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Insect-Microorganism-Plant Interactions 2nd Discussion session 5th Lecture 1 Insect-Microorganism-Plant Interactions Fact Insects and microorganisms require hosts. 2 Introduction Mutualism Parasitism Insects + Micro Plants Directly or indirectly interactions 3 Introduction Mutualism Parasitism Plant + Micro Insects Directly or indirectly 4 Plant-Microbes-Insects When an insect touches a plant, it touches Microbes (mainly bacteria) and their metabolic products. 5 Introduction What are the benefits for microbes being on plants. House that may provide shelter, suitable microclimate …etc for microbes 6 Outlines 1- Endophytic Fungi 2- Insect orientation by Bacteria 3- Entomopathogens-insect-plant interactions 4- Symbiotic bacteria in insects 8 1 Endophytic Fungi 11 Endophytic Fungi The endophyte is actually a fungus that lives, grows and thrives inside a host plant Most terrestrial plants are colonized by one or several species of endophytic fungi which can be isolated from healthy-looking host tissue Endophytes are contained within the plant without disease. Plant tissues remain entire and functional. 12 Endophytic Fungi The endophyte and the grass plant form a mutually beneficial relationship; the plant feeds and "houses" the endophyte while the endophyte helps the plant survive insects, heat and drought. 13 Endophytic Fungi Endophytes may upregulate host responses to pathogens and pests. Chaetomium globosum has been shown to increase host resistance to rust and tan spot pathogens in wheat. Direct interactions appear to be too small to measure in this case. Presence of Lecanicillium lecanii appears to reduce the feeding by aphids from leaves of cotton 14 Endophytic Fungi Plants may benefit from the presence of endophytes in many ways. Potential plant benefits have been examined in only a few cases. Rhabdocline parkeri produces a compound that reduces needle attack by borers. Metabolites produced by Phomopsis sp in cotton appear to deter larvae of Helicoverpa from feeding on leaves. The parallels with Neotyphodium are clear. In addition, aphids feeding on leaves of cotton may become colonized by Lecanicillium lecanii, when conditions permit. Thus the aphid may be killed or it may transfer the fungus to another leaf 16 2 Insect orientation by Bacteria 17 Insect orientation by Bacteria Bacteria as adult food Associated with oviposition site, host fruit surfaces, and larval infested tissues Bacteria orient fruit flies to the suitable host Fruit flies-Bacteria-plant interactions 18 3 Entomopathogens-insect-plant interactions 19 Entomopathogens-insect-plant interactions Plants can use entomopathogens as bodyguards 1) maintaining a population of bodyguards on the plant surface, (2) increasing contact rates between insect host and pathogen and (3) increasing the susceptibility of the host. 20 Entomopathogens-insect-plant interactions Plants produce many secondary compounds that confer protection against herbivore feeding (Karban and Baldwin 1997). In addition to their direct effects on herbivores, these compounds may interact with bacterial, fungal, and viral pathogens of herbivores. Secondary plant metabolites have shown both synergistic and antagonistic effects on toxicity of B. thuringiensissubsp. kurstaki against the gypsy moth. 21 Entomopathogens-insect-plant interactions For example, oak (Quercus spp.) tannins, and conifer terpenoids can decrease toxicity of B. thuringiensis subsp. kurstaki against gypsy moths (Barbosa 1988, Appel and Schultz 1994, Farrar et al. 1996). In contrast, activity against larvae feeding on aspen (Populus spp.) increased with increasing foliar levels of phenolic glycosides (Hwang et al. 1995). Secondary metabolites have been hypothesized to be responsible for the relatively low activity of B. thuringiensis subsp. kurstaki against gypsy moth larvae on willow (Salix spp.) (Farrar et al. 1996). 22 24 Entomopathogens-insect-plant interactions the fungal pathogen Beauveria bassiana sprayed on to US corn was found to grow into the plant and provide season-long control of corn borer larvae ( Bing & Lewis 1991). 25 Entomopathogens-insect-plant interactions Many plant traits will influence directly or indirectly the survival of entomopathogens. Although certainly not the only selection pressure on these traits, selection may mould them to improve pathogen persistence. Examples are canopy architecture, leaf form and leaf colour, which can greatly diminish the harmful effects of UV on these pathogen propagules. Other examples include the density of hairs, waxiness, the veins, size and shape of leaf, angle of the leaf to the stem, number of stomata and density of the canopy, which will influence the microclimate of the leaf, the thickness of the boundary layer and the effects of wind and rain 26 Entomopathogens-insect-plant interactions Aschersonia aleyrodis were found to retain infectivity to whitefly much better on some plants (e.g. cucumber) than others (e.g. poinsettia). 27 Entomopathogens-insect-plant interactions laboratory study by Ignoffo et al. (1977 ), who demonstrated that soybean seedlings can pick up spores of the fungus Nomuraea rileyi from the soil and that these spores can subsequently infect larvae of Trichoplusia ni (soybean looper). 28 Entomopathogens-insect-plant interactions Brown et al. (1995 ). The authors found that the germination of conidia of the entomophthoralean pathogen Pandora ( = Erynia) neoaphidis was inhibited by volatiles released by tobacco plants in response to herbivory by Myzus nicotianae (tobacco aphids). Germination would in this instance lead to the production of secondary conidia, which would redisperse away from the leaf. Thus, the authors hypothesized that the conidia "sit and wait" until they are picked up, whereupon they germinate to infect the aphid hosts. 29 Bacteria was sampled from fruit-free plants, plants without fruits (just were harvested) and plant with fruits. Fruit fly Bactrocera proved an olfactory to attract flies to the host plants 30 Plant-mediated interactions between pathogenic microorganisms and herbivorous insects 31 Plant-mediated interactions between pathogenic microorganisms and herbivorous insects Types:1. Direct plant-mediated effect 2. Locally plant-mediated effect 3. Systemic effect 32 Direct plant-mediated effect Effect on antagonist 2 results from the physical presence or activity of antagonist 1 and/or from plant responses induced by antagonist 1 Some of these interactions are direct, for example, when arthropods vector phytopathogenic microorganisms 33 Locally plant-mediated effect Effect on antagonist 2 results exclusively from plant response induced by antagonist 1. Both antagonists are were present on the same morphological entity of the plant (typically leaf) The term local refers to interactions in which pathogens and arthropods used the same plant tissue at the same or different times 34 Systemic effect Effect on antagonist 2 results exclusively from plant response induced by antagonist 1. Each antagonist is or was active on a different morphological entity of the plant. Indirect interactions between phytopathogenic microorganisms and herbivorous arthropods can occur when infection or infestation by a first attacker alters the shared host plant in a way that affects a second attacker that is often spatially or temporally separated from the first 35 Plant-mediated interactions between pathogenic microorganisms and herbivorous insects Objectives plant-mediated interactions will facilitate an understanding of how plants coordinate and integrate their defenses against multiple biotic threats. 36 The contemporary view of plant defense recognizes that plants contend simultaneously with myriad attackers by utilizing a multifaceted array of resistance mechanisms, involving not only primary and secondary metabolites and morphological and physicochemical traits, but also mechanisms that allow plants to tolerate attack and recruit the natural enemies of attackers Recruit = to raise or strengthen 37 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES Interactions in which virus infection has a beneficial effect on vectoring homopterans are thought to reflect a mutualistic relationship between virus and vector. Data consistent with this hypothesis were presented, for example, in a study in which rates of increase of Myzus persicae on virusinfected potatoes were positively correlated with the level of dependence of the virus on the aphid for dispersal 38 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES Similarly, recent studies demonstrated that aphid vectors such as Rhopalosiphum padi and Sitobion avenae had greater preference for, and higher fecundities and rates of increase on, cereals infected with Barley yellow dwarf virus than for healthy cereals under laboratory conditions 39 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES Even in interactions in which virus infection has no net effect or a net negative effect on vector population growth or fitness, changes in vector settling or feeding behavior may be adaptations to increase the spread of the virus For example, the leafhopper Nepotettix virescens, a vector of Tungro virus in rice, grew slower, fed less, was less fecund, and experienced higher mortality on infected rice plants than on uninfected plants. Despite this overall negative effect of infection on leafhoppers, initial settling on Tungro-infected rice plants was higher. The alterations in initial settling behavior and feeding behavior were hypothesized to increase spread of the virus. 40 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES Two studies have shown positive effects of systemic virus infection on nonhomopteran arthropods. Survival of Colorado potato beetle larvae was higher on tomato plants infected with Tobacco mosaic virus (TMV) than on healthy plants Larvae of the Mexican bean beetle fed less and grew slower on uninfected bean plants than on bean plants infected with Southern bean mosaic virus or Bean pod mottle virus (88). In the latter case, the insect vectors the viruses. 41 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES The effects on arthropods of feeding on tissues infected with pathogenic fungi or bacteria can be positive or negative. European corn borer larvae developed approximately 20% faster on maize showing symptoms of stalk rot caused by Colletotrichum graminicola than on nondiseased tissue 42 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES In contrast, systemic infection of Cirsium arvense with the necrotrophic fungus Phoma destructiva resulted in a number of negative effects on the beetle Cassida rubiginosa, including reduced oviposition, feeding, survival, rates of growth, and pupal weights 43 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES One factor that appears to influence the outcome of these types of tripartite interactions is the constitutive quality of the infected plant for the herbivore. The negative effects of rust infection on larvae of Tyria jacobaeae were greater when larvae fed on infected leaves of Tussilago farfara, a poor host for this insect species, than when larvae fed on rust-infected Senecio jacobaea, a superior host 44 FEEDING BY ARTHROPODS ON DISEASED PLANT TISSUES Similarly, the increase in relative growth rates and fecundities of black bean aphids (Aphis fabae) on bean leaves infected with the necrotroph Botrytis fabae was greater on a resistant bean variety than on a susceptible one 45 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS Several other studies have investigated the systemic effects of localized infection by bacterial and fungal pathogens on subsequent attackers. Again, both positive and negative effects have been demonstrated 46 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS Localized infection of one leaflet on young tomato plants with P. syringae pv. tomato resulted in 50% to 80% reductions in growth rates of the noctuid Helicoverpa zea on the remaining leaflets of the inoculated leaf. In contrast, infection of terminal leaflets by the hemibiotrophic oomycete Phytophthora infestans had no effect on leaf-systemic resistance to H. zea, demonstrating again that infection by different pathogens can have different results for the same herbivore 47 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS Infection of stems of peanuts with the necrotrophic white mold fungus, Sclerotium rolfsii, altered the resistance of leaves to a leaf-feeding caterpillar, Spodoptera exigua. Survival was∼20% higher, development was faster, pupae were ∼25% heavier, and feeding was greater on diseased plants 48 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS Only a few studies have directly compared the systemic effects of pathogen infection on arthropods with local effects. Local effects were usually stronger than systemic effects Local effects were usually stronger than systemic effects. Weights of larvae and pupae were reduced and development times were extended for the beetle Phaedon cochleariae feeding on cabbage leaves infected with the necrotroph Alternaria brassicae, but the effects were not systemic 49 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS Local effects were usually stronger than systemic effects.Weights of larvae and pupae were reduced and development times were extended for the beetle Phaedon cochleariae feeding on cabbage leaves infected with the necrotroph Alternaria brassicae, but the effects were not systemic 50 PLANT-MEDIATED EFFECTS OF PATHOGEN INFECTION ON INSECTS In addition, melon aphid (Aphis gossypii) produced fewer offspring on diseased leaves when leaves were almost completely necrotic, but produced more offspring on diseased leaves when symptoms were not as severe. No systemic effects on aphid reproduction were observed. In addition to showing the importance of spatial scale, this latter study also shows the importance of disease severity in determining effects on herbivores. 51 PLANT-MEDIATED EFFECTS OF ARTHROPODS ON PATHOGENS Several recent studies have documented increases in resistance to pathogens following feeding by arthropods with a sucking mode of feeding. Because sucking insects can induce biochemical responses similar to those induced by pathogens 52 PLANT-MEDIATED EFFECTS OF ARTHROPODS ON PATHOGENS Infesting rice plants with the white-backed planthopper, Sogatella furcifera, dramatically increased the resistance of plants to rice blast, Magnaporthe grisea 53 In tomato, plants previously infested with silverleaf whitefly, Bemisia argentifolii, were more resistant to powdery mildew (Erysiphe cichoracearum), but not to TMV, than were control plants 54 Feeding by chewing insects can also influence plant resistance to pathogens. Feeding by H. zea on the terminal leaflets of tomato leaves resulted in a 30% reduction in lesion numbers caused by P. syringae on undamaged leaflets of damaged leaves 55 Several investigations of tripartite interactions among pests of alfalfa have also been conducted and again show the importance of feeding type. Defoliation of alfalfa by yellowstriped armyworms did not affect severity of crown rot caused by F. oxysporum 56 Hatcher et al. (51) documented a plant-systemic increase in the resistance of Rumex sp. to the rust fungus Uromyces rumicis following feeding by G. viridula 57 MECHANISMS OF PLANTMEDIATED INTERACTIONS Activation of Plant Response Pathways DEFENSE SIGNALING IN PLANTS SECONDARY METABOLITES CROSS-EFFECTS CROSS-TALK Other Mechanisms of Plant-Mediated Interactions CHANGES IN PRIMARY METABOLISM PLANT STRESS-MEDIATED INTERACTIONS CHANGES IN PLANT GROWTH AND MORPHOLOGY INTERACTIONS MEDIATED BY NATURAL ENEMIES 58 4 Symbiotic bacteria in insects 59 Symbiotic bacteria in insects symbiotic micro-organisms that provide the insect with nutrients or detoxify plant allelochemicals They may expand the plant range of insects by improving insect utilization of otherwise marginal plants 60 Symbiotic bacteria in insects What bacteria do for the Insects. A- Degradation of plant food B- Synthesis of nutritional requisites that plants do not provide at all or provide in insufficient quantities (sterols, amino acids…. C- Detoxification of plant allelochemicals 61 Symbiotic bacteria in insects A comprehensive understanding of the biology of insects requires that they be studied in ecological context with microorganisms as an important component of the system E.A. Steinhaus (1960) 62 Symbiotic bacteria in insects 63 64