but impacts on non‐target organisms are less well understood. Using experimental plots in silage systems, we investigated impacts of herbicides (both narrow spectrum targeting broad‐leaved plants and selective and non‐selective broad spectrum) applied using traditional techniques (blanket‐ and manual spot‐spraying) and a novel application technique (automated spot‐spraying) on non‐target plant richness/diversity, target weed presence (Rumex species) and production (DM yield). All herbicides reduced non‐target plant richness/diversity and sometimes target weeds (when applied using traditional methods). Automated spot‐spraying had fewer negative effects on non‐target organisms, but did not reduce target weeds. No differences in production levels among treatments were observed. The automated spot‐spraying technique requires further research and development. Our results indicate that 20–30% weed cover does not significantly alter production and so, as herbicides are expensive, their effects on non‐target organisms and the environment can be more significant than their benefits to production. We advocate more research into the relationships between weed infestation and production in grasslands, so that the propensity to overuse herbicides is reduced.