Testing a Novel Attract-and-Kill Strategy for Drosophila suzukii (Diptera: Drosophilidae) Management.

The invasion of the spotted wing drosophila, Drosophila suzukii Matsumura, across the Americas and Europe has led to increased insecticide applications to protect fruit crops. This insecticide usage conflicts with integrated pest management programs, as well as harvest, export, and pollination services in the affected crops. A novel management tool was assessed against D. suzukii that may mitigate these conflicts. HOOK SWD, an attract-and-kill (A&K) formulation applied as a sprayable bait, was evaluated for three growing seasons in two berry crops in New Jersey and California. In blueberry crops treated with HOOK SWD, fruit infestations by D. suzukii were 2-8 times lower than in untreated crops. In trials in commercial raspberry fields, weekly or biweekly HOOK SWD applications combined with a single grower standard D. suzukii-targeted cover spray resulted in nearly 2-5 times fewer fruit infestations compared to the grower standard cover spray alone. Assays of the residual activity of HOOK SWD resulted in more than 78-93% adult D. suzukii mortality when exposed to raspberry leaves after the formulation had aged for 35 d in the field under plastic hoop houses. These results suggest that this A&K strategy can be integrated in D. suzukii management programs.

Temperature-Mediated Development Thresholds of Sparganothis sulfureana (Lepidoptera: Tortricidae) in Cranberries.

Larvae of Sparganothis sulfureana Clemens frequently attack cranberries, often resulting in economic damage to the crop. Because temperature dictates insect growth rate, development can be accurately estimated based on daily temperature measurements. To better predict S. sulfureana development across the growing season, we investigated the temperature range within which S. sulfureana larvae can feed and grow. Larvae were reared at 13 constant temperatures ranging from 6.5-38.6 °C. Larval growth rate was determined by the rate of change of larval weight across time. The respective growth rates among these temperatures were modeled using simple linear, cubic, and Lactin nonlinear development functions. These models isolated the lower temperature threshold at which growth became nonzero and the upper temperature at which growth was maximized. All three models were significantly predictive of S. sulfureana growth, but the cubic model best represented the observed growth rates, effectively isolating lower and upper thresholds of 9.97 and 29.89 °C, respectively. We propose that these thresholds be used to create a degree-day model of temperature-mediated S. sulfureana development.

Landscape factors facilitating the invasive dynamics and distribution of the brown marmorated stink bug, Halyomorpha halys (Hemiptera: Pentatomidae), after arrival in the United States.

The brown marmorated stink bug, Halyomorpha halys, a native of Asia, has become a serious invasive pest in the USA. H. halys was first detected in the USA in the mid 1990s, dispersing to over 41 other states. Since 1998, H. halys has spread throughout New Jersey, becoming an important pest of agriculture, and a major nuisance in urban developments. In this study, we used spatial analysis, geostatistics, and Bayesian linear regression to investigate the invasion dynamics and colonization processes of this pest in New Jersey. We present the results of monitoring H. halys from 51 to 71 black light traps that were placed on farms throughout New Jersey from 2004 to 2011 and examined relationships between total yearly densities of H. halys and square hectares of 48 landscape/land use variables derived from urban, wetland, forest, and agriculture metadata, as well as distances to nearest highways. From these analyses we propose the following hypotheses: (1) H. halys density is strongly associated with urban developments and railroads during its initial establishment and dispersal from 2004 to 2006; (2) H. halys overwintering in multiple habitats and feeding on a variety of plants may have reduced the Allee effect, thus facilitating movement into the southernmost regions of the state by railroads from 2005 to 2008; (3) density of H. halys contracted in 2009 possibly from invading wetlands or sampling artifact; (4) subsequent invasion of H. halys from the northwest to the south in 2010 may conform to a stratified-dispersal model marked by rapid long-distance movement, from railroads and wetland rights-of-way; and (5) high densities of H. halys may be associated with agriculture in southern New Jersey in 2011. These landscape features associated with the invasion of H. halys in New Jersey may predict its potential rate of invasion across the USA and worldwide.

Degree-Day Benchmarks for Sparganothis sulfureana (Lepidoptera: Tortricidae) Development in Cranberries.

Sparganothis sulfureana Clemens is a severe pest of cranberries in the Midwest and northeast United States. Timing for insecticide applications has relied primarily on calendar dates and pheromone trap-catch; however, abiotic conditions can vary greatly, rendering such methods unreliable as indicators of optimal treatment timing. Phenology models based on degree-day (DD) accrual represent a proven, superior approach to assessing the development of insect populations, particularly for larvae. Previous studies of S. sulfureana development showed that the lower and upper temperature thresholds for larval development were 10.0 and 29.9°C (49.9 and 85.8°F), respectively. We used these thresholds to generate DD accumulations specific to S. sulfureana, and then linked these DD accumulations to discrete biological events observed during S. sulfureana development in Wisconsin and New Jersey cranberries. Here, we provide the DDs associated with flight initiation, peak flight, flight termination, adult life span, preovipositional period, ovipositional period, and egg hatch. These DD accumulations represent key developmental benchmarks, allowing for the creation of a phenology model that facilitates wiser management of S. sulfureana in the cranberry system.

Trap designs for monitoring Drosophila suzukii (Diptera: Drosophilidae).

Drosophila suzukii (Matsumura), an invasive pest of small and stone fruits, has been recently detected in 39 states of the United States, Canada, Mexico, and Europe. This pest attacks ripening fruit, causing economic losses including increased management costs and crop rejection. Ongoing research aims to improve the efficacy of monitoring traps. Studies were conducted to evaluate how physical trap features affect captures of D. suzukii. We evaluated five colors, two bait surface areas, and a top and side position for the fly entry point. Studies were conducted at 16 sites spanning seven states and provinces of North America and nine crop types. Apple cider vinegar was the standard bait in all trap types. In the overall analysis, yellow-colored traps caught significantly more flies than clear, white, and black traps; and red traps caught more than clear traps. Results by color may be influenced by crop type. Overall, the trap with a greater bait surface area caught slightly more D. suzukii than the trap with smaller area (90 vs. 40 cm(2)). Overall, the two traps with a side-mesh entry, with or without a protective rain tent, caught more D. suzukii than the trap with a top-mesh entry and tent.

Jasmonate-Mediated Induced Volatiles in the American Cranberry, Vaccinium macrocarpon: From Gene Expression to Organismal Interactions.

Jasmonates, i.e., jasmonic acid (JA) and methyl jasmonate (MeJA), are signaling hormones that regulate a large number of defense responses in plants which in turn affect the plants' interactions with herbivores and their natural enemies. Here, we investigated the effect of jasmonates on the emission of volatiles in the American cranberry, Vaccinium macrocarpon, at different levels of biological organization from gene expression to organismal interactions. At the molecular level, four genes (BCS, LLS, NER1, and TPS21) responded significantly to gypsy moth larval feeding, MeJA, and mechanical wounding, but to different degrees. The most dramatic changes in expression of BCS and TPS21 (genes in the sesquiterpenoid pathway) were when treated with MeJA. Gypsy moth-damaged and MeJA-treated plants also had significantly elevated expression of LLS and NER1 (genes in the monoterpene and homoterpene biosynthesis pathways, respectively). At the biochemical level, MeJA induced a complex blend of monoterpene and sesquiterpene compounds that differed from gypsy moth and mechanical damage, and followed a diurnal pattern of emission. At the organismal level, numbers of Sparganothis sulfureana moths were lower while numbers of parasitic wasps were higher on sticky traps near MeJA-treated cranberry plants than those near untreated plants. Out of 11 leaf volatiles tested, (Z)-3-hexenyl acetate, linalool, and linalool oxide elicited strong antennal (EAG) responses from S. sulfureana, whereas sesquiterpenes elicited weak EAG responses. In addition, mortality of S. sulfureana larvae increased by about 43% in JA treated cranberry plants as compared with untreated plants, indicating a relationship among adult preference, antennal sensitivity to plant odors, and offspring performance. This study highlights the role of the jasmonate-dependent defensive pathway in the emissions of herbivore-induced volatiles in cranberries and its importance in multi-trophic level interactions.

Lethality of reduced-risk insecticides against plum curculio (Coleoptera: Curculionidae) in blueberries, with emphasis on their curative activity.

BACKGROUND: Historically, management of plum curculio, Conotrachelus nenuphar (Herbst), in highbush blueberries has focused on post-bloom broad-spectrum insecticide applications targeting the adults. Here, the efficacy of different classes of insecticides against various stages of C. nenuphar was compared, and a prebloom treatment with the chitin synthesis inhibitor novaluron in combination with a post-bloom insecticide application was tested. RESULTS: Novaluron decreased the number of oviposition scars and eggs on fruit and reduced larval emergence by >60% when applied prebloom. Post-bloom applications of the oxadiazine indoxacarb and the organophosphate phosmet, but not the neonicotinoid acetamiprid, showed significant adulticidal activity. The neonicotinoids acetamiprid and thiamethoxam and phosmet showed significant curative activity on C. nenuphar larvae when applied topically to infested fruit, whereas the pyrethroid fenpropathrin, indoxacarb and novaluron were weaker curative agents. Residue profiles showed that acetamiprid and phosmet residues had the highest levels while fenpropathrin and novaluron had the lowest levels of fruit penetration. CONCLUSIONS: In blueberries, novaluron showed anti-ovipositional/ovicidal activity, indoxacarb and phosmet showed adulticidal activity, while the neonicotinoids and phosmet showed best curative (larvicidal) control on C. nenuphar. A prebloom novaluron application in combination with a post-bloom treatment with an adulticidal/larvicidal insecticide is recommended for optimal multi-life-stage management of C. nenuphar. © 2013 Society of Chemical Industry.

Herbivore-induced blueberry volatiles and intra-plant signaling.

Herbivore-induced plant volatiles (HIPVs) are commonly emitted from plants after herbivore attack. These HIPVs are mainly regulated by the defensive plant hormone jasmonic acid (JA) and its volatile derivative methyl jasmonate (MeJA). Over the past 3 decades researchers have documented that HIPVs can repel or attract herbivores, attract the natural enemies of herbivores, and in some cases they can induce or prime plant defenses prior to herbivore attack. In a recent paper, I reported that feeding by gypsy moth caterpillars, exogenous MeJA application, and mechanical damage induce the emissions of volatiles from blueberry plants, albeit differently. In addition, blueberry branches respond to HIPVs emitted from neighboring branches of the same plant by increasing the levels of JA and resistance to herbivores (i.e., direct plant defenses), and by priming volatile emissions (i.e., indirect plant defenses). Similar findings have been reported recently for sagebrush, poplar, and lima beans. Here, I describe a push-pull method for collecting blueberry volatiles induced by herbivore (gypsy moth) feeding, exogenous MeJA application, and mechanical damage. The volatile collection unit consists of a 4 L volatile collection chamber, a 2-piece guillotine, an air delivery system that purifies incoming air, and a vacuum system connected to a trap filled with Super-Q adsorbent to collect volatiles. Volatiles collected in Super-Q traps are eluted with dichloromethane and then separated and quantified using Gas Chromatography (GC). This volatile collection method was used in my study to investigate the volatile response of undamaged branches to exposure to volatiles from herbivore-damaged branches within blueberry plants. These methods are described here. Briefly, undamaged blueberry branches are exposed to HIPVs from neighboring branches within the same plant. Using the same techniques described above, volatiles emitted from branches after exposure to HIPVs are collected and analyzed.

Molecular, biochemical, and organismal analyses of tomato plants simultaneously attacked by herbivores from two feeding guilds.

Previous work identified aphids and caterpillars as having distinct effects on plant responses to herbivory. We sought to decipher these interactions across different levels of biological organization, i.e., molecular, biochemical, and organismal, with tomato plants either damaged by one 3rd-instar beet armyworm caterpillar (Spodoptera exigua), damaged by 40 adult potato aphids (Macrosiphum euphorbiae), simultaneous damaged by both herbivores, or left undamaged (controls). After placing insects on plants, plants were transferred to a growth chamber for 5 d to induce a systemic response. Subsequently, individual leaflets from non-damaged parts of plants were excised and used for gene expression analysis (microarrays and quantitative real-time PCR), C/N analysis, total protein analysis, proteinase inhibitor (PI) analysis, and for performance assays. At the molecular level, caterpillars up-regulated 56 and down-regulated 29 genes systemically, while aphids up-regulated 93 and down-regulated 146 genes, compared to controls. Although aphids induced more genes than caterpillars, the magnitude of caterpillar-induced gene accumulation, particularly for those associated with plant defenses, was often greater. In dual-damaged plants, aphids suppressed 27% of the genes regulated by caterpillars, while caterpillars suppressed 66% of the genes regulated by aphids. At the biochemical level, caterpillars induced three-fold higher PI activity compared to controls, while aphids had no effects on PIs either alone or when paired with caterpillars. Aphid feeding alone reduced the foliar C/N ratio, but not when caterpillars also fed on the plants. Aphid and caterpillar feeding alone had no effect on the amount of protein in systemic leaves; however, both herbivores feeding on the plant reduced the amount of protein compared to aphid-damaged plants. At the organismal level, S. exigua neonate performance was negatively affected by prior caterpillar feeding, regardless of whether aphids were present or absent. This study highlights areas of concordance and disjunction between molecular, biochemical, and organismal measures of induced plant resistance when plants are attacked by multiple herbivores. In general, our data produced consistent results when considering each herbivore separately but not when considering them together.

New evidence for a multi-functional role of herbivore-induced plant volatiles in defense against herbivores.

A diverse, often species-specific, array of herbivore-induced plant volatiles (HIPVs) are commonly emitted from plants after herbivore attack. Although research in the last 3 decades indicates a multi-functional role of these HIPVs, the evolutionary rationale underpinning HIPV emissions remains an open question. Many studies have documented that HIPVs can attract natural enemies, and some studies indicate that neighboring plants may eavesdrop their undamaged neighbors and induce or prime their own defenses prior to herbivore attack. Both of these ecological roles for HIPVs are risky strategies for the emitting plant. In a recent paper, we reported that most branches within a blueberry bush share limited vascular connectivity, which restricts the systemic movement of internal signals. Blueberry branches circumvent this limitation by responding to HIPVs emitted from neighboring branches of the same plant: exposure to HIPVs increases levels of defensive signaling hormones, changes their defensive status, and makes undamaged branches more resistant to herbivores. Similar findings have been reported recently for sagebrush, poplar and lima beans, where intra-plant communication played a role in activating or priming defenses against herbivores. Thus, there is increasing evidence that intra-plant communication occurs in a wide range of taxonomically unrelated plant species. While the degree to which this phenomenon increases a plant's fitness remains to be determined in most cases, we here argue that within-plant signaling provides more adaptive benefit for HIPV emissions than does between-plant signaling or attraction of predators. That is, the emission of HIPVs might have evolved primarily to protect undamaged parts of the plant against potential enemies, and neighboring plants and predators of herbivores later co-opted such HIPV signals for their own benefit.

Seasonal abundance, life history, and parasitism of Caloptilia porphyretica (Lepidoptera: Gracillariidae), a leafminer of highbush blueberry.

The leafminer Caloptilia porphyretica Braun (Lepidoptera: Gracillariidae), has become a frequent pest in commercial highbush blueberries, Vaccinium corymbosum L., in New Jersey, but little is known about its seasonal abundance, life history, and parasitism rates. Monitoring programs were conducted from 2003 to 2006 on seven blueberry farms to determine the relative abundance of this leafminer by using pheromone-baited traps and by sampling vegetative and flower/fruit clusters and new shoot growth. We found at least three distinct generations per year, with populations reaching their highest peak in the second generation. Laboratory studies characterized the life history of C. porphyretica and its parasitoid Pholetesor sp. prob. salalicus (Mason). The developmental period of C. porphyretica, from egg to adult, took 927, 838, and 912 degree-days (DD) at 20, 25, and 30 degrees C, respectively, by using a developmental threshold of 4.8 degrees C. This was equivalent to an average of 892 DD to complete development to adult, which compared with 870 and 880 DD between the first and second, and second and third generations, respectively, by using pheromone trap data averaged from 2004 to 2006. Although C. porphyretica populations varied greatly, the number of larvae in cluster and new shoot samples was highly correlated with the number of adults in traps. Field parasitism rate was -29%, with the braconid Pholetesor sp. the most abundant parasitoid. Ten of the 13 parasitoid species collected belonged to the family Eulophidae. When different ages of leafminers were exposed to Pholetesor sp., we found that the parasitoid attacks 9-15-d-old instars that reside in the mines of leaves.

Optimization of pheromone deployment for effective mating disruption of oriental beetle (Coleoptera: Scarabaeidae) in commercial blueberries.

Cost is a potential limiting factor in the adoption of mating disruption to control oriental beetle, Anomala orientalis (Waterhouse) (Coleoptera: Scarabaeidae). A 3-yr study was conducted in 1-ha blueberry, Vaccinium corymbosum L., plots to test the possibility of lowering cost by reducing the number of point-source dispensers and pheromone [(Z)-7-tetradecen-2-one] concentrations, while maintaining mating disruption. Trap shutdown, as measured by the disruption index (DI), caged females, and sentinel potted-plants with tethered females were used to assess the success of mating disruption. Disrupted plots had DI values of > or = 93% in all years, and a lower percentage of mated females, compared with control plots. However, DI values were > or = 95% at > or = 50 dispensers per ha. When 25 dispensers containing 0.05 g of active ingredient (AI) were used per hectare, the numbers of males in female cages and larvae in sentinel pots were similar to controls. Thus, dispenser density was critical for successful mating disruption of oriental beetles. Male oriental beetles approach the dispensers at all times of the day according to field observations, indicative of competitive attraction as a potential mechanism for mating disruption. However, at peak activity, greater male attraction was observed to dispensers containing 0.1 g of pheromone than 0.05 or 0.025 g, demonstrating the importance of pheromone rate. Although dispensers continued to emit pheromone for at least 7 wk in the field, emission rates dropped to levels close to 0 after 3 wk. We conclude that deployment of > or = 50 dispensers/ha at > or = 0.1 g (AI) per dispenser is the most effective rate for mating disruption of oriental beetle in blueberries.

Herbivore-induced volatiles in the perennial shrub, Vaccinium corymbosum, and their role in inter-branch signaling.

Herbivore feeding activates plant defenses at the site of damage as well as systemically. Systemic defenses can be induced internally by signals transported via phloem or xylem, or externally transmitted by volatiles emitted from the damaged tissues. We investigated the role of herbivore-induced plant volatiles (HIPVs) in activating a defense response between branches in blueberry plants. Blueberries are perennial shrubs that grow by initiating adventitious shoots from a basal crown, which produce new lateral branches. This type of growth constrains vascular connections between shoots and branches within plants. While we found that leaves within a branch were highly connected, vascular connectivity was limited between branches within shoots and absent between branches from different shoots. Larval feeding by gypsy moth, exogenous methyl jasmonate, and mechanical damage differentially induced volatile emissions in blueberry plants, and there was a positive correlation between amount of insect damage and volatile emission rates. Herbivore damage did not affect systemic defense induction when we isolated systemic branches from external exposure to HIPVs. Thus, internal signals were not capable of triggering systemic defenses among branches. However, exposure of branches to HIPVs from an adjacent branch decreased larval consumption by 70% compared to those exposed to volatiles from undamaged branches. This reduction in leaf consumption did not result in decreased volatile emissions, indicating that leaves became more responsive to herbivory (or "primed") after being exposed to HIPVs. Chemical profiles of leaves damaged by gypsy moth caterpillars, exposed to HIPVs, or non-damaged controls revealed that HIPV-exposed leaves had greater chemical similarities to damaged leaves than to control leaves. Insect-damaged leaves and young HIPV-exposed leaves had higher amounts of endogenous cis-jasmonic acid compared to undamaged and non-exposed leaves, respectively. Our results show that exposure to HIPVs triggered systemic induction of direct defenses against gypsy moth and primed volatile emissions, which can be an indirect defense. Blueberry plants appear to rely on HIPVs as external signals for inter-branch communication.