Biological Control Efficacy Research Articles

Pythium Species and Isolate Diversity Influence Inhibition by the Biological Control Agent Streptomyces lydicus.

Disease control of soilborne pathogens by biological control agents (BCAs) is often inconsistent under field conditions. This inconsistency may be partly influenced by pathogen diversity if there is a differential response among pathogen species and isolates to selected BCAs. The responses of 148 Pythium isolates obtained from soil at three forest nurseries and representative of 16 Pythium spp. were evaluated in the presence of Streptomyces lydicus strain WYEC108 in an in vitro assay. Percent growth inhibition, inhibition zone distance, mortality, and growth rate were recorded for each isolate, and data were analyzed for effects of species and isolate. Responses of three Pythium spp. (Pythium irregulare, P. sylvaticum, and P. ultimum) were further analyzed for a location (nursery) effect. Although S. lydicus inhibited all Pythium isolates, differences in percent growth inhibition, inhibition zone distance, and mortality were observed among Pythium spp. and isolates. Small but significant location effects were also noted. Growth rate also varied among Pythium spp. and isolates and was found to strongly bias percent growth inhibition and, to a lesser degree, inhibition zone distance; depending on which measure was used, slower-growing isolates appeared less sensitive (growth inhibition) or more sensitive (inhibition zone) to S. lydicus than faster-growing isolates. Results illustrate the importance of using multiple, representative pathogen isolates in preliminary BCA inhibition assays as well as accounting for the effect of pathogen growth rate on pathogen inhibition by BCAs. Future studies should take pathogen diversity into account when evaluating biological control efficacy.

Density dependence, precipitation and biological control agent herbivory influence landscape‐scale dynamics of the invasive Eurasian plant Linaria dalmatica

Summary Resource availability and natural enemies are among the most commonly cited mechanisms affecting competitive ability of invasive plants, but their simultaneous effects on plant dynamics are seldom evaluated in the field. Understanding how endogenous and exogenous factors affect invasive plant abundance is essential when evaluating the impact of classical weed biological control agents because misinterpretations of the mechanisms regulating plant demography may bias inference of herbivore impact. In this study, we report results from a citizen‐science monitoring programme initiated to evaluate the effects of the stem‐mining weevil Mecinus janthiniformis on the Eurasian invasive weed Dalmatian toadflax Linaria dalmatica. We used a discrete model of population dynamics to evaluate the relative importance of endogenous and exogenous processes affecting changes in ramet density of L. dalmatica at 38 spatially replicated sites in Idaho, USA. Analysis of per‐capita population growth rates based on ramet density indicated the presence of direct density dependence, which was most likely due to intraspecific competition. Changes in ramet density were not influenced by the abundance of the resident plant community. Precipitation had a strong, positive effect on ramet density of L. dalmatica, while the abundance of M. janthiniformis had a weaker, but significant, negative effect after accounting for the density dependence. There was no support for an interactive effect between precipitation and herbivory. Synthesis and applications. Our results indicate that biological control is an important factor affecting weed population growth at the landscape scale, but they also suggest that biological control impact may vary considerably on local infestations due to site‐specific variation in rainfall and density‐dependent processes. We recommend that invasive plant management strategies integrate precipitation and biological control agent monitoring into their programmes to estimate expected biological control efficacy. Alternative control methods should be prioritized in areas where herbivore impact is expected to be low.

Bottom-up effects on top-down regulation of a floating aquatic plant by two weevil species: the context-specific nature of biological control

Summary Plant nutrition (bottom-up effects) impacts a plant's ability to withstand herbivory (top-down effects) and influences phytophagous insect fecundity. These factors confound efficacy predictions for biological control purposes. We investigated the importance of these forces with regard to invasibility of Eichhornia crassipes. We also examined whether one could have predicted whether the weevil species Neochetina eichhorniae or Neochetina bruchi would have been the better biocontrol agent. Three experiments compared E. crassipes performance when subjected to herbivory by these weevils across a range of fertilizer levels. A low initial plant density experiment examined growth and flowering while allowing density to increase. A high plant density experiment evaluated reduction in biomass and surface coverage. A third experiment evaluated plant growth unconstrained by space. Herbivory effects varied between treatments; interactions varied between experiments. Herbivory reduced plant density and biomass, except in low fertilizer conditions, and consistently reduced flowering. N. bruchi alone or both weevil species together reduced growth more than N. eichhorniae alone, elucidating N. bruchi as the better agent. However, N. eichhorniae better reduced biomass and plant density under initial conditions of complete plant coverage, suggesting it as the superior choice. Synthesis and applications. Bottom-up and top-down forces acting in concert affect invasibility of plant species. Plasticity to resource availability enables invaders to persist in conditions that are unfavourable to herbivores (enemy-free space). Consequently, biological control efficacy is nuanced and context specific, so predictability requires assessment of multiple parameters across a range of conditions. Overly simplistic evaluations risk rejection of effective agents capable of mediating non-native plant invasions. These results also argue that biological control practitioners should disregard the concept that a single best agent can be identified to control an invasive plant that exploits a broad range of habitats and environmental conditions. Aquaphyte invasions often coincide with watershed nutrification so unless weed control is carried out concurrently with measures that reduce nutrient inputs, removal of infestations will merely reduce competition thereby accelerating regrowth or facilitating replacement by other undesirable species (the ‘invasive treadmill’). The suppressive effects of biological control are valuable and complimentary, even when replacement by other, presumably less problematic, species may occur. These subtler effects should be considered with protocols developed for integrated control strategies.

Current levels of suppression of waterhyacinth in Florida USA by classical biological control agents

Waterhyacinth, Eichhornia crassipes (Mart.) Solms (Pontederiaceae), has been a global target for classical biological control efforts for decades. In Florida, herbicidal application is the primary control method employed, usually without regard for the activities of the three biological control agents introduced intentionally during the 1970s, namely Neochetina eichhorniae Warner, Neochetina bruchi, Hustache (Coleoptera: Curculionidae), and Niphograpta albiguttalis Warren (Lepidoptera: Crambidae). A series of field experiments from 2008 to 2010 was conducted at four Florida sites using an insecticide-check approach to quantify the current levels of suppression provided by these agents. In the field N. albiguttalis was rarely found while more than 99% of all Neochetina sp. adults were N. eichhorniae. Although it was not possible to disentangle the relative impacts of Neochetina sp. adults from larvae on individual plant variables, the larvae played a major role in reducing plant biomass and the number of inflorescences. Plots exposed to unrestricted herbivory contained 58.2% less biomass and produced 97.3% fewer inflorescences at the end of the experiments. Despite these large reductions, herbivory decreased waterhyacinth coverage by only 16.8% and most of this was attributed to a low-nutrient site where coverage was reduced disproportionately. Overall, coverage trended upwards during the course of the experiments and was always close to 100% when the plots were harvested. Although coverage is a somewhat arbitrary metric, especially for floating plants subject to compression and dispersion, it influences the perception of biological control efficacy which, in turn, directly influences herbicide management decisions in Florida. Despite waterhyacinth populations that now produce less than half as much biomass and up to 98% fewer seeds than before the deployment of biological control agents, the overall approach used to achieve maintenance control of the plant in Florida will probably not change unless new biological control agents, such as Megamelus scutellaris Berg (Hemiptera: Delphacidae), can reduce coverage significantly.

Establishment, Population Increase, Spread, and Ecological Host Range ofLophodiplosis trifida(Diptera: Cecidomyiidae), a Biological Control Agent of the Invasive TreeMelaleuca quinquenervia(Myrtales: Myrtaceae)

The Australian tree Melaleuca quinquenervia (Cavanilles) Blake is an invasive weed in wetland systems of Florida. A biological control program targeting M. quinquenervia has resulted in the release of the gall forming midge Lophodiplosis trifida Gagné (Diptera: Cecidomyiidae). Populations of the introduced herbivore readily established at all 24 release sites across the weed's range in Florida, and there was no evidence that founding colony size (100, 2,000, or 6,000 adults) influenced herbivore establishment or local population growth rates. Landscape level spread of L. trifida from release sites averaged nearly 6 km/yr, ranging as high as 14.4 km/yr. Prerelease host range testing predicted that L. trifida oviposits indiscriminately on test plant species but does not complete development on any of the test species, including congeners present in Florida. To test the predictability of these host range tests, L. trifida was released in a common garden consisting of 18 test plant species that were interplanted with M. quinquenervia. Plant species postulated to be at risk experienced no gall development by L. trifida while intermingled M. quinquenervia trees supported 704.8 (± 158.5) galls per plant. Historically, many introduced Cecidomyiidae have limited effect on plant performance of target weeds because of recruitment of native parasitoids that disrupt biological control efficacy. In contrast to this trend, there has been no evidence to date that parasitoids are exploiting L. trifida in Florida.

Economic Thresholds in Soybean-Integrated Pest Management: Old Concepts, Current Adoption, and Adequacy.

Increasing global demands for food underline the need for higher crop yields. The relatively low costs of the most commonly used insecticides in combination with increasing soybean market prices led growers and technical advisors to debate the adequacy of recommended economic thresholds (ETs). The adoption of ETs and pest sampling has diminished in Brazil, leading to excessive pesticide use on soybean. The reduced efficacy of natural biological control, faster pest resurgence, and environment contamination are among the side-effects of pesticide abuse. To address these problems and maximize agricultural production, pest control programs must be guided by a proper integrated pest management (IPM) approach, including the ET concept. Therefore, the most appropriate time to initiate insecticide spraying in soybean is indicated by the available ETs which are supported by experiments over the last 40years in different edapho-climatic conditions and regions with distinct soybean cultivars. Published scientific data indicate that preventive insecticide use is an expensive and harmful use of chemicals that increases the negative impact of pesticides in agroecosystems. However, the established ETs are for a limited number of species (key pests), and they only address the use of chemicals. There is a lack of information regarding secondary pests and other control strategies in addition to insecticides. It is clear then that much progress is still needed to improve ETs for pest management decisions. Nevertheless, using the current ETs provides a basis for reducing the use of chemicals in agriculture without reducing yields and overall production, thereby improving sustainability.

Characterizing the mode of action of Brevibacillus laterosporus B4 for control of bacterial brown strip of rice caused by A. avenae subsp. avenae RS-1

Biological control efficacy of Brevibacillus laterosporus B4 associated with rice rhizosphere was assessed against bacterial brown stripe of rice caused by Acidovorex avenae subsp. avenae. A biochemical bactericide (chitosan) was used as positive control in this experiment. Result of in vitro analysis indicated that B. laterosporus B4 and its culture filtrates (70%; v/v) exhibited low inhibitory effects than chitosan (5 mg/ml). However, culture suspension of B. laterosporus B4 prepared in 1% saline solution presented significant ability to control bacterial brown stripe in vivo. Bacterization of rice seeds for 24 h yielded a greater response (71.9%) for controlling brown stripe in vivo than chitosan (56%). Studies on mechanisms revealed that B. laterosporus B4 suppressed the biofilm formation and severely disrupted cell membrane integrity of A. avenae subsp. avenae, causing the leakage of intracellular substances. In addition, the expression level of virulence-related genes in pathogen recovered from biocontrol-agent-treated plants showed that the genes responsible for biofilm formation, motility, niche adaptation, membrane functionality and virulence of A. avenae subsp. avenae were down-regulated by B. laterosporus B4 treatment. The biocontrol activity of B. laterosporus B4 was attributed to a substance with protein nature. This protein nature was shown by using ammonium sulfate precipitation and subsequent treatment with protease. The results obtained from this study showed the potential effectiveness of B. laterosporus B4 as biocontrol agent in control of bacterial brown stripe of rice.

Effects of inter-annual landscape change on interactions between cereal aphids and their natural enemies

The agricultural intensification and the subsequent habitat changes in agroecosystem can strongly affect biological control services. We here examine the influence of inter-annual landscape change in wheat field area on interactions of cereal aphids and their natural enemies, as well as the efficacy of biological control using data collected from a 4-year experiment in Northwest China. Two hypotheses were tested. (i) Population densities decline following an inter-annual expansion of wheat crop proportion cover due to dilution and crowding effects. (ii) Species that are specialists or at higher trophic levels are more sensitive to bottom-up disturbance by inter-annual change in percent cover of wheat crop. Results showed the population densities of one cereal aphid (Macrosiphum avenae), one parasitic wasp (Aphidius avenae), two specialist predators (ladybirds: Hippodamia variegata and H. tredecimpunctata) and one hyperparasitic wasp (Pachyneuron aphidis) declined following the expansion of wheat crop areas, supporting the predictions of inter-annual dilution and crowding effects. In contrast, the populations of one cereal aphid (Schizaphis graminum), one parasitic wasp (A. gifuensis), two generalist predators (spiders: Pardosa astrigera; carabid beetles: Chlaenius pallipes), and two hyperparasitic wasps (Asaphes suspensus, and Alloxysta sp.) did not respond to inter-annual landscape change. The two hypotheses were partially supported but with noticeable exceptions, and the bio-control efficiency declined with the increase of the proportion cover of wheat field in agricultural landscape. Overall, different responses of cereal aphids and their natural enemies make it difficult but still possible to optimize inter-annual landscape change for enhancing the parasitism rate and predator-prey ratio.

Tritrophic effects of plant growth regulators in an aphid-parasitoid system

Plant growth regulators (PGRs) have the potential to negatively affect the outcome of biological control via plant architectural changes and plant chemical changes. Despite studies demonstrating the negative effects of PGRs on herbivore survival and development, to date, no studies have investigated the tritrophic effects of PGRs on parasitoid life history traits. In this study we investigated the effect of four commonly used PGRs on Myzus persicae abundance and suppression, and Aphidius colemani fitness in a greenhouse experiment. None of the PGRs reduced aphid abundance alone or affected aphid suppression by A. colemani. However, paclobutrazol reduced the number of mummies that developed on plants. PGRs had a range of negative effects on parasitoid fitness. No adult parasitoids eclosed from mummies on ancymidol treated plants. Paclobutrazol reduced parasitoid size, and paclobutrazol and uniconazole reduced female:total ratio. This study shows that PGRs can negatively affect parasitoid fitness, and reduce parasitism, suggesting the potential for negative long-term effects on the efficacy of biological control.

Compact plants reduce biological control of Myzus persicae by Aphidius colemani

Common horticultural practices, such as the use of plant growth regulators, may negatively influence the outcome of biological control programs. Plant growth regulators are applied to many ornamental and agricultural crops and can result in compact plants that have more branches and are bushier than untreated plants. Since plant architectural complexity can have strong effects on natural enemy foraging efficiency and pest suppression, our hypothesis was that the use of plant growth regulators would reduce aphid suppression by the parasitoid Aphidius colemani. In this study we investigated how the plant growth regulator paclobutrazol and the parasitic wasp A. colemani interact to affect the abundance and behavior of Myzus persicae. We found that paclobutrazol alone reduced aphid abundance compared to untreated plants. However, when parasitoids were present, paclobutrazol and associated changes in plant architecture reduced parasitism and increased aphid abundance compared to untreated plants. A likely mechanism for this result is that significantly more M. persicae fed in concealed locations on paclobutrazol-treated plants than on untreated plants. This study demonstrates that paclobutrazol reduced the efficacy of biological control by A. colemani and suggests that plant growth regulators could also affect biological control of other organisms.

Interactions betweenMeteorus pulchricornisandSpodoptera exiguaMultiple Nucleopolyhedrovirus

Baculoviruses may interact with parasitoids in the same host. A previous study has shown that infection of larvae with Spodoptera litura nucleopolyhedrovirus (SpltNPV) was deleterious to the survival and development of Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae). In this paper, the interactions between M. pulchricornis and Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV) in Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), a permissive host of the virus and parasitoid, were investigated. The results showed that the effect of M. pulchricornis on SeMNPV and the effect of the virus on the parasitoid both depended on the concentration of the virus and the interval between viral infection and parasitism. Whether S. exigua was treated with the parasitoid and virus simultaneously or 1 day apart, the biological activities of 105, 106, and 107 OBs/mL SeMNPV were all significantly improved by M. pulchricornis. In contrast, the biological activity of 103 OBs/mL SeMNPV was significantly decreased when the host was exposed to the virus and parasitoid simultaneously. Regarding the impact of SeMNPV on M. pulchricornis, exposing the host to the parasitoid and SeMNPV with concentrations lower than 106 occlusion bodies (OBs)/mL produced no negative effects on the parasitoid. The results also showed that ingestion of SeMNPV by adult stage M. pulchricornis significantly increased the number of parasitoid offspring that successfully emerged from the host. Furthermore, M. pulchricornis was found to transmit SeMNPV among populations of S. exigua. Taken together, these findings indicate that M. pulchricornis integrated with an appropriate concentration of SeMNPV has the potential to improve the efficacy of biological control against S. exigua.

Effect of elevated CO2 on tropical soda apple and its biological control agent Gratiana boliviana (Coleoptera: Chrysomelidae)

Rising atmospheric CO2 levels could have drastic effects on the performance of invasive weeds and their insect herbivores. Despite the importance of biological control as an effective management tool for environmental weeds, there have been few studies on the potential impact of climate change on the future efficacy of biological control. The objective of the present study was to evaluate the effect of elevated CO2 on the performance of tropical soda apple Solanum viarum (Solanaceae) and its biological control agent Gratiana boliviana (Coleoptera: Chrysomelidae). We established three levels of CO2 in environmental growth chambers: ambient (400 ppm), medium (580 ppm) and high (780 ppm). Plants growing at the high level had greater biomass compared to those growing at the ambient and medium levels of CO2. Leaf water content and the amount of leaf nitrogen were reduced at high compared to ambient or medium CO2 levels. G. boliviana immature survival and developmental time were negatively affected at high CO2 but not at medium or ambient levels. Adults were lighter and smaller when reared at the high CO2 level compared to ambient and medium treatments, while adult fecundity was higher at the medium CO2 level. Leaf area consumed by fifth instars was lower when feeding on plants grown at the high CO2 level either inside a Petri dish or on potted plants. These results suggest that beetle performance may be diminished under future climate. However, further studies should incorporate other factors such as temperature and precipitation as well as the evolutionary potential of herbivores and plants to adapt to a changing climate.

Designing the ideal habitat for entomopathogen use in nursery production

Greenhouse and nursery producers use entomopathogens (nematodes and fungi) to control soil pests. Although it is known that the physical and chemical properties of mineral soil significantly impact upon soil pathogens, the influence of soilless media used for plant production on entomopathogen performance is poorly understood. Survival and foraging distance were differently affected by sand:peat, bark and sawdust media for entomopathogenic nematodes, but not for the immobile fungus Metarhizium anisopliae. Redwood sawdust medium consistently had a negative impact upon entomopathogenic nematodes. Dividing media into individual components supported the hypothesis that redwood sawdust reduced foraging and infection abilities of S. riobrave and H. bacteriophora. Physically altering the components by adding sand significantly improved foraging and infection success for S. riobrave in media not optimum for foraging. This study is the first to highlight the importance of selecting the appropriate soilless media and pathogen species combinations to increase efficacy of biological control. H. bacteriophora was able to find hosts in a wider diversity of medium components than S. riobrave, although both nematode species performed well in peat moss and recycled plant material. These results suggest that peat moss, recycled plant material and hardwood bark are components amenable to EPN biological control programs.

Bacterial Mixture from Greenhouse Soil as a Biocontrol Agent Against Root-Knot Nematode, Meloidogyne incognita, on Oriental Melon

The biological control efficacy of a greenhouse soil bacterial mixture of Lactobacillus farraginis, Bacillus cereus, and Bacillus thuringiensis strains with antinematode activity was evaluated against the root-knot nematode Meloidogyne incognita. Two control groups planted in soil drenched with sterile distilled water or treated with the broadspectrum carbamate pesticide carbofuran were used for comparison. The results suggest that the bacterial mixture is effective as a biocontrol agent against the root-knot nematode.

Electrostatic application of antimicrobial sprays to sanitize food handling and processing surfaces for enhanced food safety

Human illnesses and deaths caused by foodborne pathogens (e.g., Salmonella enterica, Listeria monocytogenes, Escherichia coli O157:H7, etc.) are of increasing concern globally in maintaining safe food supplies. At various stages of the food production, processing and supply chain antimicrobial agents are required to sanitize contact surfaces. Additionally, during outbreaks of contagious pathogenic microorganisms (e.g., H1N1 influenza), public health requires timely decontamination of extensive surfaces within public schools, mass transit systems, etc. Prior publications verify effectiveness of air-assisted, induction-charged (AAIC) electrostatic spraying of various chemical and biological agents to protect on-farm production of food crops…typically doubling droplet deposition efficiency with concomitant increases in biological control efficacy. Within a biosafety facility this present work evaluated the AAIC electrostatic-spraying process for application of antimicrobial liquids onto various pathogen-inoculated food processing and handling surfaces as a food safety intervention strategy. Fluoroanalysis of AAIC electrostatic sprays (−7.2 mC/kg charge-to-mass ratio) showed significantly greater (p<0.05) mass of tracer active ingredient (A.I.) deposited onto target surfaces at various orientations as compared both to a similar uncharged spray nozzle (0 mC/kg) and to a conventional hydraulic-atomizing nozzle. Per unit mass of A.I. dispensed toward targets, for example, A.I. mass deposited by AAIC electrostatic sprays onto difficult to coat backsides was 6.1-times greater than for similar uncharged sprays and 29.0-times greater than for conventional hydraulic-nozzle sprays. Even at the 56% reduction in peracetic acid sanitizer A.I. dispensed by AAIC electrostatic spray applications, they achieved equal or greater CFU population reductions of Salmonella on most target orientations and materials as compared to uncharged sprays and conventional full-rate hydraulic-nozzle sprays.

Plant versus prey resources: Influence on omnivore behavior and herbivore suppression

Conservation biological control tactics, such as beetle banks, that increase habitat complexity generally increase epigeal predator abundance. Habitat complexity also increases alternative food which can attract and sustain predators but may reduce predation of target pests. Our goal was to determine how alternative food from different trophic levels (fly pupae and seeds) affects behavior and biological control efficacy of omnivorous carabid beetles. Seed subsidies increased omnivorous carabid abundance more than pupae by increasing aggregation and reducing emigration. Laboratory experiment demonstrated that both omnivorous carabid species preferred seeds and pupae over cutworms. However, in field cages seeds but not pupae resulted in greater cutworm damage to corn seedlings. Our results indicate that omnivorous carabids have a stronger behavioral response to seeds than prey such that only seeds influence aggregation, emigration, and crop damage. Interestingly, whereas seeds increased omnivorous carabid abundance, pupae had no affect on carnivore abundance. Thus, carabid guild composition is skewed in favor of omnivores when seed density increases. An important finding was that the effect of seeds on behavior, predation, and crop damage was consistent among replicate carabid species suggesting our results pertain to other omnivorous species in resource diverse habitats. Our results provide insight into the mechanisms underlying the unpredictable benefit of conservation biological control tactics that alter habitat complexity.

Searching behavior of an aphid parasitoid and its hyperparasitoid with and without floral nectar

While enhancing the searching ability of a natural enemy can give a major improvement in host population suppression, it is important to demonstrate selectivity in favor of the third trophic level when changes, such as added floral resources, are made to agro-ecosystems to enhance biological control efficacy. To explore this question, a pea-aphid endoparasitoid, Aphidius ervi, and its hyperparasitoid, Dendrocerus aphidum, were used to test the effects of prior feeding with buckwheat nectar on searching behavior of both the parasitoid and hyperparasitoid. Digital video was used to record behavior and data were scored and analyzed using Observer Video-Pro software. A. ervi females in nectar treatments spent a higher proportion of their time actively searching than those with access to water only. Fed parasitoids antennated, stung and groomed significantly more than unfed female parasitoids, and spent almost twice as much time attacking and carried out nearly twice as many attacks as unfed parasitoids. D. aphidum spent the greatest proportion of time attacking potential hosts in both ‘fed’ and ‘unfed’ treatments. However, although fed females spent a significantly longer time antennating, there was no significant difference in walking or grooming between treatments. Unfed female hyperparasitoids spent significantly more time stationary than fed females. The results demonstrate that the searching efficiency of both third and fourth trophic level parasitoids is enhanced by access to floral nectar. Nectar provision can therefore potentially have positive or negative effects on biological control, depending on the relative proportion of each species.

Spatial and temporal correspondence of sunn pest oviposition and egg parasitism by Trissolcus spp. in Iran

The sunn pest, Eurygaster integriceps Puton, is a serious pest of wheat and other cereals in the Firouzabad region of Ardabil Province, Iran. In this study, we examined the spatial distribution of sunn pest eggs and their parasitism by Trissolcus spp., primarily Trissolcus grandis Thomson, during the growing seasons of 2008 and 2009. Ten quadrats of 1 m2 were established in each of the ten experimental fields and sampled every 3 days beginning at Feekes growth stage 6 (beginning of stem elongation) through stage 11 (ripening). The numbers of sunn pest eggs, both parasitized and unparasitized, were tallied in each quadrat on each sampling date. Sunn pest oviposition peaked during the onset of flowering in both years, while parasitism peaked at the completion of anthesis in 2008, and at anthesis complete to early milk stage in 2009. Iwao’s regression provided a better fit to the data than did Taylor’s power law. Both models indicated an aggregated distribution of eggs and parasitism. Temporally, there was a negative overlap between egg density and parasitism from inflorescence emergence until the beginning of anthesis, and positive overlap from anthesis complete until the soft dough stage, such that parasitism ranged from 0% of eggs laid early, to virtually 100% of eggs laid late. Thus, despite good spatial correspondence between egg density and egg parasitism, the time lag between peak bug oviposition and peak parasitism enabled large numbers of early-laid host eggs to hatch and diminished the potential efficacy of biological control by Trissolcus spp. The implications of these results are discussed in the context of integrated management of sunn pest in wheat fields and the possible benefits of early season augmentation of the parasitoid population.

Control of Fire Blight by Pseudomonas fluorescens A506 and Pantoea vagans C9-1 Applied as Single Strains and Mixed Inocula

The biological control agents Pseudomonas fluorescens A506 and Pantoea vagans C9-1 were evaluated individually and in combination for the suppression of fire blight of pear or apple in 10 field trials inoculated with the pathogen Erwinia amylovora. The formulation of pathogen inoculum applied to blossoms influenced establishment of the pathogen and the efficacy of biological control. Pantoea vagans C9-1 suppressed fire blight in all five trials in which the pathogen was applied as lyophilized cells but in none of the trials in which the pathogen was applied as freshly harvested cells. In contrast, Pseudomonas fluorescens A506 reduced disease significantly in only one trial. A mixture of the two strains also suppressed fire blight, but the magnitude of disease suppression over all field trials (averaging 32%) was less than that attained by C9-1 alone (42%). The two biological control agents did not antagonize one another on blossom surfaces, and application of the mixture of A506 and C9-1 to blossoms resulted in a greater proportion of flowers having detectable populations of at least one bacterial antagonist than the application of individual strains. Therefore, the mixture of A506 and C9-1 provided less disease control than expected based upon the epiphytic population sizes of the antagonists on blossom surfaces. We speculate that the biocontrol mixture was less effective than anticipated due to incompatibility between the mechanisms by which A506 and C9-1 suppress disease.

A Comparative Study on the Functional Response ofWolbachia-Infected and Uninfected Forms of the Parasitoid WaspTrichogramma brassicae

Trichogramma species (Hymenoptera: Trichogrammatidae) are haplo-diploid egg parasitoids that are frequently used as biological control agents against lepidopteran pests. These wasps display two reproductive modes, including arrhenotoky (bisexuality) and thelytoky (unisexuality). Thelytokous forms are often associated with the presence of endosymbiotic Wolbachia bacteria. The use of thelytokous wasps has long been considered as a way to enhance the efficacy of biological control. The present study investigates the potential of a thelytokous Wolbachia-infected and an arrhenotokous uninfected Trichogramma brassicae Bezdenko strain as inundative biocontrol agents by evaluating their functional response towards different egg densities of the factitious host, the Angoumois grain moth, Sitotroga cerealella (Olivier) (Lepidoptera: Gelechiidae). The results revealed a type II functional response for both strains in which parasitism efficiency decreases with host egg density because of an increasing host handling time. A model with an indicator variable was used to compare the parameters of Holling's disc equation in different data sets. It was demonstrated that the two strains did not differ in host attack rate. However, the Wolbachia-infected strain did have an increased host handling time when compared to the bisexual strain. Some applied aspects of the findings are discussed.