Meloidogyne incognita Survival in Soil Infested with Paecilomyces lilacinus and Verticillium chlamydosporium

Meloidogyne incognita-infected tomato seedlings were transplanted into sterilized soil or unsterilized soil collected from 20 California tomato fields to measure suppression caused by Paecilomyces lilacinus, Verticillium chlamydosporium, and other naturally occurring antagonists. Unsterilized soils Q, A, and H contained 35, 39, and 55% fewer M. incognita second-stage juveniles (J2) than did sterilized soil 1 month after infected tomato seedlings were transplanted to these soils and placed in a greenhouse. Three months after infected seedlings were transplanted to unsterilized or sterilized soil, unsterilized soils K, L, and Q had 97, 62, and 86% fewer J2 than the corresponding sterilized soils. Unsterilized soils of M. incognita-infected seedlings that were maintained 1 month in a greenhouse followed by 1 or 2 months of post-harvest incubation contained J2 numbers equal to, or greater than, numbers in the corresponding sterilized soil. The most suppressive of the unsterilized soils, K and Q, were not infested with V. chlamydosporium. Paecilomyces lilacinus and V. chlamydosporium increased in colony forming units in unsterilized soil of all bioassays, but they were not associated with lower numbers of J2.     

Association of Verticillium chlamydosporium and Paecilomyces lilacinus with Root-knot Nematode Infested Soil

Population densities of Meloidogyne incognita and the nematophagous fungi, Paecilomyces lilacinus and Verticillium chlamydosporium, were determined in 20 northern California tomato fields over two growing seasons. Paecilomyces lilacinus was isolated from three fields, V. chlamydosporium was isolated from one field, and both fungi were isolated from 12 fields. Verticillium chlamydosporium numbers were positively correlated with numbers of M. incognita and P. lilacinus. Paecilomyces lilacinus numbers were positively correlated with V. chlamydosporium numbers, but they did not correlate with M. incognita numbers. The correlation coefficients were low (R < 0.5) but significant (P < 0.05). All P. lilacinus and V. chlamydosporium field isolates parasitized M. incognita eggs in vitro. In a greenhouse study, numbers of V. chlamydosporium and P. lilacinus increased more in soils with M. incognita-infected tomato plants than in soil with uninfected tomato plants. After 10 weeks, the Pf/ Pi of second-stage juveniles in soils infested with P. lilacinus, V. chlamydosporium, and M. incognita was 47.1 to 295.6. The results suggest V. chlamydosporium and P. lilacinus are not effectively suppressing populations of M. incognita in California tomato fields.     

Quantitative assessment of the interaction between the antagonistic fungus Talaromyces flavus and the wilt pathogen Verticillium dahliae on eggplant roots

Quantitative aspects of the interaction between the antagonist Talaromyces flavus, the pathogen Verticillium dahliae and eggplant roots, were studied. When eggplant roots were inoculated with T. flavus, prior to the infection with the pathogen, the population density of T. flavus on V. dahliae-infected roots was at least 3 times higher than on healthy uninfected roots, and the proliferation of T. flavus on diseased eggplant roots was related to the severity of wilt symptoms, in the two levels of application of T. flavus studied. However, in all classes of disease severity tested (disease index, 0–3), the population density of T. Flavus on eggplant roots treated with 10⁶ ascospores g^–1 rooting mixture was significantly (p=0.05) higher than with 10⁵ ascospores g^–1. In roots treated with 10⁵ and 10⁶T. flavus ascospores g^–1 rooting mixture, the population density of V. dahliae was reduced by 51% and 69%, respectively. When testing the relationships between the population density of V. dahliae in the roots and disease severity, no significant (p=0.05) difference was found between disease indexes 2 and 3. However, the density of V. dahliae on roots of plants with disease index 1 was significantly (p=0.05) lower than disease indexes 2 and 3. The positive relationship between the inoculum concentration of V. dahliae and the population density of T. flavus developed on eggplant roots was significant (p=0.001), linear, and highly correlated (r=0.945) on a logarithmic scale. In addition, the analysis of these data revealed a significant (p=0.05), high, negative and linear correlation (r=–0.985) between the log concentration of V. dahliae inoculum and the disease reduction achieved by T. flavus.     

Growth and survival of Verticillium chlamydosporium goddard,a parasite of nematodes,in soil

Selective media for the isolation of the nematophagous fungus Verticillium chlamydosporium, are described. These enabled densities > 500 colony forming units (CFU) g^‐1 soil to be reliably estimated. However, there was little relationship between estimates of the Verticillium biomass in a sterilized soil and the numbers of CFU which developed on the selective media. The growth and survival of the fungus infield soils were studied and estimates of the numbers of CFU in soils in which cyst‐nematode multiplication was suppressed were greater than in those in which the nematode multiplied. Isolates of the fungus differed in their ability to proliferate in soil, but some increased rapidly from applications of chlamydospores or a mixture of hyphae and conidia in alginate granules containing wheat bran. The energy source (wheat bran) was essential for the establishment of the fungus from granular applications. Numbers of CFU greatly exceeded those of chlamydospores, but there was considerable variation in the relationship in different soils. Some isolates of V. chlamydosporium proliferated in soil and survived in considerable numbers for at least 3 months. Hence, pre‐cropping applications of the fungus should survive long enough to kill nematode eggs and females that develop on roots of spring‐sown crops.     

Methods for studying the nematophagous fungus Verticillium chlamydosporium in the root environment

In order to exploit fully the biocontrol potential of the nematophagous fungus Verticillium chlamydosporium, it is important to understand the ecology of the fungus in soil, and interactions with both plant and nematode hosts. Several approaches for studying the fungus in soil and the root environment are compared. These include a semi-selective medium for V. chlamydosporium, PCR primers specific for the fungal -tubulin gene, and monoclonal antibodies. In addition to providing a target for species-specific primers, the -tubulin gene is implicated in resistance to the fungicides used in the semi-selective medium, and the genetic basis for this is investigated. Culture and PCR-based methods were used to screen for the presence of the fungus in field soils known to have been suppressive to cereal cyst nematode and that contained V. chlamydosporium populations. Monoclonal antibodies specific for either hyphae or conidia of the fungus were obtained, and their application as a tool for visualising the infection process on the root was explored.     

Relationship between agitation speed and the morphological characteristics of Verticillium lecanii CS-625 during spore production

Verticillium lecanii is recognized as an entomopathogenic fungus, and has high potential in the biological control of pests. In this study, it was investigated that the relationship between agitation speed in a 2.5 L stirred tank reactor (STR) at 25°C and initial pH 5.5, and the morphological characteristics of V. lecanii CS-625, such as hyphal length/width, spore length/width, and the number of tips during spore production. The agitation speed affected the hyphae patterns and the number of tips. The number of spores rapidly increased at 48 to 60 h of cultivation, and the highest spore productivity (2.5 × 10¹⁰ spore/L·h at 60 h) occurred with an agitation speed of 350 rpm and an aeration rate of 1.0 vvm. The number of tips increased in proportion to the increase in spore production during the same culture time. The highest number of tips (4.8 × 10⁸ tipJ.mL) was obtained at 72 h of cultivation. The shortest mean spore length (2.8 μm) was obtained at 60 h of cultivation. Therefore, it was determined that the increased number of tips and decreased mean spore length were closely related to the production of V. lecanii spores.     

Interactions between iron nutrition and Verticillium wilt resistance in tomato

The relationship between Fe nutritional status and Verticillium wilt disease in tomatoes possessing single gene resistance to Race 1 of Verticillium dahliae was investigated using hydroponic culture media. Iron limiting conditions increased the sensitivity of resistant tomatoes to the pathogen as expressed by wilting and chlorosis. Distance of fungal vascular invasion was approximately the same in both Fe replete and Fe limited treatments. Comparison of near-isolines revealed that the magnitude of disease expressed in Fe deficient Pixie II (resistant) was considerably less than that expressed by the susceptible Pixie variety. Infection of tomato did not enhance the severity of low-Fe stress as quantified by root peroxidase activity and chlorophyll content of young leaves.     

Environment factors influence in vitro interspecific interactions between A. ochraceus and other maize spoilage fungi,growth and ochratoxin production

The effect of water availability (water activity,a_w; 0.995–0.90) and temperature (18–30 °) on in vitro interactions between an ochratoxin producing strain of Aspergillus ochraceus and six other spoilage fungi was assessed in dual culture experiments on a maize meal-based agar medium. Inprimary resource capture of nutrient substrate, A. ochraceus was dominant against many of the interacting species, being able to overgrow and replace A. candidus, and sometimes A. flavus and the Eurotium spp. regardless of a_w or temperature. However, with freely available water (0.995 a_w) A. alternata and A. niger were dominant, with mutual antagonism between A. ochraceus and A. flavus at 25–30 °C. In the driest conditions tested (0.90 a_w) there was also mutual antagonism between A. ochraceus and the two Eurotium spp. Overall, under allconditions tested the Index of Dominance for A. ochraceus was much higher than for other competing species combined suggesting that A. ochraceus wasa good competitive colonist able to replace a numberof other species. However, the growth rate ofA. ochraceus was modified and decreased by the interaction with competitors. Interaction between A. ochraceus and species such as A. alternata (18°C/0.995) and Eurotium spp. (0.995–0.95 and 25–30 °C) resulted in a significant stimulation of ochratoxin production. Theresults are discussed in relation to the effect that environmental factors have on the possible competitiveness of A. ochraceus in the maizegrain ecosystem and the role of ochratoxin in nicheexclusion of competitors. This revised version was published online in June 2006 with corrections to the Cover Date.     

Microscopic observations on the interaction of the mycoparasite Verticillium biguttatum with Rhizoctonia solani and other soil-borne fungi

The mycoparasitic interactions of Verticillium biguttatum with Rhizoctonia solani and with a variety of other soil-borne fungi were investigated in dual cultures. V. biguttatum interacted with various soil fungi by appressed growth along the host hyphae and infrequent penetrations. Intracellular growth and subsequent sporulation, however, only occurred with R. solani, a few binucleate Rhizoctonia and Ceratobasidium spp., and Sclerotinia sclerotiorum. Effective mycoparasitism on sclerotia was restricted to those belonging to R. solani.Electron-microscopic observations revealed that V. biguttatum can penetrate the host cell with infection tubes. This process is probably mediated by enzymatic hydrolysis of the cell wall. Subsequently, trophic hyphae develop within the host cytoplasm, ultimately resulting in death of the host cell.     

The susceptibility of immature stages of <Emphasis Type="Italic">Bemisia tabaci</Emphasis> to the entomopathogenic fungus <Emphasis Type="Italic">Lecanicillium muscarium</Emphasis> on tomato and verbena foliage

Lecanicillium muscarium is a widely occurring entomopathogenic fungus. Laboratory studies were conducted to determine the efficacy of L. muscarium against different instars of Bemisia tabaci on tomato and verbena foliage after two incubation times (3 and 7 days). Significant reduction in B. tabaci numbers were recorded on fungus treated plants (p < 0.001). Second instar B. tabaci proved most susceptible to L. muscarium infection. There was no significant difference in mortality of B. tabaci second instars after either 3 or 7 days exposure to L. muscarium on either host plant. The importance of the speed of pest mortality following treatment and the potential of L. muscarium to be incorporated into an integrated pest management strategy for the biocontrol of B. tabaci on tomato and verbena plants are discussed.     

Seedling vaccination by stem injecting a conidial suspension of F2, a non-pathogenic Fusarium oxysporum strain, suppresses Verticillium wilt of eggplant

Several bacterial and fungal strains have been evaluated as biocontrol agents (BCAs) against Verticillium dahliae. In these studies, the BCAs were applied as a root drenching inoculum; however, this application method may have an adverse effect on the native, beneficial for the plants, microbial community. In the present study, it was evaluated whether endophytic application by stem injecting a conidial suspension of the non pathogenic Fusarium oxysporum F2 strain, isolated from a V. dahliae suppressive compost amendment, could reduce significantly Verticillium wilt symptom development in eggplants. It was revealed that stem injection of F2 seven days before transplanting the seedlings to soil infested by V. dahliae microsclerotia resulted in reduced disease severity compared to the control treatment. To visualise F2 ramification into the plant vascular system eggplant stems were injected with an EGFP transformed F2 strain. It was shown that F2 colonises effectively the plant vascular tissues over a long period of time as it was assessed by F2 DNA levels. In parallel, qPCR analysis showed that the application of F2 reduced significantly the amount of V. dahliae DNA in the stem tissues compared to the control treatment.     

Effect of temperature on the duration of larval and pupal stages of two species of sciomyzid flies, predators of the snail Lymnaea truncatula

The duration of larval and pupal stages of two species of predatory flies was investigated at five constant temperatures. Ilione albiseta (Scopoli): Mean duration of first and second instars was shortest at 23°C and there was 100% survival of first instar larvae at all temperatures. Mean third instar larval duration decreased from 104 days at 17°C to 27.5 days at 26°C. Total percentage larval survival was greatest at 17°C and least at 26°C. Mean pupal duration decreased as temperature increased and this occurred also under outdoor conditions.Pherbellia cinerella (Fallén): As temperature increased mean larval duration decreased but the percentage of larvae pupating also decreased (100% at 14°C, 33.3% at 26°C). There was a trend for shorter mean pupal durations at higher temperatures.

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Résumé L'action de la température a été examinée à 14, 17, 20, 23 et 26°C. Les 2 espèces sont prédatrices de L. truncatula, vecteur de la douve du foie en Irlande, où elle provoque à l'àgriculture une perte évaluée à 10 millions de livres par an. Les données recuellies en conditions contrôlées du laboratoire doivent permettre de définir les conditions optimales pour l'élevage continu de ces espèces. Ceci est indispensable pour évaluer, dans la nature leurs potentialités pour la lutte biologique.Ilione albiseta: C'est à 23°C que le développement des larves de premier et second stades sont les plus brefs; la survie du premier stade est totale à toutes les températures constantes examinées. La durée moyenne du 3ème stade décroît de 104 jours à 17°C jusqu'à 27,5 jours à 26°C. Le taux de survie total est plus élevé à 17°C qu'à 26°C. La durée moyenne de la nymphose diminue quand la température croît, et ceci se produit aussi à la température ambiante extérieure. On réduit de moitié la durée du développement de l'éclosion de l'oeuf à l'apparition de l'adulte, entre la nature et le laboratoire, en utilisant respectivement: 23°, 17° et 26°C pour les premier et second stades, le troisième stade, la nymphé. Cela devrait aider à multiplier la souche pour la lutte biologique.Pherbellia cinerella: Avec élévation de la température la durée du développement larvaire diminue, mais aussi le nombre de larves parvenant à la pupaison (100% à 14° et 33,3% à 26°C). La nymphose tendait à être plus brève aux températures les plus élevées.

     

The relationships between temperature and rate of development in two geographic stocks of Aphidius sonchi in the laboratory

The developmental periods from egg to adult of two stocks of the aphid parasite, Aphidius sonchi Marshall (Hymenoptera: Aphidiidae), were determined under various constant and alternating temperature regimes. The relationship between temperature and rate of development for each of the two stocks under constant conditions was described satisfactorily by the logistic curve. Based on the constant temperature curves, comparisons showed that temperature alternations did not accelerate or decelerate rate of development in either stock. Further analysis indicated that rates of development at extreme temperatures were well described by the logistic curve.The data and subsequent analyses showed that (1) because of the curvilinear form of the relationship, durations of development can differ considerably between constant and varying conditions if comparisons are made using daily mean temperatures; (2) as a result, the development threshold and thermal constant estimated by a hyperbolic transformation of data referred to daily mean temperatures will vary with the amplitude and pattern of fluctuations; and (3) temperature relationships derived from laboratory data can be used safely to predict the development of insects in the field.

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Résumé La durée du développement de l'oeuf à l'adulte de deux lignées d'A. sonchi Marshall (Hymenoptera: Aphidiidae) a été déterminée en températures constantes et avec des thermopériodes. La courbe logistique a décrit correctement la relation entre la température et la vitesse de développement pour chacune des deux lignées en conditions constantes. En se basant sur les courbes à température constante, les comparaisons ont montré que l'alternance thermique n'accélère, ni ne ralentit le développement des lignées. Une analyse complémentaire a montré que le développement aux températures extrêmes était correctement décrit par la courbe logistique. Les données et les analyses consécutives ont montré: (1) que par suite de l'aspect curviligne de la relation, les durées de développement en conditions constantes et thermopériodiques peuvent différer considérablement si les comparaisons portent sur les températures quotidiennes moyennes; (2) qu'en conséquence, le seuil de développement et la constante thermique estimés par une transformation hyperbolique des données concernant les températures quotidiennes moyennes varieront avec l'amplitude et le mode de fluctuation; et (3) que les relations thermiques obtenues à partir des données de laboratoire ne peuvent être utilisées en confiance pour prédire le développement des insectes dans la nature.

     

Effect of vesicular-arbuscular mycorrhizal fungi on verticillium wilt of cotton

The development of vesicular-arbuscular mycorrhizal fungi (VAMF): Glomus mosseae (Nicol and Gerd.) Gerdemann and Trappe, Glomus versiforme (Karsten) Berch, Sclerocystis sinuosa Gerdemann and Bakhi and Verticillium dahliae and the effects of the VAMF on the verticillium wilt of cotton (Gossypium hirsutum L. and Gossypium barbadense L.) were studied with paper pots, black plastic tubes and clay pots under natural growth conditions. All of the tested VAMF were able to infect all the cotton varieties used in the present experiment and typical vesicles and arbuscules were formed in the cortical cells of the cotton roots after inoculation. The cap cells, meristem, differentiating and elongating zones of the root tip were found to be colonized by the VAMF. In the case of most V. dahliae infection, the colonization occurred mostly from the root tip up to 2 cm. VAMF and V. dahliae mutually reduced their percentage of infection when inoculated simultaneously. VAMF inoculation reduced the numbers of germinable microsclerotia in the soil of the mycorrhizosphere, while the quantity of VAM fungal spores in the soil was not influenced by infection of with V. dahliae. The % of arbuscule colonization in roots was negatively correlated with the disease grades, while the numbers of vesicles in roots were not. These results suggest that certain vital competition and antagonistic reactions exist between VAMF and V. dahliae. VAMF reduced the incidence and disease indices of verticillium wilt of cotton during the whole growth phase. It is evident that cotton seedling growth was promoted, flowering was advanced, the numbers of flowers and bolls were increased, and this resulted in an increase in the yield of seed cotton. Among the VAMF species, Glomus versiforme was the most effective, and Sclerocystis sinuosa was inferior. So far as the author is aware, such an effect of VAMF on the increase of cotton wilt tolerance/resistance is reported here far the first time.     

Genetic interactions between FLM and other flowering-time genes in Arabidopsis thaliana

FLOWERING LOCUS M (FLM) is a MADS-domain gene that acts as an inhibitor of flowering in Arabidopsis. Here we describe the genetic interaction of FLM with genes in the photoperiod and autonomous flowering pathways. Although the sequence of FLM is most similar to that of FLC, FLM and FLC interact with different flowering pathways. It has been previously shown that flc lesions suppress the late-flowering phenotype of FRI-containing lines and autonomous-pathway mutants. However, flm lesions suppress the late-flowering phenotype of photoperiod-pathway mutants but not that of FRI-containing lines or autonomous-pathway mutants. Another MADS-domain flowering repressor with a mutant phenotype similar to FLM is SVP. The late-flowering phenotype of FLM over-expression is suppressed by the svp mutation, and an svp flm double mutant behaves like the single mutants. Thus FLM and SVP are in the same flowering pathway which interacts with the photoperiod pathway. Abbreviations: CO, CONSTANS; FLC, FLOWERING LOCUS C; FLM, FLOWERING LOCUS M; FRI, FRIGIDA; GI, GIGANTEA; LD, LUMINIDEPENDENS; SVP, SHORT VEGETATIVE PHASE; FCA is not an abbreviation     

Effect of temperature on infection, development and reproduction of the parasitic nematode Thripinema nicklewoodi in Frankliniella occidentalis

The parasitic nematodeThripinema nicklewoodi Siddiqi (Tylenchida:Allantonematidae) is currently underinvestigation for use in inoculative releasestrategies against western flower thrips (WFT),Frankliniella occidentalis Pergande(Thysanoptera: Thripidae) infesting greenhousefloricultural crops. The aim was to determinewhether temperatures within greenhouses wouldpermit the establishment of T.nicklewoodi. The abilities of T.nicklewoodi to infect, develop and reproducein WFT were assessed under a range of constantand fluctuating temperatures in the laboratory.At constant temperatures, T. nicklewoodiinfected WFT over the range of 1–30 °C,although the temperature-related infectionprofile followed an asymmetric distributionaround an optimum 20 °C (80%infection). The lower and upper thresholds forT. nicklewoodi in vivo development andreproduction were higher than for infection, at10 °C and 35 °C, respectively.Climate data recorded over 1999–2000 in acommercial greenhouse (Texas) revealed atemperature range of 15 °C to31 °C from early March through mid June,when WFT were most abundant. While low(nighttime) greenhouse temperatures areconducive for T. nicklewoodi, upperdaytime temperatures are close to the upperthreshold for infection and may reducereproductive output. However, fluctuatingtemperature bioassays in the laboratorydemonstrated that T. nicklewoodimaintained separately at the upper thresholdtemperatures for infection (30 °C) anddevelopment (35 °C) readily infected anddeveloped in WFT when they were allowedintermittent (10 h daily) exposure to apermissive temperature in the range10–20 °C. Drawing on the results, thediurnal temperature-fluctuations of variousgreenhouses growing ornamentals would permitthe establishment of T. nicklewoodi.     

Biological control of postharvest fungal rot of yam (<Emphasis Type="Italic">Dioscorea</Emphasis> spp.) with<Emphasis Type="Italic">Bacillus subtilis</Emphasis>

The potential of isolates of Bacillus subtilis from yam farm soil to control rot of yam in storage barns was investigated. Yam tubers inoculated in vivo with B. subtilis showed no rot while those inoculated with Aspergillus niger, Botryodiploidia theobromae or Penicillium oxalicum showed considerable rot. The set of yams in which B. subtilis and the fungi were simultaneously inoculated produced rot whereas those in which B. subtilis was inoculated a day before the fungi was inoculated were totally reduced or free of rot. Many fewer fungi were isolated from the surface of tubers treated with B. subtilis than from the untreated (control) and there was high recovery of B. subtilis (99–100%) throughout the period of storage. Rot build up was faster in uninoculated control tubers or those inoculated with a spoilage fungus, while those treated with the antagonist were totally reduced or free of rot. The culture filtrate of B. subtilis prevented spore germination in some spoilage fungi. The importance of this study in relation to farmers in developing countries is discussed.     

Verticillium dahliae toxin induced alterations of cytoskeletons and nucleoli in Arabidopsis thaliana suspension cells

Summary. In plant cells, cytoskeletons play important roles in response to biotic and abiotic stresses. However, little is known about the dynamics of cytoskeletons when cells are attacked by unphysical stress factors such as elicitors and toxins. We report here that the toxin of Verticillium dahliae (VD toxin) induced changes of microfilaments (MFs) and microtubules (MTs) in Arabidopsis thaliana suspension-cultured cells. When cells were treated with a low concentration of VD toxin, MFs were disrupted ordinally from the cortex to the perinuclear region, and then recovered spontaneously; but the MTs persisted. The MFs in the perinuclear region showed more resistance to VD toxin than the cortical ones. In contrast, when cells were treated with a high concentration of VD toxin, MFs and MTs were disrupted sooner and more severely and did not recover spontaneously. Treatments with high concentrations of VD toxin also induced changes of nucleoli. At the early stages of treatment, a nucleus had a single ring-shaped nucleolus. At the later stages, multiple smaller and more brightly fluorescing nucleoli emerged in a single nucleus. Disrupted MFs could be recovered by removing the VD toxin before the ringshaped nucleoli appeared. All these results showed that MFs and MTs play important roles in the early defense responses against VD toxin in Arabidopsis suspension cells. The cytoskeletons may be used as sensors and effectors monitoring the defense reactions. The changes of nucleoli induced by VD toxin should be important characteristics of cell death. Correspondence and reprints: Department of Plant Sciences, College of Biological Sciences, China Agricultural University, Beijing 100094, People’s Republic of China.     

Interacting effects of time,temperature, pH and simple sugars on biomass and toxic metabolite production by three Alternaria spp.

Biomass and toxic metabolite production by Altemaria hemicota, A. solani and A. tenuis in modified Czapek Dox broth (mCDB) as affected by incubation time (up to 4 wk) temperature (6, 15 and 25 °C), pH (3.0, 5.0 and 8.0) and sugars (glucose, fructose and sucrose) was studied. Biomass production was greatest at 25 ° C and pH 8.0 but toxin production was enhanced at 15 °C. In general, biomass production was promoted by up to 9% glucose and sucrose and 6% fructose; however, toxin production was simultaneously reduced by elevated sugars in mCDB over a 4-wk incubation period.     

Role of plant phenology in mediating interactions between two biological control agents for spotted knapweed

The role of spotted knapweed phenology on the attack rate of two seed-head insects Urophora affinis and Larinus minutus was assessed in a series of field studies at four study sites in south-eastern British Columbia, Canada. Slow or later developing knapweed plants had more seed heads that contained only single or multiple U. affinis whereas early or faster developing plants had more seed heads containing L. minutus alone or in combination with U. affinis. L. minutus did not distinguish between seed heads with or without U. affinis larvae when laying eggs. However, seed heads with multiple U. affinis present, produced fewer L. minutus adults than expected. The probability of single or multiple U. affinis galls being present increased with seed-head diameter but was not affected by seed-head height. Attack by L. minutus increased with seed-head diameters >5 mm and was lower at plant heights above 50 cm. These results demonstrate two mechanisms that enable U. affinis to successfully coexist with L. minutus: differences between the species in their response to the developmental phenology of knapweed heads, and increased survivorship of U. affinis in heads with multiple U. affinis galls through niche interference competition. These mechanisms provide a possible explanation for the persistence of U. affinis populations on spotted knapweed, in spite of high levels of within seed-head mortality that have been observed with increasing L. minutus populations.