High Redundancy as well as Complementary Prey Choice Characterize Generalist Predator Food Webs in Agroecosystems.

Food web structure influences ecosystem functioning and the strength and stability of associated ecosystem services. With their broad diet, generalist predators represent key nodes in the structure of many food webs and they contribute substantially to ecosystem services such as biological pest control. However, until recently it has been difficult to empirically assess food web structure with generalist predators. We utilized DNA-based molecular gut-content analyses to assess the prey use of a set of generalist invertebrate predator species common in temperate agricultural fields. We investigated the degree of specialization of predator-prey food webs at two key stages of the cropping season and analysed the link temperature of different trophic links, to identify non-random predation. We found a low level of specialization in our food webs, and identified warm and cool links which may result from active prey choice or avoidance. We also found a within-season variation in interaction strength between predators and aphid pests which differed among predator species. Our results show a high time-specific functional redundancy of the predator community, but also suggest temporally complementary prey choice due to within-season succession of some predator species.

Diet of generalist predators reflects effects of cropping period and farming system on extra- and intraguild prey.

The suppression of agricultural pests by natural enemies, including generalist arthropod predators, is an economically important regulating ecosystem service. Besides pests, generalist predators may also consume non-pest extraguild and intraguild prey, which can affect their impact on pest populations. This may either reduce the impact of generalist predators on pest populations, because they are diverted from pest predation, or increase it, as it helps them survive periods of low pest availability. However, the availability of pest prey and alternative, non-pest prey can vary over the crop growing season and between farming systems, potentially affecting predator-prey interactions and the levels of biological control. We have limited information about how farming systems and environmental variation over the crop growing season influence predator diets. This limits our ability to predict the importance of generalist predators as natural enemies of agricultural pests. Here we utilize molecular gut content analyses to assess detection frequencies of extra- and intraguild prey DNA in generalist predator communities in replicated organically and conventionally managed cereal fields at two key periods of the cropping season for aphid biological control. This is done in order to understand how farming system, crop season, prey availability and predator community composition determine the composition of predator diets. Aphid pests and decomposers (springtails) were equally important prey for generalist predators early in the growing season. Later in the season, the importance of aphid prey increased with increasing aphid densities while springtail predation rates were positively correlated to abundance of this prey at both early and late crop growth stages. Intraguild predation was unidirectional: carabids fed on spiders, whereas spiders rarely fed on carabids. Carabids had higher detection frequencies for the two most common spider families in organically compared to conventionally managed fields. Our study documents that predation by generalist predator communities on aphid pests increases with pest numbers independently of their generally widespread consumption of alternative, non-pest prey. Therefore, conservation strategies in agricultural fields could promote biological control services by promoting high levels of alternative non-pest prey for generalist predator communities.

Interactive effects of pests increase seed yield.

Loss in seed yield and therefore decrease in plant fitness due to simultaneous attacks by multiple herbivores is not necessarily additive, as demonstrated in evolutionary studies on wild plants. However, it is not clear how this transfers to crop plants that grow in very different conditions compared to wild plants. Nevertheless, loss in crop seed yield caused by any single pest is most often studied in isolation although crop plants are attacked by many pests that can cause substantial yield losses. This is especially important for crops able to compensate and even overcompensate for the damage. We investigated the interactive impacts on crop yield of four insect pests attacking different plant parts at different times during the cropping season. In 15 oilseed rape fields in Sweden, we estimated the damage caused by seed and stem weevils, pollen beetles, and pod midges. Pest pressure varied drastically among fields with very low correlation among pests, allowing us to explore interactive impacts on yield from attacks by multiple species. The plant damage caused by each pest species individually had, as expected, either no, or a negative impact on seed yield and the strongest negative effect was caused by pollen beetles. However, seed yield increased when plant damage caused by both seed and stem weevils was high, presumably due to the joint plant compensatory reaction to insect attack leading to overcompensation. Hence, attacks by several pests can change the impact on yield of individual pest species. Economic thresholds based on single species, on which pest management decisions currently rely, may therefore result in economically suboptimal choices being made and unnecessary excessive use of insecticides.

Insecticide resistance in pollen beetles over 7 years - a landscape approach.

BACKGROUND: In spite of considerable interest in the impact of pesticides on pest populations, few attempts have been made to link resistance patterns of insect pests to land-use features across spatial and temporal scales. We hypothesise that pollen beetle pesticide resistance increases in areas with a high proportion of oilseed rape and with an even mixture of winter and spring oilseed rape owing to high pesticide selection pressure in such areas. RESULTS: Here, we investigated 7 years of lambda-cyhalothrin (Karate(®) ) resistance in field-collected pollen beetle adults from a total of 180 sampling points across ten regions in Sweden. We found a positive effect on pollen beetle pesticide resistance of proportion of oilseed rape and even spring-winter oilseed rape mixture. However, this was true only for the regional spatial scale. Significant land-use effects in the long-term models, with oilseed rape data averaged over a longer (4 years) period of time, suggested an effect of regional landscape history on current pest resistance. CONCLUSION: For successful control of pollen beetle pesticide resistance, we suggest a long-term regional strategy for oilseed rape management. This land-use approach provides a framework for further investigations that integrate resistance management into landscape research.

Management intensity at field and landscape levels affects the structure of generalist predator communities.

Agricultural intensification is recognised as a major driver of biodiversity loss in human-modified landscapes. Several agro-environmental measures at different spatial scales have been suggested to mitigate the negative impact of intensification on biodiversity and ecosystem services. The effect of these measures on the functional structure of service-providing communities remains, however, largely unexplored. Using two distinct landscape designs, we examined how the management options of organic farming at the field scale and crop diversification at the landscape level affect the taxonomic and functional structure of generalist predator communities and how these effects vary along a landscape complexity gradient. Organic farming as well as landscapes with longer and more diversified crop rotations enhanced the activity-density of spiders and rove beetles, but not the species richness or evenness. Our results indicate that the two management options affected the functional composition of communities, as they primarily enhanced the activity-density of functionally similar species. The two management options increased the functional similarity between spider species in regards to hunting mode and habitat preference. Organic farming enhanced the functional similarity of rove beetles. Management options at field and landscape levels were generally more important predictors of community structure when compared to landscape complexity. Our study highlights the importance of considering the functional composition of generalist predators in order to understand how agro-environmental measures at various scales shape community assemblages and ecosystem functioning in agricultural landscapes.

Varied response of Spodoptera littoralis against Arabidopsis thaliana with metabolically engineered glucosinolate profiles.

Upon herbivory glucosinolates are known to be degraded into a cascade of secondary products that can be detrimental for certain herbivores. We performed herbivory bioassays using first and second instar generalist Lepidoptera larvae Spodoptera littoralis on Arabidopsis thaliana engineered to overexpress novel glucosinolates. A differential response in larval feeding patterns was observed on the plants engineered with novel glucosinolates. Larvae fed on plants overexpressing 4-hydroxybenzyl glucosinolate and isopropyl glucosinolate showed little response. Larvae fed on 35S:CYP79A2 plants engineered to overexpress benzyl glucosinolates, however, showed reduced larval and pupal weights. Upon herbivory a high expression of JA signalling gene LOX2 was observed on the 35S:CYP79A2 plants compared to the PR1a and VSP2 expression. To confirm the role of benzyl isothiocyanate (BITC), a degradation product of benzyl glucosinolate overexpressing plants, in the retarded larval growth we used Virus Induced Gene Silencing (VIGS) approach to silence LOX2 expression in the 35S:CYP79A2 plants. S. littoralis larvae fed on LOX2 silenced 35S:CYP79A2 plants exhibited a retarded larval growth thus indicating that BITC played a pivotal role in anti-herbivory and not only the JA signalling pathway.

The influence of metabolically engineered glucosinolates profiles in Arabidopsis thaliana on Plutella xylostella preference and performance.

The oviposition preference and larval performance of the diamondback moth (DBM), Plutella xylostella, was studied using Arabidopsis thaliana plants with modified glucosinolate (GS) profiles containing novel GSs as a result of the introduction of individual CYP79 genes. The insect parameters were determined in a series of bioassays. The GS content of the plants as well as the number of trichomes were measured. Multivariate analysis was used to determine the possible relationships among insect and plant variables. The novel GSs in the tested lines did not appear to have any unequivocal effect on the DBM. Instead, the plant characteristics that affected larval performance and larval preference did not influence oviposition preference. Trichomes did not affect oviposition, but influenced larval parameters negatively. Although the tested A. thaliana lines had earlier been shown to influence disease resistance, in this study no clear results were found for P. xylostella.

Species-specific primers for predation studies of the pollen beetle, Meligethes aeneus (Coleoptera, Nitidulidae).

Species-specific primers were developed for the pollen beetle (a pest in oilseed rape) for studies of predation by natural insect predators. Two forward and three reverse primers were designed within the mitochondrial COI gene and used in combination to amplify fragments in the size range of 163-290 bp. Remains of pollen beetle DNA were consistently detected in Pardosa spiders up to 24 h after ingestion but dropped drastically at 48 h. These primers will facilitate studies on biological control of this oilseed rape pest. Detection time was not correlated with fragment length as might be expected as the DNA gradually degrades into progressively shorter fragments over time.

Spatial occurrence of a habitat-tracking saproxylic beetle inhabiting a managed forest landscape.

Because of the dynamic nature of many managed habitats, proper evaluation of conservation efforts calls for models that take into account both spatial and temporal habitat dynamics. We develop a metapopulation model for successional-type systems, in which habitat quality changes over time in a predictable fashion. The occupancy and recruitment of the predatory saproxylic (dependent on dead wood) beetle Harminius undulatus was studied in a managed boreal forest landscape, covering 24,449 ha, in central Sweden. In a first step, we analyzed the beetle's occupancy pattern in relation to stand characteristics, and the amounts of present and past habitat in the surrounding landscape. Managed forest is suitable habitat when > or =60 years old, and immediately after cutting, but not between the ages of 10 and 60 years. The observed occupancy of H. undulatus was positively correlated with the stand's age as habitat. We used a metapopulation model to predict the current probability of occurrence in each forest stand, given the spatiotemporal distribution of suitable forest stands during the last 50 years. Metapopulation parameters were estimated by matching predicted spatial distributions with observed spatial distributions. The model predicted observed spatial distributions better than a similar model that assumed constant habitat quality of each forest stand. Thus, metapopulation models for successional-type systems, such as dead wood dependent organisms in managed forest landscapes, should include habitat dynamics. An estimated 82% of the landscape-wide recruitment took place in managed stands, which covered 87% of the forest area, in comparison with 18% in unmanaged stands, which covered 13% of the forest area. Among the managed stand types, > or =60-year-old stands and 3-7-year-old clear-cuttings contributed to 79% of the total recruitment while 8-59-year-old stands only contributed 3%. The results suggest the following guidelines to improve conditions for H. undulatus and other species with similar habitat requirements: (1) the proportion of the landscape constituted by younger stands should not be allowed to grow too large, (2) the rotation period of managed stands should not be allowed to be too short, and (3) dead wood should be retained and created at final cutting.

Complex formation of myrosinase isoenzymes in oilseed rape seeds are dependent on the presence of myrosinase-binding proteins.

The enzyme myrosinase (EC 3.2.3.1) degrades the secondary compounds glucosinolates upon wounding and serves as a defense to generalist pests in Capparales. Certain myrosinases are present in complexes together with other proteins such as myrosinase-binding proteins (MBP) in extracts of oilseed rape (Brassica napus) seeds. Immunhistochemical analysis of wild-type seeds showed that MBPs were present in most cells but not in the myrosin cells, indicating that the complex formation observed in extracts is initiated upon tissue disruption. To study the role of MBP in complex formation and defense, oilseed rape antisense plants lacking the seed MBPs were produced. Western blotting and immunohistochemical staining confirmed depletion of MBP in the transgenic seeds. The exclusive expression of myrosinase in idioblasts (myrosin cells) of the seed was not affected by the down-regulation of MBP. Using size-exclusion chromatography, we have shown that myrosinases with subunit molecular masses of 62 to 70 kD were present as free dimers from the antisense seed extract, whereas in the wild type, they formed complexes. In accordance with this, MBPs are necessary for myrosinase complex formation of the 62- to 70-kD myrosinases. The product formed from sinalbin hydrolysis by myrosinase was the same whether MBP was present or not. The performance of a common beetle generalist (Tenebrio molitor) fed with seeds, herbivory by flea beetles (Phyllotreta undulata) on cotyledons, or growth rate of the Brassica fungal pathogens Alternaria brassicae or Lepthosphaeria maculans in the presence of seed extracts were not affected by the down-regulation of MBP, leaving the physiological function of this protein family open.

The pollen beetle, Meligethes aeneus, changes egg production rate to match host quality.

Motivation-based models dominate current theory concerning host plant selection for oviposition by herbivorous insects. A female searching for a host plant will be more likely to accept a host which is of inferior quality for her offspring if motivation is high, e.g. a large eggload or long time since last oviposition. This implies that insects will accumulate eggs if exposed to hosts of low acceptability and after a time lay eggs on such hosts. An alternative strategy for insects when exposed to less acceptable hosts is to stop producing, instead of accumulating, eggs. Thus, resources would be saved until a more acceptable host is found. If this hypothesis is true, a herbivorous insect would cease egg production when exposed to hosts of low acceptability and resume egg production when exposed to hosts of high acceptability. Previous exposure should not affect oviposition rate when an insect encounters a new host of a different quality. In an earlier study pollen beetles, Meligethes aeneus (F.) (Coleoptera: Nitidulidae), did not accumulate eggs in the absence of high quality hosts. In this study we monitored the daily oviposition rate of female pollen beetles on hosts plants of low (Sinapis alba L.), intermediate (Brassica nigra Koch) or high (B. napus L.) acceptability over a 5-day period. Individuals were then switched to an oviposition resource of a different acceptability. Beetles moved from high- to low-acceptability plants reduced their oviposition rate considerably. In the opposite case, low to high acceptability, the rate of oviposition increased markedly after the switch. When M. aeneus were moved from the high-acceptability host to that of intermediate acceptability oviposition rate was modified accordingly. However, when moved to the intermediate host from a host of low acceptability oviposition on B. nigra was much less than would normally be expected. A possible mechanism for this finding is discussed. M. aeneus, by adjusting oviposition rate to host acceptability, maximizes the average host quality for offspring, even at the cost of a lower egg-laying rate.