Long-term fertilization and cultivation impacts on nematode abundance and community structure in tall fescue turfgrass.

Impacts of long-term fertilization and cultivation were evaluated on nematode communities associated with tall fescue turfgrass following 11 years of treatment applications. Fertilizer treatments of biosolid, synthetic, and plant-based fertilizers and cultivation treatments of 0×, 1×, and 2× aerification passes were applied to randomized and replicated tall fescue plots at the University of Maryland Paint Branch Turfgrass facility in College Park, Maryland. Free-living and plant-parasitic nematodes were identified, enumerated, and categorized into functional groups. Nematode count data were compared using generalized linear mixed modeling with negative binomial distribution and two-way ANOVA was used to compare nematode ecological indices. Biosolid treatments resulted in lower omnivore-predator densities than plant-based fertilizer treatments (p ≤ .001) and significantly greater Hoplolaimus densities than plant-based fertilizer plots (p ≤ .05). Synthetic fertilizer applications resulted in the greatest Eucephalobus (p ≤ .05) and total bacterivore densities (p ≤ .001) of all fertilizer treatments. Plant-based fertilizer-treated plots had the largest Maturity Index cp 2-5 and Structure Index (p ≤ .05). Cultivation of 1× resulted in fewer total bacterivore densities than 2× (p ≤ .01) while omnivore-predator densities were greater in 1× than 0× (p ≤ .001). Plant health, as measured by NDVI, was lowest in biosolid-treated turfgrass (p ≤ .05). These findings suggest that long-term turfgrass management practices can have variable impacts on nematode abundance and community structure in tall fescue and provide insights into ecological impacts of turfgrass management practices.

Sublethal effects of imidacloprid-contaminated honey stores on colony performance, queens, and worker activities in fall and early winter colonies.

Neonicotinoid-contaminated sugar stores can have both near term and long term effects on honey bees due to their persistence in honey stores. Effects of imidacloprid food stores contaminants were examined in subtropical colonies that experience reduced brood rearing and foraging during overwintering. Colonies were given treatment sugar syrup containing 0 ppb (control), 20 ppb (field relevant), or 100 ppb (above field relevant) imidacloprid over six weeks to simulate contaminated fall nectar. Colonies were evaluated immediately (post-treatment) and 10 weeks (mid-winter) after treatment to compare proximal and latent effects. Post-treatment 0 ppb and 20 ppb colonies had more workers than 100 ppb colonies while 0 ppb colonies more brood than 20 ppb or 100 ppb colonies. Mid-winter 0 ppb and 20 ppb colonies had more workers than 100 ppb colonies and 0 ppb colonies more brood than 100 ppb colonies. Colonies experienced seasonal declines in stored pollen but no treatment effects. Lower 100 ppb colony performance was associated with reduced effort rather than lifespan. RFID (Radio Frequency Identification) tracking revealed that workers had similar adult lifespans across treatments; however, 100 ppb workers engaged in activities outside the colony for less time than 0 ppb workers. Imidacloprid exposure affected queen but not worker nutritional physiology. Nurses retained well-developed hypopharyngeal glands (as indicated by head protein) across treatments. Mid-winter queens from 0 ppb colonies had marginally higher ovary protein than queens from 100 ppb colonies and more ovary lipids than queens from 20 ppb colonies. However, queen nutrient stores in non-reproductive tissues (fat bodies) did not differ across treatments. Queens from different treatments were attended by comparable numbers of retinue workers and had similar gland contents of four QMP (Queen Mandibular Pheromone) components essential to queen care. High levels of imidacloprid in sugar stores can negatively affect colony performance months after initial storage.

Honey bee retinue workers respond similarly to queens despite seasonal differences in Queen Mandibular Pheromone (QMP) signaling.

Honey bee colonies maintain viable queens in part through communication with Queen Mandibular Pheromone (QMP), a mixture that signals the queen's presence and reproductive quality to workers. In turn, workers are thought to provide retinue queen care or replace queens partially based on QMP profiles. We examined the effects of seasonal dearth (overwintering in a warm subtropical location) on queen-worker interactions. Retinue worker responses to continuously ovipositing queens were considered in view of QMP signaling and queen reproductive quality. QMP signaling was estimated from QMP residues recovered from nest worker bodies, which is the primary mode of QMP transfer from the queen to the colony at large. QMP residues varied seasonally but not at all with queen reproductive quality (spermatheca sperm storage, ovary protein and lipid contents). 9-HDA and 9-ODA were lower in January than other months. HOB decreased from July to January, while HVA, a component associated with mated queens, increased sharply in January. Despite these seasonal signaling differences, retinue workers attended queens at similar levels through the months. In terms of reproductive quality, queens did not differ over the months in matedness (spermatheca sperm storage) or physiological age (protein carbonyl content), but varied in nutrient allocation to reproductive and non-reproductive tissues. Queen ovaries contained more protein in September than in November, and more lipid in July and September than in November and January. Queen fat bodies had more protein in July than September or November, but less lipid in July and September than November or January. Retinue worker responses did not vary with seasonal QMP changes, but reflected overall continuous brood rearing efforts and queen matedness throughout the year. The absence of seasonal differences in worker responses to QMP should be considered in the broader context of continuous reproductive efforts in warm subtropical colonies.

Diet and pheromones interact to shape honey bee (Apis mellifera) worker physiology.

Honey bee colony health is a function of the individuals, their interactions, and the environment. A major goal of honey bee research is to understand how colonies respond to stress. Individual-level studies of the bee stress response are tractable, but their results do not always translate to the colony level. Nutritional stress is an important factor in colony declines. Nutrition studies are typically conducted on individual nurse workers (nurses), who are primarily responsible for converting pollen into brood. Nurse physiology is sensitive to both pollen and pheromones, which communicate signals among colony members. Here, we asked whether pheromones influence nurse nutrient pathways involved in brood care, and whether diet influences colony communication. We exposed caged, nurse-aged workers to different combinations of pheromones and pollen, and measured traits related to brood care. We found that pheromones enhanced pollen-dependent processes such as hypopharyngeal gland growth and mrjp1 expression, and buffered the negative effects of starvation. Pollen also enhanced how nurse phenotypes respond to pheromones. Therefore, diet and pheromones interact to influence nurse nutritional physiology and aspects of brood care. These findings have implications for studying colony function and health in an increasingly stressful climate.

Correction: Honey bees preferentially consume freshly-stored pollen.

[This corrects the article DOI: 10.1371/journal.pone.0175933.].

Exposure to sublethal concentrations of methoxyfenozide disrupts honey bee colony activity and thermoregulation.

Methoxyfenozide is an insect growth regulator (IGR) commonly used in agriculture to simultaneously control pests and preserve beneficial insect populations; however, its impact on honey bees in not fully understood. We conducted field and laboratory experiments to investigate bee health in response to field-relevant concentrations of this pesticide. Significant effects were observed in honey bee colony flight activity and thermoregulation after being exposed over 9 weeks to supplemental protein patty containing methoxyfenozide. Compared to bee colonies in the control group, colonies fed pollen patty with 200 ppb methoxyfenozide (as measured by residue analysis) had: 1) a significantly reduced rate of weight loss due to forager departure in the morning; and 2) higher temperature variability during the winter. Colonies in the 100 ppb (as measured by residue analysis) treatment group had values between the 200 ppb group and control for both response variables. The dusk break point, which is the time associated with the end of forager return, differed among all treatment groups but may have been confounded with direction the hives were facing. Bee colony metrics of adult bee mass and brood surface area, and measurements of bee head weight, newly-emerged bee weight, and hypopharyngeal gland size were not significantly affected by methoxyfenozide exposure, suggesting that there may be significant effects on honey bee colony behavior and health in the field that are difficult to detect using standard methods for assessing bee colonies and individuals. The second experiment was continued into the following spring, using the same treatment groups as in the fall. Fewer differences were observed among groups in the spring than the fall, possibly because of abundant spring forage and consequent reduced treatment patty consumption. Residue analyses showed that: 1) observed methoxyfenozide concentrations in treatment patty were about 18-60% lower than the calculated concentrations; 2) no residues were observed in wax in any treatment; and 3) methoxyfenozide was detected in bee bread only in the 200 ppb treatment group, at about 1-2.5% of the observed patty concentration.

Honey bees overwintering in a southern climate: longitudinal effects of nutrition and queen age on colony-level molecular physiology and performance.

Honey bee colony nutritional ecology relies on the acquisition and assimilation of floral resources across a landscape with changing forage conditions. Here, we examined the impact of nutrition and queen age on colony health across extended periods of reduced forage in a southern climate. We measured conventional hive metrics as well as colony-level gene expression of eight immune-related genes and three recently identified homologs of vitellogenin (vg), a storage glycolipoprotein central to colony nutritional state, immunity, oxidative stress resistance and life span regulation. Across three apiary sites, concurrent longitudinal changes in colony-level gene expression and nutritional state reflected the production of diutinus (winter) bees physiologically altered for long-term nutrient storage. Brood production by young queens was significantly greater than that of old queens, and was augmented by feeding colonies supplemental pollen. Expression analyses of recently identified vg homologs (vg-like-A, -B, and -C) revealed distinct patterns that correlated with colony performance, phenology, and immune-related gene transcript levels. Our findings provide new insights into dynamics underlying managed colony performance on a large scale. Colony-level, molecular physiological profiling is a promising approach to effectively identify factors influencing honey bee health in future landscape and nutrition studies.

Using within-day hive weight changes to measure environmental effects on honey bee colonies.

Patterns in within-day hive weight data from two independent datasets in Arizona and California were modeled using piecewise regression, and analyzed with respect to honey bee colony behavior and landscape effects. The regression analysis yielded information on the start and finish of a colony's daily activity cycle, hive weight change at night, hive weight loss due to departing foragers and weight gain due to returning foragers. Assumptions about the meaning of the timing and size of the morning weight changes were tested in a third study by delaying the forager departure times from one to three hours using screen entrance gates. A regression of planned vs. observed departure delays showed that the initial hive weight loss around dawn was largely due to foragers. In a similar experiment in Australia, hive weight loss due to departing foragers in the morning was correlated with net bee traffic (difference between the number of departing bees and the number of arriving bees) and from those data the payload of the arriving bees was estimated to be 0.02 g. The piecewise regression approach was then used to analyze a fifth study involving hives with and without access to natural forage. The analysis showed that, during a commercial pollination event, hives with previous access to forage had a significantly higher rate of weight gain as the foragers returned in the afternoon, and, in the weeks after the pollination event, a significantly higher rate of weight loss in the morning, as foragers departed. This combination of continuous weight data and piecewise regression proved effective in detecting treatment differences in foraging activity that other methods failed to detect.

Longitudinal Effects of Supplemental Forage on the Honey Bee (Apis mellifera) Microbiota and Inter- and Intra-Colony Variability.

Honey bees (Apis mellifera) provide vital pollination services for a variety of agricultural crops around the world and are known to host a consistent core bacterial microbiome. This symbiotic microbial community is essential to many facets of bee health, including likely nutrient acquisition, disease prevention and optimal physiological function. Being that the bee microbiome is likely involved in the digestion of nutrients, we either provided or excluded honey bee colonies from supplemental floral forage before being used for almond pollination. We then used 16S rRNA gene sequencing to examine the effects of forage treatment on the bees' microbial gut communities over four months. In agreement with previous studies, we found that the honey bee gut microbiota is quite stable over time. Similarly, we compared the gut communities of bees from separate colonies and sisters sampled from within the same hive over four months. Surprisingly, we found that the gut microbial communities of individual sisters from the same colony can exhibit as much variation as bees from different colonies. Supplemental floral forage had a subtle effect on the composition of the microbiome during the month of March only, with strains of Gilliamella apicola, Lactobacillus, and Bartonella being less proportionally abundant in bees exposed to forage in the winter. Collectively, our findings show that there is unexpected longitudinal variation within the gut microbial communities of sister honey bees and that supplemental floral forage can subtly alter the microbiome of managed honey bees.

Text Simplification Using Consumer Health Vocabulary to Generate Patient-Centered Radiology Reporting: Translation and Evaluation.

BACKGROUND: Radiology reporting is a clinically oriented form of documentation that reflects critical information for patients about their health care processes. Realizing its importance, many medical institutions have started providing radiology reports in patient portals. The gain, however, can be limited because of medical language barriers, which require a way for customizing these reports for patients. The open-access, collaborative consumer health vocabulary (CHV) is a terminology system created for such purposes and can be the basis of lexical simplification processes for clinical notes. OBJECTIVE: The aim of this study was to examine the comprehensibility and suitability of CHV in simplifying radiology reports for consumers. This was done by characterizing the content coverage and the lexical similarity between the terms in the reports and the CHV-preferred terms. METHODS: The overall procedure was divided into the following two main stages: (1) translation and (2) evaluation. The translation process involved using MetaMap to link terms in the reports to CHV concepts. This is followed by replacing the terms with CHV-preferred terms using the concept names and sources table (MRCONSO) in the Unified Medical Language System (UMLS) Metathesaurus. In the second stage, medical terms in the reports and general terms that are used to describe medical phenomena were selected and evaluated by comparing the words in the original reports with the translated ones. The evaluation includes measuring the content coverage, investigating lexical similarity, and finding trends in missing concepts. RESULTS: Of the 792 terms selected from the radiology reports, 695 of them could be mapped directly to CHV concepts, indicating a content coverage of 88.5%. A total of 51 of the concepts (53%, 51/97) that could not be mapped are names of human anatomical structures and regions, followed by 28 anatomical descriptions and pathological variations (29%, 28/97). In addition, 12 radiology techniques and projections represented 12% of the unmapped concepts, whereas the remaining six concepts (6%, 12/97) were physiological descriptions. The rate of lexical similarity between the CHV-preferred terms and the terms in the radiology reports was approximately 72.6%. CONCLUSIONS: The CHV covered a high percentage of concepts found in the radiology reports, but unmapped concepts are associated with areas that are commonly found in radiology reporting. CHV terms also showed a high percentage of lexical similarity with terms in the reports, which contain a myriad of medical jargon. This suggests that many CHV terms might not be suitable for lay consumers who would not be facile with radiology-specific vocabulary. Therefore, further patient-centered content changes are needed of the CHV to increase its usefulness and facilitate its integration into consumer-oriented applications.

Honey bees preferentially consume freshly-stored pollen.

Honey bees (Apis mellifera) collect and store both honey and pollen in preserved forms. Pollen storage involves the addition of honey or nectar and oral secretions to pollen granules. It is controversial whether the duration of pollen storage alters the palatability or nutritive value of the pollen storage medium. We examined how bees utilize different-aged stored pollen during an extended pollen flow. The deposition of pollen into wax cells and subsequent consumption were monitored daily on 18 brood frames from 6 colonies over an 8d observation period. Despite a greater abundance of older stored pollen cells on brood frames, bees showed a marked preference for the consumption of freshly-stored pollen. Two to four day-old pollen cell contents were significantly more likely to be consumed, while pollen cell contents more than seven days old were eaten at much lower rates. Similar experiments that controlled for cell abundance and spatial effects using cage assays yielded the same result. One day-old stored pollen was consumed approximately three times more often than 10d-old stored pollen, and two times more often than 5d-old stored pollen. These consumption preferences for freshly-stored pollen occurred despite a lack of clear developmental advantages. Young adult workers reared for 7 days on 1d-, 5d-, or 10d-old stored pollen showed no difference in body mass, stored pollen consumption, hindgut fecal material accumulation, or hypopharyngeal gland (HPG) protein titers, suggesting that different-aged pollen stores did not vary in their nutritional value to adult bees. These findings are inconsistent with the hypothesis promoting a period of microbially-mediated, "beebread maturation" that results in greater palatability or nutritive value for aged pollen stores. Rather, stored pollen that is not eaten in the first few days accumulates as excess stores preserved in a less preferred, but nutritionally-similar state.

Sublethal Effects of Imidacloprid on Honey Bee Colony Growth and Activity at Three Sites in the U.S.

Imidacloprid is a neonicotinoid pesticide heavily used by the agricultural industry and shown to have negative impacts on honey bees above certain concentrations. We evaluated the effects of different imidacloprid concentrations in sugar syrup using cage and field studies, and across different environments. Honey bee colonies fed sublethal concentrations of imidicloprid (0, 5, 20 and 100 ppb) over 6 weeks in field trials at a desert site (Arizona), a site near intensive agriculture (Arkansas) and a site with little nearby agriculture but abundant natural forage (Mississippi) were monitored with respect to colony metrics, such as adult bee and brood population sizes, as well as pesticide residues. Hive weight and internal hive temperature were monitored continuously over two trials in Arizona. Colonies fed 100 ppb imidacloprid in Arizona had significantly lower adult bee populations, brood surface areas and average frame weights, and reduced temperature control, compared to colonies in one or more of the other treatment groups, and consumption rates of those colonies were lower compared to other colonies in Arizona and Arkansas, although no differences in capped brood or average frame weight were observed among treatments in Arkansas. At the Mississippi site, also rich in alternative forage, colonies fed 5 ppb imidacloprid had less capped brood than control colonies, but contamination of control colonies was detected. In contrast, significantly higher daily hive weight variability among colonies fed 5 ppb imidacloprid in Arizona suggested greater foraging activity during a nectar flow post treatment, than any other treatment group. Imidacloprid concentrations in stored honey corresponded well with the respective syrup concentrations fed to the colonies and remained stable within the hive for at least 7 months after the end of treatment.

Hive-stored pollen of honey bees: many lines of evidence are consistent with pollen preservation, not nutrient conversion.

Honey bee hives are filled with stored pollen, honey, plant resins and wax, all antimicrobial to differing degrees. Stored pollen is the nutritionally rich currency used for colony growth and consists of 40-50% simple sugars. Many studies speculate that prior to consumption by bees, stored pollen undergoes long-term nutrient conversion, becoming more nutritious 'bee bread' as microbes predigest the pollen. We quantified both structural and functional aspects associated with this hypothesis using behavioural assays, bacterial plate counts, microscopy and 454 amplicon sequencing of the 16S rRNA gene from both newly collected and hive-stored pollen. We found that bees preferentially consume fresh pollen stored for <3 days. Newly collected pollen contained few bacteria, values which decreased significantly as pollen were stored >96 h. The estimated microbe to pollen grain surface area ratio was 1:1 000 000 indicating a negligible effect of microbial metabolism on hive-stored pollen. Consistent with these findings, hive-stored pollen grains did not appear compromised according to microscopy. Based on year round 454 amplicon sequencing, bacterial communities of newly collected and hive-stored pollen did not differ, indicating the lack of an emergent microbial community co-evolved to digest stored pollen. In accord with previous culturing and 16S cloning, acid resistant and osmotolerant bacteria like Lactobacillus kunkeei were found in greatest abundance in stored pollen, consistent with the harsh character of this microenvironment. We conclude that stored pollen is not evolved for microbially mediated nutrient conversion, but is a preservative environment due primarily to added honey, nectar, bee secretions and properties of pollen itself.

An amino acid substitution inhibits specialist herbivore production of an antagonist effector and recovers insect-induced plant defenses.

Plants respond to insect herbivory through the production of biochemicals that function as either direct defenses or indirect defenses via the attraction of natural enemies. While attack by closely related insect pests can result in distinctive levels of induced plant defenses, precise biochemical mechanisms responsible for differing responses remain largely unknown. Cowpea (Vigna unguiculata) responds to Fall armyworm (Spodoptera frugiperda) herbivory through the detection of fragments of chloroplastic ATP synthase γ-subunit proteins, termed inceptin-related peptides, present in larval oral secretions (OS). In contrast to generalists like Fall armyworm, OS of the legume-specializing velvetbean caterpillar (VBC; Anticarsia gemmatalis) do not elicit ethylene production and demonstrate significantly lower induced volatile emission in direct herbivory comparisons. Unlike all other Lepidoptera OS examined, which preferentially contain inceptin (Vu-In; +ICDINGVCVDA-), VBC OS contain predominantly a C-terminal truncated peptide, Vu-In(-A) (+ICDINGVCVD-). Vu-In(-A) is both inactive and functions as a potent naturally occurring antagonist of Vu-In-induced responses. To block antagonist production, amino acid substitutions at the C terminus were screened for differences in VBC gut proteolysis. A valine-substituted peptide (Vu-In(ΔV); +ICDINGVCVDV-) retaining full elicitor activity was found to accumulate in VBC OS. Compared with the native polypeptide, VBC that previously ingested 500 pmol of the valine-modified chloroplastic ATP synthase γ-subunit precursor elicited significantly stronger plant responses in herbivory assays. We demonstrate that a specialist herbivore minimizes the activation of defenses by converting an elicitor into an antagonist effector and identify an amino acid substitution that recovers these induced plant defenses to a level observed with generalist herbivores.

The attraction of Spodoptera frugiperda neonates to cowpea seedlings is mediated by volatiles induced by conspecific herbivory and the elicitor inceptin.

Neonate fall armyworms [FAW; Spodoptera frugiperda (Smith)] often encounter conspecific herbivore damage as they disperse from an egg mass to an initial feeding site. We investigated the orientation responses of dispersing neonates to herbivore damage in cowpea seedlings, specifically examining whether neonate behaviors were affected by inceptin, the primary elicitor of FAW-induced defenses in cowpea leaves. We focused on responses to damage caused by conspecific first instars, as might occur during the dispersal of siblings from an egg mass. Inceptin contents of damaging first instar FAW were controlled through their diets, with leaf-fed FAW producing inceptins in their oral secretions, and root-fed or starved FAW lacking these elicitors. In a bioassay designed to evaluate neonate dispersal off a host plant, a higher percentage of neonates remained on herbivore-induced or inceptin-treated plants than on undamaged plants, mechanically damaged plants, freshly damaged plants, or on plants damaged by FAW lacking inceptins. Further investigations of neonate responses to plant odors with a four-arm olfactometer demonstrated that neonate attraction to odors from 4-h old FAW damage was strongly dependent on previous diet of the damaging larvae. Neonates were attracted to odors from 4-h old FAW damage over odors from undamaged plants or fresh FAW damage, provided that the damaging larvae had previously ingested leaf material. In a direct comparison of odors from induced plants, plants damaged by leaf-fed FAW were as attractive as plants treated with synthetic inceptin. GC-MS analysis confirmed that (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) was the major volatile induced by FAW herbivory. While both DMNT and undamaged plant odors were more attractive than air, neonates preferred DMNT-supplemented plant odors. These results suggest that neonate FAW exploit herbivore-induced plant volatiles as host plant location and recognition cues.

Cowpea chloroplastic ATP synthase is the source of multiple plant defense elicitors during insect herbivory.

In cowpea (Vigna unguiculata), fall armyworm (Spodoptera frugiperda) herbivory and oral secretions (OS) elicit phytohormone production and volatile emission due to inceptin [Vu-In; (+)ICDINGVCVDA(-)], a peptide derived from chloroplastic ATP synthase gamma-subunit (cATPC) proteins. Elicitor-induced plant volatiles can function as attractants for natural enemies of insect herbivores. We hypothesized that inceptins are gut proteolysis products and that larval OS should contain a mixture of related peptides. In this study, we identified three additional cATPC fragments, namely Vu-(GE+)In [(+)GEICDINGVCVDA(-)], Vu-(E+)In [(+)EICDINGVCVDA(-)], and Vu-In(-A) [(+)ICDINGVCVD(-)]. Leaf bioassays for induced ethylene (E) production demonstrated similar effective concentration(50) values of 68, 45, and 87 fmol leaf(-1) for Vu-In, Vu-(E+)In, and Vu-(GE+)In, respectively; however, Vu-In(-A) proved inactive. Shortly following ingestion of recombinant proteins harboring cATPC sequences, larval OS revealed similar concentrations of the three elicitors with 80% of the potential inceptin-related peptides recovered. Rapidly shifting peptide ratios over time were consistent with continued proteolysis and preferential stability of inceptin. Likewise, larvae ingesting host plants with inceptin precursors containing an internal trypsin cleavage site rapidly lost OS-based elicitor activity. OS containing inceptin elicited a rapid and sequential induction of defense-related phytohormones jasmonic acid, E, and salicylic acid at 30, 120, and 240 min, respectively, and also the volatile (E)-4,8-dimethyl-1,3,7-nonatriene. Similar to established peptide signals such as systemin and flg22, amino acid substitutions of Vu-In demonstrate an essential role for aspartic acid residues and an unaltered C terminus. In cowpea, insect gut proteolysis following herbivory generates inappropriate fragments of an essential metabolic enzyme enabling plant non-self-recognition.

Attraction of Spodoptera frugiperda larvae to volatiles from herbivore-damaged maize seedlings.

Plants respond to insect attack with the induction of volatiles that function as indirect plant defenses through the attraction of natural enemies to the herbivores. Despite the fact that volatiles are induced in response to caterpillar attack, their reciprocal effects on the host location behaviors of the same foraging herbivores are poorly understood. We examined orientation responses of sixth instar fall armyworm [FAW; Spodoptera frugiperda (Smith)] to odors from herbivore-damaged and undamaged maize seedlings (Zea mays var. Golden Queen) in y-tube olfactometer bioassays. While both damaged and undamaged maize seedlings were attractive compared with air, sixth instars preferred odors from damaged maize seedlings over odors from undamaged maize seedlings. Gas chromatography-mass spectrometry analysis of plant volatiles revealed that linalool and 4,8-dimethyl-1,3,7-nonatriene were the major volatiles induced by FAW herbivory 6 hr after initial damage. Given its prominence in induced plants and established attractiveness to adult FAW, linalool was evaluated both as an individual attractant and as a supplemental component of whole plant odors. Volatile linalool was more attractive than air to sixth instar FAW over a broad range of release rates. FAW also responded selectively to different amounts of linalool, preferring the higher amount. The orientation preferences of FAW were readily manipulated through capillary release of linalool into the airstream of whole plant odors. FAW preferred linalool over undamaged plant odors, and linalool-supplemented plant odors over unsupplemented plant odors, indicating that olfactory preferences could be changed by alteration of a single volatile component. These results suggest that although many induced volatiles attract natural enemies of herbivores, these defenses may also inadvertently recruit more larval herbivores to an attacked plant or neighboring conspecifics.

Fragments of ATP synthase mediate plant perception of insect attack.

Plants can perceive a wide range of biotic attackers and respond with targeted induced defenses. Specificity in plant non-self-recognition occurs either directly by perception of pest-derived elicitors or indirectly through resistance protein recognition of host targets that are inappropriately proteolyzed. Indirect plant perception can occur during interactions with pathogens, yet evidence for analogous events mediating the detection of insect herbivores remains elusive. Here we report indirect perception of herbivory in cowpea (Vigna unguiculata) plants attacked by fall armyworm (Spodoptera frugiperda) larvae. We isolated and identified a disulfide-bridged peptide (+ICDINGVCVDA-), termed inceptin, from S. frugiperda larval oral secretions that promotes cowpea ethylene production at 1 fmol leaf(-1) and triggers increases in the defense-related phytohormones salicylic acid and jasmonic acid. Inceptins are proteolytic fragments of chloroplastic ATP synthase gamma-subunit regulatory regions that mediate plant perception of herbivory through the induction of volatile, phenylpropanoid, and protease inhibitor defenses. Only S. frugiperda larvae that previously ingested chloroplastic ATP synthase gamma-subunit proteins and produced inceptins significantly induced cowpea defenses after herbivory. Digestive fragments of an ancient and essential plant enzyme, inceptin functions as a potent indirect signal initiating specific plant responses to insect attack.

Lutein sequestration and furanocoumarin metabolism in parsnip webworms under different ultraviolet light regimes in the montane west.

Both biotic and abiotic selection pressures can contribute to geographic variation in allelochemical production in plants. We examined furanocoumarin production in western North American populations of Heracleum lanatum and Pastinaca sativa that, at different latitudes and altitudes, experience different ultraviolet (UV) light regimes. Total furanocoumarins and linear furanocoumarins of fruits were negatively correlated with UV irradiance, whereas amounts of angular furanocoumarins, which are generally less phototoxic, were not. Another factor potentially influencing furanocoumarin production is the presence of the parsnip webworm Depressaria pastinacella, (Lepidoptera: Oecophoridae), an herbivore that feeds on reproductive structures of both plant species. These insects sequester lutein from their host plants; this carotenoid acts to ameliorate furanocoumarin toxicity. Although the concentration of lutein in fruits did not vary with UV irradiance, lutein sequestration by sixth instars was positively correlated with UV irradiance. Webworm populations are variably infested with the polyembryonic webworm parasitoid Copidosoma sosares Walker (Hymenoptera: Encyrtidae). H. lanatum fruits from populations with webworms parasitized by C. sosares had lower concentrations of furanocoumarins, with the exception of sphondin, than fruits from plants infested with webworms free from parasitism. Lower levels of these furanocoumarins may reduce negative effects on the fitness of this parasitoid. In contrast with the variation in furanocoumarin content, the ability of webworms to metabolize furanocoumarins by cytochrome P450 did not differ significantly among populations from New Mexico to Alberta.