Multicomponent alarm pheromones in the mandibular glands of major workers of the African weaver ant, Oecophylla longinoda

ABSTRACT. The complex system of alarm communication in Oecophylla longinoda is described. The mandibular gland secretion of major workers releases in other major workers a complex pattern of behaviour, which includes components of alerting, attraction and biting. The main constituents of the secretion are hexanal and 1-hexanol, which release alerting and attraction respectively. Of the thirty or so trace components, at least two, 2-butyl-2-octenal and 3-undecanone, act as markers for attack. In combination, these components produce a sequential message in space and time, dependent on their relative proportions, volatilities and threshold concentrations for behavioural reponses.     

The volatile components of the mandibular gland secretion of workers of the ants Myrmica lobicornis and Myrmica sulcinodis

The mandibular glands of workers of M. lobicornis produce a mixture of 3-alkanones and 3-alkanols in the C₆–C₁₀ carbon chain length range, in addition to nanogram amounts of acetone, ethanal and 2-methylpropanal. Ethological studies have shown that the three major constituents, 3-octanol, 3-octanone and 3-decanol, are also the major pheromonally active components. When presented together they stimulate an alarm response in the workers similar to that induced by a worker's crushed head.In M. sulcinodis the mandibular gland secretion is composed of a mixture of the same ketones and alcohols, but in different proportions.     

Phéromones agrégeant les ouvrières de Myrmica rubra

The workers of Myrmica rubra aggregate around a source of one of their secretions, which can be called ‘alarm pheromone’, and also around workers of Lasius flavus. The mechanism of these aggregations differ.Both L. flavus workers and a solution in liquid paraffin of 3-octanol, one of the mandibular gland compounds, act as an arrestant for the workers of M. rubra. Both Dufour's gland secretion and a source of 3-octanone, the major compound of the mandibular gland secretion, are true attractants.The poison gland secretion, a mixture of 3-octanone and 3-octanol in liquid paraffin and a solution in liquid paraffin of 3-nonanone, a minor mandibular gland compound, all induce klinokinesis. The secretion of the mandibular glands and the secretion of the venom apparatus both cause positive klinokinesis and taxis. These locomotory reactions increase the probability that an object, marked by nest mates with these secretions, will be detected by several workers.When presented alone, 3-octanone is the only attractive compound in the mandibular gland secretion. However, a mixture of 3-octanone and 3-octanol (15 per cent of 3-octanol in the vapour phase) is detected more easily by the ants. The diffusion coefficients of the two compounds are different, and a mixture of these substances creates not only a quantitative but also a qualitative odour gradient. This may explain the synergy of the mixture.     

Electroantennogram and behavioral responses of workers of the stingless bee Oxytrigona mediorufa to mandibular gland volatiles

Gas chromatography-mass spectrometric analysis showed that the mandibular gland secretion of workers of the stingless bee Oxytrigona mediorufa (Cockerell) (Hymenoptera: Apidae) is composed of 3-hepten-2,5-dione, 3-nonene-2,5-dione, 2-heptanone, hydrocarbons, tetradecyl, and hexadecyl acetates and minor compounds related to these. Behavioral and electroantennographic tests were carried out on conspecific workers to observe the effect of worker mandibular gland secretion, individual compounds (2-heptanone, 2-nonanone, and tetradecyl acetate), and a blend of these three. Bioassays showed that biting responses of O. mediorufa to the odor of the mandibular gland secretion were similar to that of 2-heptanone and to the three-component blend. Oxytrigona mediorufa workers exhibited antennal olfactory responses to the gland extract, 2-heptanone, and the synthetic mixture. The function of the mandibular gland secretion and 2-heptanone is discussed in terms of an alarm pheromone and defensive substances.     

The physicochemical transmission of two components of a multiple chemical signal in the African weaver ant, (Oecophylla longinoda)

The behavioural thresholds of the two principal components of the mandibular gland secretion in major workers of the weaver ant Oecophylla longinoda, hexanal and 1-hexanol, were found to be 1.99×10¹⁰ and 3.35×10¹¹ molecules cm^?3 respectively. The mandibular gland contains an average of 2.896×10¹⁵ molecules of hexanal, and 1.274×10¹⁶ molecules of 1-hexanol. From these measurements, molecular diffusion theory predicts behavioural active spaces for 1-hexanol (an attractant) which agree with those observed in response to the whole secretion, but there is no such agreemnet for haxanal (a releaser of behaviour with no directionial compnent). The effect of microturbulence on the transmission of releaser and chemotactic stimuli is discussed.     

The ontogenetic pattern of mandibular gland components in queenless worker bees (Apis mellifera capensis Esch.)

The quantity and composition of the six major mandibular gland components of young queenless workers of the Cape honeybee (Apis mellifera capensis) were determined. The total amount of the six components increased with age. The relative quantities in the mandibular gland secretion of queenless caged workers were found to change rapidly during the first 4 days after emergence and to become dominated by the queen substance, 9-keto-2(E)-decenoic acid. Also the relative amounts of 9-hydroxy-decenoic acid, a precursor of the queen substance, showed an increase of an order of magnitude within the first 4 days of imaginal life. The relative amounts of the aromatic compounds typical to the queen pheromone remained similar in this developmental time window. The increase of queenlike compounds is particularly strong between days two and three after emergence. These queen-like pheromones play a major role in the development of reproductive hierarchies among workers under queenless conditions. This may be an important factor in the socio-parasitic pathway of A. m. capensis.     

The mandibular gland secretion of the ant, Myrmica scabrinodis

ABSTRACT. The volatile secretion of the mandibular gland of the common elbowed red ant, Myrmica scabrinodis Nyl., is shown to consist of ethanal, ethanol, acetone, 3-hexanone, 3-hexanol, 3-heptanone, 3-heptanol, 3-octanone, 3-octanol, 6-methyl-3-octanone, 6-methyl-3-octanol, 3-nonanone, 3-nonanol, 3-decanone and 3-undecanone. The electroantennographic response to the major components was recorded and compared with some related compounds. Behavioural tests were carried out on the major constituents, showing that 3-octanol is an attractant for workers, that 3-octanone increases the effect of 3-octanol, and 3-nonanone augments the linear speed of the ants.     

Effect of gland extracts on digging and residing preferences of red imported fire ant workers (Hymenoptera: Formicidae)

There is evidence that ant‐derived chemical stimuli are involved in regulating the digging behavior in Solenopsis invicta Buren. However, the source gland(s) and chemistry of such stimuli have never been revealed. In this study, extracts of mandibular, Dufour's, postpharyngeal, and poison glands were evaluated for their effect on ant digging and residing preferences of S. invicta workers from three colonies. In the intracolonial bioassays, workers showed significant digging preferences to mandibular gland extracts in 2 of 3 colonies and significant residing preferences in 1 of 3 colonies; significant digging preferences to Dufour's gland extracts in 1 of 3 colonies and significant residing preferences in 2 of 3 colonies. No digging and residing preferences were found for postpharyngeal and poison gland extracts. In intercolonial bioassays, significant digging and residing preferences were found for mandibular gland extracts in 3 of 6 colony combinations. Significant digging preferences to Dufour's gland extracts were found in 4 of 6 colony combinations and significant residing preferences in all 6 colony combinations. For postpharyngeal gland extracts, significant digging preferences were found only in 1 of 6 colonial combinations and no significant residing preferences were found. For poison gland extracts, no significant digging preferences were found; significant residing preferences were found in 1 of 6 colony combinations. However, a significant residing deterrence (negative residing preference index) was found for 2 of 6 colony combinations. Statistical analyses using data pooled from all colonies showed that mandibular and Dufour's gland extracts caused significant digging and residing preferences in both intracolonial and intercolonial bioassays but not postpharyngeal and poison gland extracts. By analyzing the data pooled from the same three colonies used for gland extract bioassays, it was found that, in no cases, workers showed significant digging and residing preferences to 2‐ethyl‐3,6‐dimethylpyrazine, an alarm pheromone component from mandibular gland.     

蜜蜂上颚腺及其分泌物研究进展

上颚腺是蜜蜂重要的外分泌腺体,其分泌物是维系蜂群社会性结构的重要物质。蜂王和工蜂上颚腺分泌物合成均以硬脂酸为合成前体,但在脂肪酸的β-氧化过程中表现出级型差异性,导致分泌物组分比例不同。蜂王上颚腺分泌物以9-羰基-2癸烯酸(9-ODA)为主,有吸引工蜂和雄蜂、抑制工蜂卵巢发育等作用;工蜂上颚腺分泌物以10-羟基-2癸烯酸(10-HDA)和10-羟基癸酸(10-HDAA)为主,是蜂王浆的重要组成部分。同时,这种具备典型级型差异的分泌物组成又具有级型间可塑性,在不同蜂种间也存在区别。近年来在转录水平和蛋白水平的一些研究进一步揭示了级型间差异的分子基础。针对蜜蜂上颚腺及其分泌物的研究在蜜蜂生物学、行为学和蜂产品质量控制等方面具有重要的意义。本文通过总结国内外相关研究进展,旨在为上颚腺分泌物的作用机制、生物合成机制等领域的进一步深入研究提供借鉴。     

Exocrine chemistry of the myrmicine ant Zacryptocerus pusillus (Hymenoptera: Formicidae)

Minor workers of the ant Zacryptocerus pusillus have unusual exocrine secretions in both their mandibular and Dufour glands. The mandibular glands contain a 3:1 mixture of 4-heptanone and 4-heptanol, a mixture found only in the related species Z. varians. The Dufour gland contains a mixture of 13 aldehydes from C9 to C18, not previously encountered in ant secretions. The venom glands gave variable results with only nonanal present consistently.     

Mandibular gland chemistry of four Caribbean species of Camponotus (Hymenoptera: Formicidae)

The volatile components of whole-body extracts of males, females and workers were analyzed in four species of Neotropical ants in the formicine genus, Camponotus. The species, C. kaura, C. sexguttatus, C. ramulorum and C. planatus, represent three different subgenera. Volatile mandibular gland components were found only in male extracts in three of the species. In C. ramulorum, volatile components were found in male and female reproductives and workers. 3,4-Dihydro-3,5-dihydroxy-6-methylpyran-4-one and octanic acid were found in different sets of three of the species. Methyl 6-methyl salicylate was found in two species and the isocoumarin, mellein, was found in a third species. The significance of the mandibular gland secretion for formicid systematics is discussed.     

Functional morphology of the mandibular gland of queens of the ant Monomorium pharaonis (L.)

In this study, the functional morphology of the mandibular glands of pharaoh ant queens is investigated through histological and behavioural observations. The mandibular glands of queens of different age stages and mating status were examined at the light microscopical as well as the ultrastructural level. The results clearly show a high activity of the gland at the time of hatching, followed by a rapid degeneration independent of the queens' mating status. We therefore hypothesize the glandular secretion serves to stimulate workers to remove the queens' pupal skin. Experiments to compare hatching ability of isolated pupae and pupae assisted by workers confirmed the necessity of worker assistance, which is even more crucial to allow proper wing formation in queens and males. Together with the histological data, this suggests that the mandibular gland may indeed play a role in the social facilitation of hatching, although bioassays are required to give decisive answers about the mandibular gland's function.     

Mandibular gland secretions of workers of Myrmica rugulosa and M.schencki: comparison with four other Myrmica species

ABSTRACT. The mandibular glands of the two species of ant, Myrmica schencki Em. and Myrmica rugulosa Nyl., contain mixtures of similar compounds, but in different proportions. M.rugulosa produces 3-pentanol, 3-hexanone, 3-hexanol, 3-heptanone, 3-heptanol, 3-octanone (by far the most abundant component), 3-octanol, 3-nonanol, 3-decanone and 6-methyl-3-octanone, in addition to small amounts of ethanal, acetone and methylpropanal. M.schencki produces most of these (though much less 3-octanone and much more 3-octanol), but also produces significant amounts of 3-nonanol, 3-decanol and 6-methyl-3-octanol, while producing no detectable 3-pentanol or 3-hexanone. The mandibular gland secretions of these two species attract the workers, increase their linear speed, and reduce their sinuosity of movement. In M.schencki these behavioural activities are caused by 3-octanol and 3-octanone, the effect of a synthetic mixture of the two being exactly like that of an isolated mandibular gland; the two compounds act together to cause attraction and increase linear speed, and in synergy to reduce the workers' sinuosity of movement. In M.rugulosa , 3-octanol, 3-octanone and 6-methyl-3-octanone are the major active constituents. 3-Octanone attracts the workers, its effect being enhanced by 3-octanol; it also increases the ants' linear speed, this effect being moderated slightly by the 3-octanol. Presented together these two substances act synergistically to decrease the workers' sinuosity of movement, and reproduce exactly the overall behavioural activity of an isolated mandibular gland. The chemical and behavioural results are combined with those previously reported to explain the responses of M.rubra, M.ruginodis, M.rugulosa, M.sabuleti, M.schencki and M.scabrinodis workers to isolated mandibular glands of these species.     

The labial gland in the termiteSchedorhinotermes lamanianus (Isoptera: Rhinotermitidae): Morphology and function during communal food exploitation

Summary InSchedorhinotermes lamanianus, size and cellular differentiation of the labial gland exhibit a caste-specific polymorphism. The acini of workers are composed of three distinct types of secretory cells, one of which is absent in soldiers and alates. The labial gland of workers releases a chemical signal for intraspecific communication. During the communal exploitation of a food source, labial gland secretion makes workers aggregate at gnawing sites where this signal is deposited. A newly developed bioassay is demonstrated.     

Exocrine glands of the ant Myrmoteras iriodum

This paper describes the morphological characteristics of nine major exocrine glands in workers of the formicine ant Myrmoteras iriodum. The elongate mandibles reveal along their entire length a conspicuous intramandibular gland, which contains both class‐1 and class‐3 secretory cells. The secretory cells of the mandibular glands show a peculiar appearance, with a branched end apparatus, which is unusual for ants. The other major glands (pro‐ and postpharyngeal gland, infrabuccal cavity gland, labial gland, metapleural gland, venom gland and Dufour gland) show common features for formicine ants. The precise function of the glands could not yet be experimentally demonstrated, and to clarify this will depend on the availability of live material of these enigmatic ants in future.     

The antimicrobial activity of the mandibular gland secretion of a formicine ant, Calomyrmex sp. (Hymenoptera: Formicidae)

The mandibular gland secretion from mature workers of the formicine ant Calomyrmex sp. exhibits strong antimicrobial activity when tested against selected soil microorganisms. The activity against bacteria is both inhibitory and bacteriocidal while that against fungi is inhibitory. The white secretion from young workers appears to be much weaker in its antibiotic effects.     

Identification of a component of the trail pheromone of the ant Pheidole pallidula (Hymenoptera: Formicidae)

ABSTRACT The poison gland of minor workers of P.pallidula. Nyl. contains 3-ethyl-2,5-dimethylpyrazine which induces trail-following in other workers, but does not account for the full trail-following effect of one worker's poison gland. No pyrazines were detected in major workers and their glands do not contain the pheromone.     

Analysis of the enzymatic formation of citral in the glands of sweet basil

Basil glands of the Sweet Dani cultivar contain high levels of citral, a mixture of geranial and its cis-isomer neral, as well as low levels of geraniol and nerol. We have previously reported the identification of a cDNA from Sweet Dani that encodes an enzyme responsible for the formation of geraniol from geranyl diphosphate in the glands, and that these glands cannot synthesize nerol directly from geranyl diphosphate. Here, we report the identification of two basil cDNAs encoding NADP+-dependent dehydrogenases that can use geraniol as the substrate. One cDNA, designated CAD1, represents a gene whose expression is highly specific to gland cells of all three basil cultivars examined, regardless of their citral content, and encodes an enzyme with high sequence similarity to known cinnamyl alcohol dehydrogenases (CADs). The enzyme encoded by CAD1 reversibly oxidizes geraniol to produce geranial (which reversibly isomerizes to neral via keto-enol tautomerization) at half the efficiency compared with its activity with cinnamyl alcohol. CAD1 does not use nerol and neral as substrates. A second cDNA, designated GEDH1, encodes an enzyme with sequence similarity to CAD1 that is capable of reversibly oxidizing geraniol and nerol in equal efficiency, and prolonged incubation of geraniol with GEDH1 in vitro produces not only geranial and neral, but also nerol. GEDH1 is also active, although at a lower efficiency, with cinnamyl alcohol. However, GEDH1 is expressed at low levels in glands of all cultivars compared with its expression in leaves. These and additional data presented indicate that basil glands may contain additional dehydrogenases capable of oxidizing geraniol.     

Behavioral and olfactory antennal responses of Solenopsis geminata (Fabricius) (Hymenoptera: Formicidae) workers to their dufour gland secretion

Behavioral and electrophysiological tests were performed to evaluate the responses of workers of the ant Solenopsis geminata (Fabricius) from different size categories to Dufour gland extracts. Morphometric measures based in head widths across eyes were used to determine worker sizes. Trail following response of different worker sizes to Dufour gland extract from workers of different sizes was assessed. For each worker size category olfactory responses to Dufour gland extracts were determined using electroantennography (EAG). Gas chromatography and mass spectrometry (GC-MS) were used to determine the chromatographic profile of Dufour gland secretion for each worker size. Morphometric measures permitted to classify the workers of S. geminata as large, medium and small workers. Medium S. geminata workers displayed a significantly higher behavioral response to Dufour gland extracts produced by medium size workers. Similarly, medium workers showed a significantly higher EAG response to Dufour gland extracts produced by medium sized workers. Chromatographic profile of Dufour gland secretions produced by workers showed that each size category exhibited a characteristic profile of the three main components considered as potential trail pheromone constituents. This work showed that medium workers of S. geminata exhibited a high trail-following behavior as well as a high antennal response to Dufour gland secretion. This and their relative abundance in field foraging areas, suggest that medium-sized workers are specialized in foraging activities.     

Mechanism of secretion of the contents of the mandibular glands of Plodia interpunctella larvae

ABSTRACT. Larvae of Plodia interpunctella deposit droplets of mandibular gland secretion onto silk filaments connecting particles within the substrate they inhabit, and on to the substrate itself. Active participation of the mouthparts is necessary for both the formation and deposition of these droplets along the filaments and occurs as the silk is spun. Deposition of secretion directly on the substratum does not involve the mouthparts and appears to result from contact between the lower surface of the head and the substratum. Larvae from which the mandibular glands have been removed by excision neither produce droplets on silk nor deposit them on the substratum. Those which are incapable of spinning silk due to cauterization of the spinneret are still able to deposit secretion on the substratum, however. Deposition of secretion on silk webbing does not occur when the setae which regulate formation of droplets on filaments are removed by shaving. Droplets are not characteristic of a particular larval instar but are produced by larvae of all instars as a normal function which does not depend upon intraspecific encounters. Four other Lepidopteran pests, Ephestia elutella, Ephestia cautella, Anagasta kuehniella and Antigasta catolaunalis , were also found to produce droplets of mandibular gland secretion in a similar manner to larvae of P. interpunctella.