Monocrotaline presence in the Crotalaria (Fabaceae) plant genus and its influence on arthropods in agroecosystems / Presença de monocrotalina em plantas do gênero Crotalaria (Fabaceae) e a sua influência sobre artrópodes nos agroecossistemas

Abstract Crotalaria (Fabaceae) occurs abundantly in tropical and subtropical regions and has about 600 known species. These plants are widely used in agriculture, mainly as cover plants and green manures, in addition to their use in the management of phytonematodes. A striking feature of these species is the production of pyrrolizidine alkaloids (PAs), secondary allelochemicals involved in plant defense against herbivores. In Crotalaria species, monocrotaline is the predominant PA, which has many biological activities reported, including cytotoxicity, tumorigenicity, hepatotoxicity and neurotoxicity, with a wide range of ecological interactions. Thus, studies have sought to elucidate the effects of this compound to promote an increase in flora and fauna (mainly insects and nematodes) associated with agroecosystems, favoring the natural biological control. This review summarizes information about the monocrotaline, showing such effects in these environments, both above and below ground, and their potential use in pest management programs.

Resumo Crotalaria (Linnaeus, 1753) (Fabaceae) ocorre abundantemente em regiões tropicais e subtropicais e tem cerca de 600 espécies conhecidas. Estas plantas são amplamente utilizadas na agricultura, principalmente como cobertura e adubos verdes, além da sua utilização no manejo de fitonematoides. Uma característica marcante destas espécies é a produção de alcalóides pirrolizidinicos (APs), aleloquímicos secundários envolvidos na defesa das plantas contra os herbívoros. Nas espécies de Crotalaria, a monocrotalina é a AP predominante, que tem muitas atividades biológicas relatadas, incluindo citotoxicidade, tumorigenicidade, hepatotoxicidade e neurotoxicidade, além de uma vasta gama de interações ecológicas. Assim, estudos têm procurado elucidar os efeitos desse composto para promover um incremento na flora e fauna (principalmente insetos e nematoides) associados aos agroecossistemas, favorecendo o controle biológico natural. Esta revisão compila informações sobre a monocrotalina, mostrando tais efeitos nesses ambientes, tanto acima como abaixo do solo e a sua potencial utilização em programas de manejo de pragas.

Monocrotaline presence in the Crotalaria (Fabaceae) plant genus and its influence on arthropods in agroecosystems

Abstract Crotalaria (Fabaceae) occurs abundantly in tropical and subtropical regions and has about 600 known species. These plants are widely used in agriculture, mainly as cover plants and green manures, in addition to their use in the management of phytonematodes. A striking feature of these species is the production of pyrrolizidine alkaloids (PAs), secondary allelochemicals involved in plant defense against herbivores. In Crotalaria species, monocrotaline is the predominant PA, which has many biological activities reported, including cytotoxicity, tumorigenicity, hepatotoxicity and neurotoxicity, with a wide range of ecological interactions. Thus, studies have sought to elucidate the effects of this compound to promote an increase in flora and fauna (mainly insects and nematodes) associated with agroecosystems, favoring the natural biological control. This review summarizes information about the monocrotaline, showing such effects in these environments, both above and below ground, and their potential use in pest management programs.

Resumo Crotalaria (Linnaeus, 1753) (Fabaceae) ocorre abundantemente em regiões tropicais e subtropicais e tem cerca de 600 espécies conhecidas. Estas plantas são amplamente utilizadas na agricultura, principalmente como cobertura e adubos verdes, além da sua utilização no manejo de fitonematoides. Uma característica marcante destas espécies é a produção de alcalóides pirrolizidinicos (APs), aleloquímicos secundários envolvidos na defesa das plantas contra os herbívoros. Nas espécies de Crotalaria, a monocrotalina é a AP predominante, que tem muitas atividades biológicas relatadas, incluindo citotoxicidade, tumorigenicidade, hepatotoxicidade e neurotoxicidade, além de uma vasta gama de interações ecológicas. Assim, estudos têm procurado elucidar os efeitos desse composto para promover um incremento na flora e fauna (principalmente insetos e nematoides) associados aos agroecossistemas, favorecendo o controle biológico natural. Esta revisão compila informações sobre a monocrotalina, mostrando tais efeitos nesses ambientes, tanto acima como abaixo do solo e a sua potencial utilização em programas de manejo de pragas.

Monocrotaline presence in the Crotalaria (Fabaceae) plant genus and its influence on arthropods in agroecosystems.

Crotalaria (Fabaceae) occurs abundantly in tropical and subtropical regions and has about 600 known species. These plants are widely used in agriculture, mainly as cover plants and green manures, in addition to their use in the management of phytonematodes. A striking feature of these species is the production of pyrrolizidine alkaloids (PAs), secondary allelochemicals involved in plant defense against herbivores. In Crotalaria species, monocrotaline is the predominant PA, which has many biological activities reported, including cytotoxicity, tumorigenicity, hepatotoxicity and neurotoxicity, with a wide range of ecological interactions. Thus, studies have sought to elucidate the effects of this compound to promote an increase in flora and fauna (mainly insects and nematodes) associated with agroecosystems, favoring the natural biological control. This review summarizes information about the monocrotaline, showing such effects in these environments, both above and below ground, and their potential use in pest management programs.

Infection by the semi-persistently transmitted Tomato chlorosis virus alters the biology and behaviour of Bemisia tabaci on two potato clones.

Insect-borne plant viruses usually alter the interactions between host plant and insect vector in ways conducive to their transmission ('host manipulation hypothesis'). Most studies have tested this hypothesis with persistently and non-persistently transmitted viruses, while few have examined semi-persistently transmitted viruses. The crinivirus Tomato chlorosis virus (ToCV) is semi-persistently transmitted virus by whiteflies, and has been recently reported infecting potato plants in Brazil, where Bemisia tabaci Middle East Asia Minor 1 (MEAM1) is a competent vector. We investigated how ToCV infection modifies the interaction between potato plants and B. tabaci in ways that increase the likelihood of ToCV transmission, in two clones, one susceptible ('Agata') and the other moderately resistant (Bach-4) to B. tabaci. Whiteflies alighted and laid more eggs on ToCV-infected plants than mock-inoculated plants of Bach-4. When non-viruliferous whiteflies were released on ToCV-infected plants near mock-inoculated plants, adults moved more intensely towards non-infected plants than in the reverse condition for both clones. Feeding on ToCV-infected plants reduced egg-incubation period in both clones, but the egg-adult cycle was similar for whiteflies fed on ToCV-infected and mock-inoculated plants. Our results demonstrated that ToCV infection in potato plants alters B. tabaci behaviour and development in distinct ways depending on the host clone, with potential implications for ToCV spread.

Direct and indirect resistance of sugarcane to Diatraea saccharalis induced by jasmonic acid.

Treating plants with synthetic jasmonic acid (JA) induces a defensive response similar to herbivore attack, and is a potential strategy for integrated pest management. Despite the importance of sugarcane, its JA-induced defences have not yet been studied. We investigated the effects of JA treatment on the direct and indirect resistance of sugarcane to the key-pest and specialist herbivore Diatraea saccharalis and the generalist Spodoptera frugiperda. Indirect defences were examined by testing the attraction of Cotesia flavipes, a sugarcane-borer parasitoid, to JA-induced volatile. The results showed that JA-treated sugarcane did not affect the weight gain of the two larvae. However, in dual-choice assays, both species preferred to feed on mock rather than JA-treated plants. Leaf colorimetric analyses showed that visual cues are unlikely to be involved in larval preference, whereas results from olfactometric assays revealed that D. saccharalis preferred JA-induced over mock plant volatiles. After 48 h of treatment, JA-treated plants emitted a volatile blend attractive to C. flavipes, comprised mainly of sesquiterpenes. However, the parasitoid did not discriminate JA-treated from host-damaged plant volatiles. When the wasps were given a choice between JA-treated and JA-treated + host-damaged plants, they preferred the latter, which emitted a more complex blend, suggesting that JA treatment likely does not hamper host-finding. We concluded that JA induces the emission of volatiles that are attractive to the sugarcane borer parasitoid, as well as an antixenosis type of resistance in sugarcane against the two pests, although neither volatiles nor visual cues alone are involved in the underlying mechanism.

Description of the immatures of Scaptocoris carvalhoi Becker (Hemiptera: Cydnidae).

Nymphs and adults of the burrower bug Scaptocoris carvalhoi Becker feed on vegetal sap of their host plants through the roots, and little is known on the morphology and biology of its immature stage. Therefore, we aimed to characterize the immatures of S. carvalhoi by describing the egg and the morphology of each instar. Eggs of S. carvalhoi have a smooth chorion surface without visible micropylar processes. The presence of five instars was confirmed by the coefficient of determination (R (2) > 0.95) and by the growth constant (K between 1.2 and 1.6). Nymphs have an elliptical body and fossorial scythe-like forelegs. The tarsi are absent as in adults, and the prototarsal insertion region becomes visible only in the fourth instar. Nymphs from first to fourth instar of S. carvalhoi showed the presence of 1 + 1 trichobothria in urosternites III to VII, close to the anterior margin and inside the spiracles; besides these trichobothria, fifth instars presented 1 + 1 pre-trichobothria in urosternites III to V located posteriorly, almost in the row of spiracles close to the posterior margin of the urosternites. This is the first detailed morphological record of immatures belonging to Scaptocoris.

Herbivore-induced plant volatiles to enhance biological control in agriculture.

Plants under herbivore attack synthetize defensive organic compounds that directly or indirectly affect herbivore performance and mediate other interactions with the community. The so-called herbivore-induced plant volatiles (HIPVs) consist of odors released by attacked plants that serve as important cues for parasitoids and predators to locate their host/prey. The understanding that has been gained on the ecological role and mechanisms of HIPV emission opens up paths for developing novel strategies integrated with biological control programs with the aim of enhancing the efficacy of natural enemies in suppressing pest populations in crops. Tactics using synthetic HIPVs or chemically/genetically manipulating plant defenses have been suggested in order to recruit natural enemies to plantations or help guiding them to their host more quickly, working as a "synergistic" agent of biological control. This review discusses strategies using HIPVs to enhance biological control that have been proposed in the literature and were categorized here as: (a) exogenous application of elicitors on plants, (b) use of plant varieties that emit attractive HIPVs to natural enemies, (c) release of synthetic HIPVs, and (d) genetic manipulation targeting genes that optimize HIPV emission. We discuss the feasibility, benefits, and downsides of each strategy by considering not only field studies but also comprehensive laboratory assays that present an applied approach for HIPVs or show the potential of employing them in the field.

Mating behavior of Diabrotica speciosa (Coleoptera: Chrysomelidae).

Diabrotica speciosa (Germar) is an economically important pest of Neotropical cultures and represents a quarantine risk for Neartic and Paleartic Regions. Despite its agricultural importance, few studies have been done on mating behavior and chemical communication, which has delayed the development of behavioral techniques for population management, such as the use of pheromone traps. In this study, we determined 1) the age at first mating; 2) diel rhythm of matings; 3) number of matings over 7 d; 4) the sequence of D. speciosa activities during premating, mating, and postmating; 5) the duration of each activity; and 6) response to male and female conspecific volatiles in Y-tube olfactometer. The first mating occurred between the third and seventh day after adult emergence and the majority of pairs mated on the fourth day after emergence. Pairs of D. speciosa showed a daily rhythm of mating with greater sexual activity between the end of the photophase and the first half of the scotophase. During the 7 d of observation, most pairs mated only once, although 30% mated two, three, or four times. In a Y-tube olfactometer, males were attracted by virgin females as well as by the volatile compounds emitted by females. Neither males nor their volatiles were attractive to either sex. Our observation provide information about mating behavior of D. speciosa, which will be useful in future research in chemical communication, such as identification of the pheromone and development of management techniques for this species using pheromone traps.

Fall Armyworm, spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), female moths respond to herbivore-induced corn volatiles.

In response to herbivore attack, plants release herbivore-induced plant volatiles (HIPVs) that represent important chemical cues for herbivore natural enemies. Additionally, HIPVs have been shown to mediate other ecological interactions with herbivores. Differently from natural enemies that are generally attracted to HIPVs, herbivores can be either attracted or repelled depending on several biological and ecological parameters. Our study aimed to assess the olfactory response of fall armyworm-mated female moths toward odors released by mechanically and herbivore-induced corn at different time intervals. Results showed that female moths strongly respond to corn volatiles, although fresh damaged corn odors (0-1 h) are not recognized by moths. Moreover, females preferred volatiles released by undamaged plant over herbivore-induced plants at 5-6 h. This preference for undamaged plants may reflect an adaptive strategy of moths to avoid competitors and natural enemies for their offspring. We discussed our results based on knowledge about corn volatile release pattern and raise possible explanations for fall armyworm moth behavior.

Herbivore-induced plant volatiles can serve as host location cues for a generalist and a specialist egg parasitoid.

Herbivore-induced plant volatiles are important host finding cues for larval parasitoids, and similarly, insect oviposition might elicit the release of plant volatiles functioning as host finding cues for egg parasitoids. We hypothesized that egg parasitoids also might utilize HIPVs of emerging larvae to locate plants with host eggs. We, therefore, assessed the olfactory response of two egg parasitoids, a generalist, Trichogramma pretiosum (Tricogrammatidae), and a specialist, Telenomus remus (Scelionidae) to HIPVs. We used a Y-tube olfactometer to tests the wasps' responses to volatiles released by young maize plants that were treated with regurgitant from caterpillars of the moth Spodoptera frugiperda (Noctuidae) or were directly attacked by the caterpillars. The results show that the generalist egg parasitoid Tr. pretiosum is innately attracted by volatiles from freshly-damaged plants 0-1 and 2-3 h after regurgitant treatment. During this interval, the volatile blend consisted of green leaf volatiles (GLVs) and a blend of aromatic compounds, mono- and homoterpenes, respectively. Behavioral assays with synthetic GLVs confirmed their attractiveness to Tr. pretiosum. The generalist learned the more complex volatile blends released 6-7 h after induction, which consisted mainly of sesquiterpenes. The specialist T. remus on the other hand was attracted only to volatiles emitted from fresh and old damage after associating these volatiles with oviposition. Taken together, these results strengthen the emerging pattern that egg and larval parasitoids behave in a similar way in that generalists can respond innately to HIPVs, while specialists seems to rely more on associative learning.