Onion fly and little house fly host finding selectively mediated by decomposing onion and microbial volatiles

Responses of onion flies,Hylemya antiqua (Meigen), to various synthetic onion and microbial volatiles as well as volatiles from microbial cultures and decomposing onions were tested to characterize the most effective host-finding stimuli. Of nine onion and microbial volatiles tested individually, only the known attractant,n-dipropyl disulfide, caught significant numbers of flies. However, a blend of these volatiles attracted more flies than any single chemical, includingn-dipropyl disulfide. In another experiment, agar plates inoculated with microorganisms from decomposing onions did not attract onion flies. However, cut onions inoculated with microorganisms and conditioned 4 days caught more onion flies than freshly cut onions andn-dipropyl disulfide. These results suggest that a blend of chemicals, rather than a single key chemical, is the more effective host-finding stimulus, and that microbial activity enhances the attractancy of a blend of onion volatiles. Large numbers ofFannia canicularis (L.), the little house fly, responded to the microbial cultures, demonstrating the existence of a potent attractant for this muscid.Diptera: Anthomyiidae.Diptera: Muscidae.Paper No. 9470 of the Michigan State University Agricultural Experiment Station. Received for publication May 23, 1980.     

Chemical attractants tested against the Australian bush flyMusca vetustissima (Diptera: Muscidae)

A number of chemical compositions known to attract several synanthropic and pest fly species were tested against the Australian bush flyMusca vetustissima, a severe and persistent pest of man and domesticated and wild mammals. A standard composition containing small quantities of trimethylamine and indole, blended with large amounts of ammonium sulfate and anchovy meal, showed good attractancy against this fly. Deletion of ammonium sulfate did not significantly alter attractancy. Similarly, incorporation ofn-butyric acid and linoleic acid into the formulation did not alter attractancy, nor did addition of several carbonates and bicarbonates. Skatole, a closely related compound to indole, decreased attractancy significantly. In all the tests, the attractant compositions lured greater number of females than males. The attractancy of the standard composition was studied and compared with equal amount of fresh cattle dung. Chemical attractants lured bush flies and not dung beetles, while dung attracted both flies and beetles. The dung resource was 4.6 × more attractive to flies than the standard attractant composition. This suggests that there are other chemical attractants emanating from dung that warrant isolation and identification work. Nevertheless, the chemical attractants studied here provide a starting point for further investigations. The ratio of female to male flies attracted to humans was greater than the ratio of flies attracted to chemicals. The sex ratio of flies netted from human host was 3.0, while the sex ratio of flies attracted to chemical attractants was 1.8. The preponderance of females attracted to both sources was probably due to physiological requirements of the sexes.     

A novel attractant for Mexican fruit fly,Anastrepha ludens,from fermented host fruit

Chemicals from fermented chapote fruit were identified and evaluated as attractants for hungry adult Mexican fruit flies in laboratory and greenhouse bioassays. Twenty-eight chemicals identified from an attractive gas-chromatography fraction were as attractive as a chapote volatiles extract (CV) when mixed in the same amounts found in CV. Sixteen of the chemicals were slightly attractive to flies when tested individually. A mixture containing 15 of the chemicals by design and the 16th as an impurity, in arbitrary concentrations, was at least as attractive as the original CV. In a series of experiments, the number of chemicals was reduced to three by elimination of unnecessary components. The three-component mixture retained the attractiveness of the 15-component mixture. The three chemicals were 1,8-cineole, ethyl hexanoate, and hexanol (CEH). Attractiveness of the three-chemical mixture was equal to the sum of the attractiveness of the three individual components, suggesting that each chemical binds to a different receptor type that independently elicits partial attraction behavior. Optimal ratios were 1011 of the three chemicals, respectively. Optimal test quantities ranged between 0.4–4g of 1,8-cineole and 40–400 ng each of ethyl hexanoate and hexanol applied to filter paper in the laboratory bioassays. A neat 1011 mixture of the chemicals was 1.8 times more attractive than aqueous solutions ofTorula dried yeast and borax to starved 2-day-old flies when the lures were tested in competing McPhail traps in a large greenhouse cage.Diptera: Tephritidae.     

Partial characterization and HPLC isolation of bacteria-produced attractants for the Mexican fruit fly,Anastrepha ludens

Methods were developed to collect and isolate volatile chemicals produced by aStaphylococcus bacterium in tryptic soy culture that are attractive to protein-hungry adult Mexican fruit flies. Centrifugation of bacteria culture yielded a slightly attractive pellet containing most of the bacteria cells and a highly attractive supernatant. Supernatant filtered to remove the remaining bacteria was as attractive as the unfiltered supernatant. Filtrate at pH 7 and above was much more attractive than filtrate at pH 5 and below. Most of the attractiveness was retained on strong cation exchange media under acidic conditions and eluted with base. Attractive principles could not be trapped on adsorbents such as Porapak Q or extracted with organic solvents from aqueous preparations, but they were easily collected by headspace sweeping with steam. The attractive components were efficiently concentrated by rotary evaporation of steam distillate at pH 5, but at higher pH much of the attractiveness distilled. A reverse-phase HPLC method using a negative counter-ion was developed to separate and collect attractive components of concentrated steam distillate. Attractive fractions collected using this method were concentrated and injected onto silica HPLC. Activity eluted from silica in two distinct bands. Results suggest that the most attractive components of the bacterial odor are highly polar, low-molecular-weight amines.Diptera: Tephritidae.     

Identification of odors from overripe mango that attract vinegar flies,Drosophila melanogaster

Bioassays with a variety of overripe fruits, including mango, plum, pear, and grape, and their extracts showed that odors from overripe mango were most attractive to adult vinegar flies, Drosophila melanogaster. Combined gas chromatography–electroantennographic detection (GC-EAD) analyses of solid-phase microextraction (SPME) and Tenax extracts of overripe mango odors showed that several volatile compounds, including ethanol, acetic acid, amyl acetate, 2-phenylethanol, and phenylethyl acetate elicited significant EAG responses from antennae of female flies. Most of the volatile compounds in the extracts were identified by mass spectral and retention index comparisons with synthetic standards. In cage bioassays, lures with a blend of ethanol, acetic acid, and 2-phenylethanol in a ratio of 1:22:5 attracted six times more flies than any single EAG-active compound. This blend also attracted four times more flies than traps baited with overripe mango or unripe mango. However, in field trials, the blend was not as attractive as suggested by the laboratory bioassay.     

Chemically Mediated Host-Plant Selection by the Milfoil Weevil: A Freshwater Insect–Plant Interaction

The milfoil weevil Euhrychiopsis lecontei is a specialist aquatic herbivore that feeds, oviposits, and mates on the invasive freshwater macrophyte Myriophyllum spicatum. We characterized the weevil's preference for M. spicatum, and through bioassay-driven fractionation, isolated and identified two chemicals released by M. spicatum that attract E. lecontei. Mass spectrometry and nuclear magnetic resonance spectroscopy were used to identify the attractive compounds as glycerol and uracil. Dose-response curves for glycerol and uracil indicated that weevil preference increased as sample concentration increased. Weevils were attracted to a crude sample of M. spicatum-released chemicals from 0.17 to 17 mg/l, to glycerol from 18 to 1800 μM (0.0017–0.17 mg/l), and to uracil from 0.015 to 15 μM (0.00014–1.4 mg/l). Although glycerol and uracil are ubiquitous, weevils are likely responding to high concentrations that are released as a result of the rapid growth of M. spicatum. Uracil concentration was greater in the exudates of M.spicatum than other Myriophyllum spp. E. lecontei was attracted to glycerol at a concentration similar to that at which terrestrial insects are attracted to sugar alcohols. This is the first example of a freshwater specialist insect being attracted to chemicals released by its host plant. Analysis of the water milfoil–weevil interaction provides further understanding as to how insects locate their host plants in aquatic systems.     

Purine metabolizing capability of Enterobacter agglomerans affects volatiles production and attractiveness to Mexican fruit fly

We investigated two strains of Enterobacter agglomerans that differ in their ability to metabolize uric acid for (1) attractiveness to sugar-fed Mexican fruit flies, and (2) production of volatile chemicals that may be responsible for the attractiveness. The two strains were cultured on a medium that contained uric acid as the primary nitrogen source to simulate bird feces, a natural substrate for this bacterium. Active cultures of both strains were more attractive than uninoculated uric acid medium to both sexes of sugar-fed flies in wind-tunnel bioassays. The uricase(+) strain was more attractive than the uricase(–) strain to males and to females <9 days old, but not to older females. Volatiles found by solid-phase microextraction in greater amounts in headspace above active cultures of both strains than above uninoculated medium were ammonia, dimethyldisulfide, 3-methylbutanol, 2-phenylethanol, 2,5-dimethylpyrazine, and trimethylpyrazine. The uricase(+) strain produced more ammonia, dimethyldisulfide, and trimethylpyrazine than the uricase(–) strain. An additional chemical, 3-hydroxybutanone, appears to be produced exclusively by the uricase(+) strain. The uricase(–) strain produced more 2-phenylethanol than the uricase(+) strain. Differences in volatiles are consistent with the generally greater attractiveness of the uricase(+) strain compared with the uricase(–) strain as ammonia, 3-hydroxybutanone, and trimethylpyrazine have been demonstrated attractive to sugar-fed Mexican fruit flies.     

Attractants for synanthropic flies

Fermented molasses or sucrose solutions are known to attract several species of filth-breeding flies. To identify the volatile attractants produced in fermenting sucrose solutions with yeast, these solutions were fractionated, and the chemical constituents identified and bioassayed against filth-breeding flies includingFannia canicularis (L.),Muscina stabulans (Fallén), andMusca domestica (L.). Distillation of a fermented sucrose solution gave an active distillate and an inactive residue. Gas Chromatographic analysis of the distillate showed the presence of acetaldehyde, ethyl acetate, ethanol, 1-pro-panol, 2-methyl-l-propanol, and 3-methyl-l-butanol. Ethanol constituted by far the greatest proportion of compounds present in the distillate. An aqueous solution of ethanol exhibited the same level of attractancy as the distillate, the fermented sucrose solution, and a reconstituted distillate containing all compounds identified. Ethanol was thus identified as the sole attractant emanated from fermented carbohydrate solutions that elicited positive responses in pest flies, especially inF. canicularis.Diptera: Muscidae.     

Identification of Floral Volatiles From Ligustrum japonicum that Stimulate Flower-Visiting by Cabbage Butterfly, Pieris rapae

Floral scent compounds of Ligustrum japonicum that affect the foraging behavior of Pieris rapae adults were examined by means of chemical analyses, electroantennogram (EAG) responses, and behavioral bioassays; the behavioral biossays consisted of two tests: reflex extension of proboscis (REP) in response to odor, and attraction to scented and unscented artificial flowers. More than 30 compounds, including 2-phenylethanol, benzyl alcohol, and methyl phenylacetate as the major components were identified from L. japonicum flowers. Of these, 22 compounds were tested for their effect on foraging behavior. Phenylacetaldehyde (PA), 2-phenylethanol (PE), and 6-methylhept-5-en-2-one (MHO) elicited the highest REP responses, and benzaldehyde (BA) and methyl phenylacetate (MPA) evoked intermediate REP responses. EAG responses were not necessarily correlated with REP activities; the three high-REP compounds gave only moderate EAG responses, whereas two other compounds (ethyl phenylacetate and 2-phenylethyl acetate) that released high EAG responses showed low REP activities. In two-choice behavioral bioassays, flower models scented with any one of these high-REP compounds attracted significantly more adults, while compounds with low REP activities exhibited weak or no appreciable attractiveness. This suggests that the REP responsiveness closely reflects the attractiveness of a compound and could be an effective measure in elucidating which chemical attractants are involved in flower-visiting. A synthetic blend of five floral chemicals (PA, PE, MHO, BA, and MPA) displayed an attractiveness that was comparable to that of the floral extract and was more effective in attractiveness than the compounds tested singly. Consequently, it is highly likely that the flower-visiting by P. rapae to L. japonicum is mediated largely by floral scent chemicals and that a synergistic effect of the five floral components would be most responsible for attraction of the butterfly to this flower. The present results also strongly suggest that specific floral volatiles may facilitate close-range flower location by P. rapae, could serve in part as a cue for recognizing food sources, and also be profoundly implicated in flower preference.     

Flying the Fly: Long-range Flight Behavior of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> to Attractive Odors

The fruit fly, Drosophila melanogaster Meigen (Diptera: Drosophilidae), is a model for how animals sense, discriminate, and respond to chemical signals. However, with D. melanogaster our knowledge of the behavioral activity of olfactory receptor ligands has relied largely on close-range attraction, rather than on long-range orientation behavior. We developed a flight assay to relate chemosensory perception to behavior. Headspace volatiles from vinegar attracted 62% of assayed flies during a 15-min experimental period. Flies responded irrespective of age, sex, and mating state, provided they had been starved. To identify behaviorally relevant chemicals from vinegar, we compared the responses to vinegar and synthetic chemicals. Stimuli were applied by a piezoelectric sprayer at known and constant release rates. Re-vaporized methanol extracts of Super Q-trapped vinegar volatiles attracted as many flies as vinegar. The main volatile component of vinegar, acetic acid, elicited significant attraction as a single compound. Two other vinegar volatiles, 2-phenyl ethanol and acetoin, produced a synergistic effect when added to acetic acid. Geosmin, a microbiological off-flavor, diminished attraction to vinegar. This wind tunnel assay based on a conspicuous and unambiguous behavioral response provides the necessary resolution for the investigation of physiologically and ecologically relevant odors and will become an essential tool for the functional analysis of the D. melanogaster olfactory system.     

(S)-2-pentadecyl acetate and 2-pentadecanone Components of aggregation pheromone ofDrosophila busckii

(S)-2-Pentadecyl acetate and 2-pentadecanone were identified as the major aggregation pheromone components, inDrosophila busckii. Both sexes of flies were attracted equally in a wind-tunnel olfactometer. The flies also responded to racemic 2-pentadecyl acetate but not to the pureR enantiomer. In bioassay, (S)-2-pentadecyl acetate and 2-pentadecanone were each active alone, and a mixture of both increased the number of flies responding ca. twofold. The aggregation pheromone components are found in the ejaculatory bulb of sexually mature males and are transferred primarily to the female cuticle during mating. One third of the pheromone transferred is released by the female to the surrounding environment in a few hours after mating. None of the aggregation pheromone components remained on the mated female's cuticle, leaving two thirds unaccounted for. The same results were obtained when racemic 2-pentadecyl acetate was topically applied to immature and mature virgin males and females. BothD. mulleri andD. busckii were attracted to (S)-2-acetates of 13, 14 and 15 carbons, butD. mulleri preferred (S)-2-tridecyl acetate andD. busckii preferred (S)-2-pentadecyl acetate.     

Aggregation pheromones ofDrosophila immigrans,D. phalerata,andD. subobscura

Aggregation pheromones ofDrosophila immigrans, D. phalerata andD. subobscura were demonstrated by testing attraction of adult flies to hexane extracts of the flies in a windtunnel bioassay. Extracts of adult males of all species attracted conspecific males and females. However,D. subobscura flies were attracted only when the extract (cVA) in the extracts of adult maleD. immigrans andD. phalerata. Both species were attracted to synthetic cVA. Male and femaleD. phalerata. Both species were attracted to synthetic cVA. Male and femaleD. subobscura produced 5,9-pentacosadiene, 5-pentacosene, 2-methylhexacosene and 5,9-heptacosadiene, while only maleD. subobscura produced (Z)-5-tricosene and minor amounts of cVA.     

Specific foraging kairomones used by a generalist parasitoid

In general, it is assumed that generalist natural enemies do not innately use specific cues for the location of their host or prey species. This hypothesis was tested using naïve females of the generalist parasitoid Lariophagus distinguendus Förster and two of its hosts, larvae of the lesser grain borer Rhyzopertha dominica (F.) and of the granary weevil Sitophilus granarius L., feeding in wheat grains. In a four-chamber olfactometer, female parasitoids were attracted to volatiles emanating from the feces of both host species. Chemical analysis of the volatiles from the feces of R. dominica revealed the presence of dominicalure 1 and 2, the species specific aggregation pheromones of R. dominica. The main compounds in the volatiles from feces of S. granarius were identified as chemicals related to mites that are associated with hosts of L. distinguendus. Because these mites are not specific for S. granarius but also co-occur with other hosts, the mite chemicals have to be considered as general cues. In bioassays, synthetic dominicalure was attractive to naïve L. distinguendus, explaining the attraction of feces volatiles from R. dominica. Synthetic mite chemicals and sitophilate, the aggregation pheromone of S. granarius, had no effect on naïve parasitoids. It remains to be determined which innate chemical cues from feces of S. granarius are used by L. distinguendus. In contrast to our initial hypothesis, the generalist L. distinguendus is innately using specific cues for foraging. Two ideas are provided to explain this result.     

Allelopathic interactions among algae

Five species of green algae isolated from the Cleveland County, Oklahoma, area and a commercially obtained yellow-green alga were tested for allelopathic interactions. Axenic cultures ofPandorina morum Bory were stimulated in three sterile filtrates from nonaxenic cultures ofScenedesmus incrassatulus var.mononae G.M. Smith, with the oldest filtrate showing the least stimulation. By measuring the growth of axenic cultures inoculated into sterile filtrates of old cultures, instances of inhibition and stimulation were recorded in three screening experiments.Cosmarium vexatum West filtrate was inhibitory to the five other species, a characteristic probably important in its role of producing waterblooms in ponds and swamps. The allelopathic effects of filtrates of the five other species onBolrydium becherianum Vischer could be a key to its restriction to terrestrial sites. Instances of heat lability of the active substances were noted. These interactions, both allelopathic and stimulatory, probably play a significant role in the succession of algal blooms in nature.     

Production of onion fly attractants and ovipositional stimulants by bacterial isolates cultured on onion

Decomposing onions at certain microbial successional stages produce potent volatile attractants and ovipositional stimulants of the onion fly,Delia antiqua (Diptera: Anthomyiidae). A reproducible source of these compounds was obtained by culturingErwinia carotovora var.carotovora (EC) on sterile onion tissue. In laboratory choice tests, EC-inoculated onion was more attractive thanKlebsiella pneumoniae (KP) cultured on onion, EC cultured on potato (a nonhost of onion fly), or the chemical synthetic baits dipropyl disulfide and an aqueous solution of 2-phenylethanol and pentanoic acid. Onion flies were mildly attracted to potato after inoculation with EC, but females did not accept EC-inoculated potato for oviposition. This work emphasizes that sources of semiochemicals may need to be defined microbiologically as well as physically and chemically.Diptera: Anthomyiidae.Paper No. 12106 of the Michigan State University Agricultural Experiment Station.     

Volatiles from Duck Feces Attractive to Mexican Fruit Fly

Aqueous extracts of three types of bird feces were attractive to laboratory-colony Mexican fruit flies in laboratory bioassays. Extracts of black-bellied whistling duck feces were chosen for further analysis. Duck feces extract was attractive to both sugar-fed and sugar-starved flies but was more attractive to sugar-fed flies. Protein deprivation of flies had no effect on their response. Duck feces incubated in water for 24 hr were more attractive than those incubated for longer periods. Duck feces extract was attractive at pH 5–9, but was most attractive at pH 9, indicating that more than one class of chemicals contributed to attractiveness of the feces. Major components of headspace of duck feces extracts collected by solid-phase microextraction and identified by GC-MS, GC-FID, and GC-FTD included ethanol, propanol, phenol, ammonia, low-molecular weight amines, and pyrazines. In general, chemicals containing nitrogen proved most attractive to sugar-fed flies. However, dimethylamine and 1-pyrroline were nearly equally attractive to sugar-fed and sugar-deprived flies. Two chemicals without nitrogen were attractive to sugar-deprived flies; none were attractive to sugar-fed flies. A synthetic mixture was prepared containing ammonia, methylamine, dimethylamine, trimethylamine, 1-pyrroline, phenol, and 2-ethylhexanol in amounts found in odor of duck feces extract. The synthetic mixture was 96% as attractive as duck feces extract to sugar-fed flies and 80% as attractive to sugar-deprived flies.     

Characterization of a Behaviorally Active,Gender-Specific Volatile Compound from the Male Asparagus Fly <Emphasis Type="Italic">Plioreocepta poeciloptera</Emphasis>

Adult male asparagus flies exhibit typical calling behaviors (suggestive of pheromone production) during which they emit a single volatile compound that was identified as isopropyl (S)-5-hydroxyhexanoate. In laboratory bioassays, synthetic samples elicited an arrestant response in females, but did not appear to attract females. On the other hand, the synthetic material attracted conspecific males in olfactometer bioassays.     

Attraction of 3-Methyl-1-Butanol and Ammonia Identified from Enterobacter agglomerans to Anastrepha suspensa

Tests demonstrated that volatile chemicals emitted from Enterobacter agglomerans, a bacterium that has been isolated from adults as well as fruit infested with larvae of the Caribbean fruit fly, Anastrepha suspensa (Loew) and other pest fruit flies, are attractive to female A. suspensa in laboratory bioassays. 3-Methyl-1-butanol and ammonia were identified as the two primary volatile chemicals released from active cultures of E. agglomerans. No 3-methyl-1-butanol and little ammonia (16.0 g/hr) are released from sterile tryptic soy agar plates. E. agglomerans-inoculated tryptic soy agar plates, however, released an average of 1.5 ± 0.53 g/hr 3-methyl-1-butanol and 332.9 ± 239.16 g/hr ammonia after 24 hr of growth. 3-Methyl-1-butanol lures were formulated in a membrane-based system to provide a constant release rate of synthetic chemical. Release rates ranged from 0.046 ± 0.007 to 12.16 ± 2.76 g/hr. In laboratory tests, equal numbers of females were captured in response to ammonium carbonate lures that released ammonia at the rate of 100 g/hr and to 3-methyl-1-butanol lures that released 12.16 ± 2.756 g/hr of synthetic material. The combination of the two lures was more attractive than ammonia alone. Availability of lures formulated for a range of 3-methyl-1-butanol release rates will facilitate field tests of this putative microbial attractant and may lead to a better understanding of the role of bacteria in the ecology of pest fruit flies.     

Differences among sibling speciesRhagoletis mendax andR. pomonella (diptera: ephritidae) in their antennal sensitivity to host fruit compounds

Prior electrophoretic and morphological studies have identified two closely related, economically important tephritid flies,R. mendax (Curran) andR. pomonella (Walsh), which infest the fruits of ericaceous and rosaceous plants, respectively. Further studies also have shown consistent differences among these species in their ovipositional preferences for apples and highbush blueberries and have determined that their ovipositional behavior is elicited by extracts obtained from these fruits. In this paper we report the results of an experiment that tested whether these species show distinct electroantennogram (EAG) responses to a large array of compounds present in gas chromatograph-fractionated pentane extracts of apples and highbush blueberries.R. mendax andR. pomonella flies were found to have significant differences in their antennal sensitivity to 11 blueberry and nine apple extract peaks, which correspond to 24.4% of all blueberry and 25.0% of all apple peaks that elicited a measurable EAG response from either species. Interspecific differences in peripheral sensitivity were more pronounced for blueberry than apple extract;R. pomonella flies were most sensitive to blueberry compounds with low retention times, whereasR. mendax flies responded to blueberry compounds with a broader range of retention times. Both species were most sensitive to apple peaks with high retention times. The retention times of most apple and blueberry peaks that elicited EAG responses fromR. mendax andR. pomonella flies were different from the retention times of seven attractant fruit esters that were previously identified by Fein et al. (1982). The identification of these unknown apple and blueberry compounds could lead to the discovery of new chemical cues that mediate the host-plant preferences of these sibling species.     

Chemicals Attractive to Mexican Fruit Fly from Klebsiella pneumoniae and Citrobacter freundii. Cultures Sampled by Solid-Phase Microextraction

Headspace above tryptic soy broth culture filtrates of Klebsiella pneumoniae contained greater amounts of ammonia, methylamine, 3-methylbutanamine, 1-pyrroline, 2,3,4,5-tetrahydropyridine, and two pyrazines than were found above tryptic soy broth. It also contained chemicals not found above tryptic soy broth, including dimethyldisulfide and several alcohols and ketones. Headspace above tryptic soy broth culture filtrates of Citrobacter freundii contained greater amounts of ammonia, 1-pyrroline, and several pyrazines than were found above tryptic soy broth. It also contained chemicals not found above tryptic soy broth including dimethyldisulfide, some of the same alcohols as above K. pneumoniae filtrates, a different ketone, and phenol. Additional chemicals were detected above filtrates that were saturated with sodium chloride or had their pH adjusted up or down. In laboratory bioassays with protein-starved, sugar-fed Mexican fruit flies, chemicals that did not contain protonizable nitrogen were not attractive. All chemicals containing protonizable nitrogen, except 2-methylpyrazine, were attractive. Synthetic mixtures of ammonia, trimethylamine, 1-pyrroline, 3-methylbutanamine, pyrazine, 2,3,4,5-tetrahydropyridine, 2,5-dimethylpyrazine, and trimethylpyrazine in concentrations similar to those in filtrates of the two bacteria were 73–87% as attractive as bacterial filtrates.