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Sexualpheromone sind im gesamten Tierreich weit verbreitet. Vor allem Insekten nutzen diese Art der Partnerfindung. Im Vergleich zu vielen Insektenarten, bei welchen die Art der pheromonbildenden Drüsenzellen, ihre Lage im Körper sowie die chemische Zusammensetzung ihrer Pheromone identifiziert wurde (s. Review von Blomquist & Vogt 2003) ist bei Schnellkäfern (Elateridae) noch sehr wenig bekannt. Die verschiedenen mitteleuropäischen Schnellkäferarten lassen sich anhand ihrer Pheromonspeicherung in zwei unterschiedliche Gruppen unterteilen. Die Weibchen der Unterfamilie Elaterinae, besitzen im letzten von außen sichtbaren Abdominalsegment ein paariges Drüsenreservoir in welchem das produzierte Sexualpheromon gespeichert wird. Im Gegensatz dazu fehlt bei den Weibchen der übrigen Unterfamilien ein solches Pheromonreservoir und es ist unklar, wo bei diesen Arten die Pheromone produziert werden. Beobachtungen von Ivastschenko & Adamenko (1980) deuten darauf hin, dass sich bei diesen Arten die pheromonproduzierenden Zellen im Bereich der Intersegmentalmembran zwischen dem Abdominalsegment VIII und IX befinden. Um zu klären, ob die Pheromondrüsen tatsächlich in diesem Bereich zu finden sind, wurden im Rahmen der vorliegenden Arbeit die Intersegmentalmembranen bei Weibchen und Männchen von Athous haemorrhoidalis und A. subfuscus auf mögliche Drüsenzellvorkommen untersucht.
Die Schlupfwespe Pimpla turionellae parasitiert versteckte Lepidopterenpuppen und orientiert sie sich bei der Wirtssuche multisensorisch mittels visueller Reize und aktiver Vibrationsortung mit selbst produzierten Schwingungen (Vibrational-Sounding). Die Studie untersucht, inwieweit die Wespen bei Änderung der Umgebungstemperaturen von 8-26°C (1) zwischen der temperatur-sensitiven vibratorischen und der -insensitiven visuellen Orientierung wechseln und (2) gegebenenfalls selbst die Körpertemperatur regulieren können, um die sehr präzise Vibrationsortung bei niedrigen Temperaturen aufrechtzuerhalten. Messungen mit Infrarot-Thermographie zeigen, dass suchende Wespen leicht erhöhte Körpertemperaturen während der vibratorischen Wirtssuche aufweisen, welche auf metabolische Wärmeproduktion zurückzuführen sind. Wahlexperimente unter kontrollierten Temperaturen zeigen zudem, dass die Nutzung der temperatur-sensitiven vibratorischen Reize bei pessimalen Temperaturen abnimmt und die Wespen auf fast ausschließliche visuelle Orientierung wechseln. Folglich wird die Relevanz einzelner Reize bei der multisensorischen Orientierung direkt vom Faktor Temperatur beeinflusst. Solange ein zuverlässiger Reiz vorhanden ist, nimmt dabei auch die Präzision der Lokalisation insgesamt nicht ab.
Erkenntnisse über das Wirt-Parasit-Verhältnis zwischen Hopfen Humulus lupulus L. und Hopfenblattlaus Phorodon humuli (Schrank) sind von entscheidender Bedeutung für die Selektion blattlausresistenter Sorten. Die Besiedelung des Wirts durch Blattläuse wird durch in den Pflanzen befindliche Substanzen erheblich beeinflusst (Nault & Styer 1972; Leath & al. 1974; Klingauf & al. 1978; Kendall & al. 1980; Singh 1980; Powell & al. 1999; Campo & al. 2003). Die in dieser Hinsicht wichtigsten Bestandteile des Hopfens sind die Bittersäuren, welche für den Parasit die Funktion von Signalstoffen haben können. In frischem Hopfen liegen sie überwiegend in Form der alpha-Bittersäuren (Humulon, co-Humulon, ad-Humulon) und der beta-Bittersäuren (Lupulon, co-Lupulon, ad-Lupulon) vor (Backleh 2001). Die Gehalte an Bitterstoffen im Pflanzengewebe des Hopfens schwanken stark im Lauf der Vegetationsperiode. Da ein Insekt sich an quantitative wie auch qualitative Unterschiede im Nährstoffangebot anzupassen vermag, sollten sich Änderungen im Wirtswahlverhalten bei der Nahrungsaufnahme von Blattläusen durch die saisonbedingten Stoffwechselschwankungen der Pflanzen ergeben. Um diese Veränderungen im Verhalten der Aphiden zu erfassen, war es notwendig, die Verhaltensbeobachtungen im Verlauf der Vegetationsperiode mehrfach zu wiederholen. Die Methode der Wahl zur Untersuchung quantitativer Unterschiede im Verhalten bei der Nahrungsaufnahme von Aphiden ist der Electrical Penetration Graph (EPG) (McLean & Kinsey 1964; Spiller 1988; Prado & Tjallingii 1994; Calatayud & al. 2001; 1978, 1988, 2006). In der vorliegenden Studie wurde diese Methode mit gleichzeitigen HPLC - Analysen der Bittersäurezusammensetzung des Gewebes relevanter Organe (Blätter und Zapfen) von Hopfen im Verlauf der Vegetationsperiode kombiniert.
Thysanopteren wehren sich gegen Prädatoren und Parasiten, indem sie einen Analtropfen mit einem Wehrsekret produzieren. Dieser Tropfen wird in einigen Fällen hoch gehalten, meist jedoch auf den Angreifer aufgetupft (Lewis 1973). Über die Wehrsekrete von Thripsen der Unterordnung Terebrantia ist bisher wenig bekannt. Von der einzigen chemisch genauer untersuchten Art Frankliniella occidentalis weiß man, dass deren Larven hauptsächlich mit einem Gemisch aus Decyl- und Dodecylacetat arbeiten (Teerling & al. 1993a, MacDonald & al. 2003). Diese Substanzen sind nur in beschränktem Umfange als Allomon wirksam. Hauptsächlich wirken sie als Alarmpheromon (Bakker & Sabelis 1987, 1989), indem sie die Aufmerksamkeit der Larven erhöhen. Nachteilig ist die relativ hohe Flüchtigkeit dieser Acetate. Sie können dadurch zusätzlich als Kairomon wirken und Prädatoren anlocken (Teerling & al. 1993b). Ein weiteres Problem ist, dass solche Acetate schnell verdunsten und nur für kurze Zeit auf der Oberfläche der Sinnesorgane potenzieller Angreifer verbleiben. Nicht untersucht ist, ob sie eventuell durch Umesterung, durch Freisetzung von Essigsäure oder durch andere Mechanismen schädigend wirken. Denkbar wäre hier auch eine ungünstige Beeinflussung der Epicuticula der Sensillen oder der im Sensilleninneren enthaltenen Pheromon-Bindungsproteine. Bei den Vertretern der Unterordnung Tubulifera wird die Handhabung des Wehrsekrettropfens besonders unterstützt durch einen Tubus, der in Extremfällen (beispielsweise bei Leeuwenia gladiatrix) fast die Hälfte der Körperlänge ausmachen kann. Mit diesem Tubus kann der Wehrsekrettropfen sehr geschickt über den Kopf hinweg gerichtet, auf sichere Entfernung, appliziert werden. Eigene Beobachtungen und die anderer Autoren (beispielsweise Howard & al. 1987) zeigen, dass bei allen bisher untersuchten Tubulifera der Sekrettropfen, wenn er nicht genutzt wurde, sorgfältig wieder „eingezogen“ wird.
Das Wirtswahlverhalten von Blattläusen wird neben primären Nahrungskomponenten auch durch sekundäre Pflanzeninhaltsstoffe gesteuert (Klingauf & al. 1978; Singh 1980; Powell & al. 1999; Campo & al. 2003). Häufig entziehen sich diese jedoch einer gezielten Untersuchung ihrer Wirkung auf die Aphiden, da sie als Substanzgemisch vorliegen. Beziehungen zwischen Substanzkonzentration und Verhaltensmustern aus Electrical Penetration Graphs (EPG) sind nur mit mäßigem Erfolg über statistische Verfahren darzustellen. Eine Versuchsanordnung zur gezielten Prüfung verhaltensbeeinflussender Substanzen mittels EPG wäre in diesem Falle hilfreich. Ein geeignetes einfaches Testsystem müsste mindestens folgende Komponenten beinhalten: – eine von den Aphiden akzeptierte Oberfläche, welche die Kutikula / Epidermis eines natürlichen Systems nachbildet – diese Oberfläche sollte evtl. durch den Versuchsansteller manipulierbar sein – eine dem Parenchym entsprechende Schicht, in welche die Testsubstanzen einzubringen wären – eine flüssige Phase, die dem Phloem entspricht und ebenfalls in ihrer Zusammensetzung variiert werden kann.
Cantharidin, which is mainly found in blister beetles (Coleoptera: Meloidae), is one of the most intensively studied natural products of insect (Dettner, 1997; McCormick & Carrel, 1987). The involvement of cantharidin in courtship behaviour has been already confirmed for certain canthariphilous insects (Eisner & al. 1996a,b; Frenzel & Dettner 1994; Frenzel & al. 1992; Schütz & Dettner, 1992; Hemp & al. 1999). The function and intrinsinc role of cantharidin in the courtship behaviour of Meloids has been never fully established. McCormick & Carrel (1987) only suggested that cantharidin might be used by female meloids when selecting a mate at close range. Pinto (1974, 1975) was, in fact, the first to consider male cuticular pores as being involved in the courtship behaviour of species from the genus Linsleya and Tegrodera (Meloidae). Based on morphology and chemical analyses of Cyaneolytta sp. (Coleoptera: Meloidae), we have hereby provided some further evidences that cantharidin may act as an infochemical in courtship behaviour of meloid beetles.
Gögler, Julia ; Stökl, Johannes ; Sramkova, Anna ; Twele, Robert ; Francke, Wittko ; Cortis, Pierluigi ; Scrugli, Antonio ; Giotta, Cesario ; Piccitto, Marcello ; Ayasse, Manfred
Sexual deception of male bees is one of the most remarkable mechanisms of pollination (Ackermann 1986, Proctor & al. 1996). Flowers of the orchid genus Ophrys mimic females of their pollinator species, usually bees and wasps, to attract males, which try to copulate with the flowers. During this so-called “pseudocopulation” the male removes the pollinia and transfers them to another flower to ensure pollination. Apart from visual and tactile cues, floral scent was shown to be most important for eliciting mating behaviour in males (Kullenberg 1961, Schiestl & al. 1999, Ayasse & al. 2003). Pollination in Ophrys is highly specific and usually each Ophrys species attracts only one pollinator species (Paulus & Gack 1990). The high degree of specialization provides the means of reproductive isolation between the intercrossable Ophrys-species (Ehrendorfer 1980). The complex odour-bouquets released by the flowers are species-specific and often consist of more than 100 different chemical compounds (Borg-Karlson & al. 1985, Ayasse 2006). Speciation in Ophrys-orchids may be brought about by changes in the pollinator attracting floral scent. The attraction of a new pollinator may act as a pre-zygotic isolation barrier (Stebbins 1970, Paulus & Gack 1990, Soliva & al. 2001). We investigated three sympatrically occuring Ophrys-species on Sardinia. O. chestermanii and O. normanii are endemic and are both pollinated by males of the bumblebee B. vestalis. O. tenthredinifera is pollinated by Eucera nigrilabris. There are different opinions concerning the taxonomic status of O. normanii. It has been described as an actual hybrid between O. chestermanii and O. tenthredinifera (Wood 1983). Paulus & Gack (1995) suggested that it is an own species, that either has developed from a hybrid between O. chestermanii and O. normanii or that has evolved by radiation from O. tenthredinifera. By conducting behavioural-tests with B. vestalis males, performing gas chromatographic analyses and electrophysiological studies we wanted to identify pollinator attracting scent and to clarify the taxonomic status of O. normanii.
The orchid genus Epipactis is represented by 25 species in Europe (Richards 1982). Epipactis helleborine (L.) Crantz is the most common and widely distributed species of the genus (Wiefelspütz 1970), and is a prime example for wasp-flowers, because it is mainly pollinated by social wasps (Hymenoptera: Vespidae), like Vespula vulgaris and V. germanica (Müller 1873). Darwin (1888) already noticed that E. helleborine is almost exclusively ignored by bees and bumblebees, an observation that was confirmed in recent investigations (Keppert 2001). The flowers of E. helleborine show morphological, physiological and phenological adaptations to the visit and the pollination by Vespidae (Keppert 2001). They possess a reddish-brown or dirty purple coloration of the inflorescence (Keppert 2001), have relatively small, mostly bulbous blossoms with a broad entrance and bulbous widened, nectar-rich juice holders (Müller 1873, 1881; Schremmer 1962). Although there is much reported about wasp-pollinated flowers there is little known about the signals that are responsible for the attraction of wasps. Wiefelspütz (1970) proclaimed the statement that only the visual stimulus is responsible for the wasp attraction. Recently studies, however, assumed that odour is involved in the wasp attraction (Keppert 2001). Hölzler (2003) showed that the main attraction of the wasp-flower Epipactis for pollinators is its olfactory stimulus. It remains an unanswered question why E. helleborine flowers almost exclusively attracts social wasps, as opposed to bees and bumblebees. In this study we analysed the role of floral volatiles which are responsible for the specific attraction of social wasps. We supposed a mimicry-system in E. helleborine for the specific attraction of pollinators for the following reasons. So-called “green leaf volatiles” (GLVs) are emitted by plants while herbivorous insects, for example caterpillars, feed on them. GLVs thereby attract predators or parasitoids of the herbivorous insects (Dicke & Sabelis 1988; Turlings & al. 1990, 1995; Dicke & Vet 1999). Among the GLVs so far identified in former studies there are aldehydes, compounds that were also found in flower extracts of E. helleborine (Hölzler 2003). Therefore, we postulated that E. helleborine flowers produce GLVs in order to attract prey hunting social wasps for pollination. We performed bioassays and analysed flower odour gained to headspace-sampling using gas chromatography (GC), mass spectrometry (GC-MS) and gas chromatography coupled with electrophysiological analysis (GC-EAD) to investigate the hypothesis that E. helleborine flowers mimic “green leaf volatiles” (GLVs) to attract their pollinators.
Die Glucosinolate (GS) sind charakteristische sekundäre Pflanzeninhaltsstoffe, vorkommend in der Gruppe der Brassicaceae und anderen Familien der Ordnung Brassicales (Halkier & Gershenzon 2006). Bisher sind mehr als 120 verschiedene GS beschrieben, welche eine gemeinsame Grundstruktur mit variablem Seitenkettenrest kennzeichnet (Fahey et al. 2001). Je nach chemischer Natur der Seitenkette werden die GS in aliphatische, aromatische und Indolyl-GS unterteilt. Alle GS-enthaltenden Pflanzen besitzen zusätzlich räumlich getrennt von den GS hydrolysierende Enzyme, so genannte Myrosinasen. Erst nach Zellbeschädigung kommen die beiden Komponenten in Kontakt zueinander und weitere biologisch aktive Verbindungen wie z. B. Isothiocyanate und Nitrile werden freigesetzt (Rask et al. 2000). Das GS-Myrosinase-System ist ein effektives Abwehrsystem insbesondere gegenüber generalistischen Insekten, Pathogenen und Bakterien, allerdings dienen vielen spezialisierten Insekten diese Stoffe zur Wirtspflanzenfindung und -akzeptanz (Renwick 2002, Halkier & Gershenzon 2006). Die Modellpflanze Arabidopsis thaliana L. enthält als Vertreter der Brassicaceae GS als Fraßabwehrstoffe. In A. thaliana als auch in Brassica ist das aliphatische GS-Muster sehr variabel, wohingegen die Indolyl-GS weit verbreitet sind (Kliebenstein 2001, Li & Quiros 2002). Allerdings fehlen Studien zur Funktion dieser GS-Klassen innerhalb der Pflanzenresistenz gegenüber Phytophagenfraß. Deshalb wurden zwei A. thaliana -Mutanten mit verändertem aliphatischen bzw. Indolyl-GS-Profil im Vergleich zu Columbia WT auf die Wirtspflanzeneignung für drei verschieden spezialisierte Lepidoptera-Arten getestet.
Solitary bees are important pollinators of flowers. Besides nectar they collect pollen at flowers mainly to provide their larvae with food. Many bee species collect pollen only on a few closely related plant species (oligolecty) (Müller & al. 1997). Little is known about the visual and olfactorial signals they use for host-plant finding (Wcislo & Cane 1996). However, bees can olfactorily distinguish between different pollen species (von Frisch 1923), and a species-specific chemistry of pollen odour is known for some plant species (Bergström & al. 1995, Dobson & al. 1999). Further, it was shown that naïve oligolectic bees recognize their host-plant on the basis pollen volatiles (Dobson & Bergström 2000) and that flower-experienced bees could use pollen odours to assess pollen availability (Dobson & al. 1999). Besides scent, also visual cues are of relevance for host-plant finding, and bees orientate especially spectral contrasts. Biotests with dummy flowers revealed that colour contrast and not intensity and dominant wavelengths are influencing innate behavioural responses (Lunau 1990). Further it was shown that naïve bumblebees were most motivated to land on a flower when visual stimuli from the antheres are combined with olfactorial stimuli from the pollen (1992). We choose Osmia adunca P., which is highly specialized on Echium L., as a model to investigate the importance of floral cues for an oligolectic bee. Because bees learn to associate odours with reward more rapidly than visual cues (Menzel 1985), we hypothesize that scent plays a major role in attraction flower-experienced O. adunca females. We used gas chromatography to compare the scent of three Echium species with the scent of a closely related Anchusa species, and a spectrometer to compare the colour of the three Echium species. Additionally we conducted a biotest to determine the importance of visual and olfactorial signals of Echium for host-plant finding of experienced O. adunca females.