Attract The Opposite Sex With Pheromones

Pheromone Advantage

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Pheromone Advantage Overview

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Background And Further Reading

Jurenka and G. J. Blomquist, Insect pheromones - an overview of biosynthesis and endocrine regulation, Insect Biochemistry and Molecular Biology, 1999, 29, 481-514. www-pherolist.slu.se for a comprehensive list of lepidopteran sex pheromones, species and pictures.

Cephalic sensory structures

Numerous sensory organs, or sensilla (singular sensillum), in the form of hairs, pegs, pits, or cones, occur on antennae and function as chemoreceptors, mechanoreceptors, thermoreceptors, and hygrore-ceptors (Chapter 4). Antennae of male insects may be more elaborate than those of the corresponding females, increasing the surface area available for detecting female sex pheromones (section 4.3.2).

Intraspecific Interactions

Adult emergence occurs as synchronous mass flights. Once near or on the host plants, female pheromones attract male scarabs. Ruteline scarabs utilize sex pheromones derived via fatty acid biosynthesis, whereas scarabs in the not distantly related Melolonthinae utilize amino acid derivatives and terpenoid compounds. The compounds of different classes are released from glands on different parts of the body ruteline pheromones from epithelial cells lining inner surfaces of the abdominal apical segments, for example, or melolonthine pheromones from eversible glands on the abdominal apex. Pheromones used in the other beetle superfamilies span these scarab pheromone classes (e.g., terpenoids in the Curcu-lionidae, fatty-acid-derived aldehydes and acetates in the Elateridae, esters in the Dermestidae).

Alkadienes and Alkatrienes

Are oxidized and cleaved then act as sex pheromones of the yellow-headed spruce sawfly, Pikonema alaskensis (Bartelt et al. 1982) and two 7,11-alkadienes from D. melanogaster are involved in stimulating male courtship of females (Antony and Jallon 1982). The major female sex pheromone from the almond seed wasp, Eurytoma amygdali, was identified as a mixture of two dienes, 6,9-tricosadiene and 6,9-pentacosadiene (Krokos et al. 2001 Mazomenos et al. 2004). An example of an alkatriene is found in the arctiid moth, Utethesia ornatrix, which uses 3,6,9-henei-cosatriene as short range orientation cues for males (Conner et al. 1980). Additional examples can be found in the extensive review by Lockey (1988).

Methyl Branched Alkanes

Twenty-one of the twenty-four cuticular hydrocarbons with known biological activity are methyl-branched alkanes (Nelson 1993). These were functioning as sex pheromones, kairomones and anti-aphrodisiacs. In the last 10 years the number of methyl-branched cuticular hydrocarbons suspected to be involved in chemical communication continues to grow e.g. colony recognition in Polistes wasps (Dani et al. 2001) and fertility signal in ants (Endler et al. 2004). Although numerous studies identified methyl-branched alkanes thought to be involved in various aspects of communication, it has been very difficult to prove this with bioassays. Synthesising these compounds is currently difficult and laborious. Furthermore, establishing the chirality of an isomer and subsequent synthesis of the correct isomer is difficult. However, the diversity of identified methyl-branched alkanes is far greater than any other group of cuticular hydrocarbons (Martin and Drijfhout, in press), since not only can the...

Do Hydrocarbon Profiles Change

Several factors are important for changing the hydrocarbon profile of an insect. In the above section some have already been mentioned. These are genetic factors such as the age of a bee or the gender of a fly. In general the age or gender of an insect can influence the profile of the hydrocarbons present. This genetical component of hydrocarbon production has been well studied in Drosophila spp. (Ferveur 2005). Several genes are known to either induce or reduce hydrocarbon production. Much of this work has been carried out in relation to the sex pheromones in D. domestica that is (Z9)-tricosene. The (Z7) isomer which is produced by the males can either be up-regulated or down-regulated depending on the enzyme. However, the amount of saturated versus unsaturated hydrocarbons can also be altered, as when the PGa14 transposon was inserted within desat1 in D. melano-gaster production of unsaturated hydrocarbons was reduced and saturated hydrocarbon production increased (Marcillac et al....

Nervous And Chemical Integration

The deutocerebrum is largely composed of the paired antennal lobes (Homberg et al., 1989 Hannson and Anton, 2000). These two neuropiles include both sensory and motor neurons and are responsible for initiating both responses to antennal stimuli, especially olfactory and mechanosensory, and movements of the antennae. In species where females produce sex-pheromones the antennal lobes often show sexual dimorphism, being larger with additional interneurons in males. From the antennal lobes, interneurons convey

Box 42 The electroantennogram

Of different sizes and several ultrastructural morphologies. Sensilla respond specifically to sex-signaling chemicals produced by the female (sex pheromones see below). As each sensillum has up to 3000 pores, each 10-15 nm in diameter, there are some 45 million pores per moth. Calculations concerning the silkworm moth suggest that just a few molecules could stimulate a nerve impulse above the background rate, and behavioral change may be elicited by less than a hundred molecules.

Bringing The Sexes Together

To another member of the species (section 4.3.2). Substances emitted with the intention of altering the sexual behavior of the recipient are termed sex pheromones. Generally, these are produced by the female and announce her presence and sexual availability to conspecific males. Recipient males that detect the odor plume become aroused and orientate from downwind towards the source. More and more insects investigated are found to have species-specific sex pheromones, the diversity and specificity of which are important in maintaining the reproductive isolation of a species.

Cannibalistic mating in mantids Mantodea

Courtship displays may be complex or absent, depending on species, but generally the female attracts the male via sex pheromones and visual cues. Typically, the male approaches the female cautiously, arresting movement if she turns her head towards him, and then he leaps onto her back from beyond her strike reach. Once mounted, he crouches to elude his partner's grasp. Copulation usually lasts at least half an hour and may continue for several hours, during which sperm are transferred from the male to the female in a spermatophore. After mating, the male retreats hastily. If the male were in no danger of becoming the female's meal, his distinctive behavior in the presence of the female would be inexplicable. Furthermore, suggestions of gains in reproductive fitness of the male via

The Hows And Whys Of Investigating Natural Enemy Mating Behaviour

Invaluable additions to the predictions made by population models of host-parasitoid dynamics (Luck, 1990). Studying mating behaviour may be critical in predicting the success of biological control introductions and establishments (e.g. Hopper and Roush, 1993) as well as in developing efficient techniques for the mass-rearing of insects that are required in large numbers at particular times, e.g. for field or greenhouse releases (Waage et al, 1985 Hall, 1993 Heinz, 1998). Understanding responses of males to female sex pheromones may be useful in monitoring parasitoid populations (Suckling et al., 2002).

Types of behaviour modifying chemicals

Sex pheromones are used by males to locate females for mating. Typically, female Lepidoptera emit pheromone for a specific period during the day in order to attract males. The pheromone gland in the final segments of the abdomen is extruded so that the pheromone is released to move downwind as a plume of odour. A resting or flying male that detects the pheromone will fly into the wind and follow the plume by upwind anemotaxis (orientation with respect to the direction of the wind). On arriving near the female, the high concentration of odour perceived by the male causes a reduction in forward flight speed and further orientation over the last few centimetres will be both by chemical sensing and visual means (Shorey, 1977). Copulation may then take place. A large number of the sex pheromones of Lepidoptera have now been identified but the behavioural sequences involved in the detection and tracking leading to successful mating has rarely been studied. If insect behaviour is to be...

Mating Behaviour As A Source Of Taxonomic Characters

The success of control programmes is likely to be influenced by the presence of reproductive incompatibility between released wasps. Initial investigations revealed that there were three reproductively incompatible groups among these cultures, and more detailed investigations of response to sex pheromone, courtship behaviour and sperm transfer were carried out to determine the basis of the incompatibility. Males responded normally to the sex pheromones (section 4.3) of incompatible females and courted them actively. However, courtship by incompatible males did not usually induce receptivity by females, which did not stop moving or elevate their abdomens for copulation, and in most cases there was no genital contact or sperm transfer. Reproductive incompatibility is apparently the result either of the failure of females to recognise males as potential mating partners, or of active rejection of males. Molecular investigations showed that the genetic distances between the three...

Functional Terminology For Behavioral Chemicals

Chemical cues involved in host-seeking behavior of parasitoids. All chemical attractants, arrestants, and so on that are important as modifiers of behavior between different organisms are grouped under the general term semiochemicals. Pheromones are semiochemicals that serve to communicate between organisms of the same species sex pheromones are an obvious example. Allelochemicals have effects between species and are further divided into those depending on whether the producing or receiving organism is helped or hurt by the signal. If the species producing the material is helped and the receiving one is harmed, the chemical is called an allomone. Examples of allomones include repellents that a stinkbug may produce to ward off predators such as ants or birds. A substance that harms the producing species but helps the receiving one is called a kairomone. The chemicals produced by insect hosts that serve as cues to parasitoids are kairomones because the parasitoid exploits them to the...

Biosynthesis and detection of pheromones and plant volatiles introduction and overview

The first insect sex pheromone identified was bombykol, (E, Z)-10,12-hexadecadien-l-ol (Butenandt et al., 1959) from the silkworm moth, Bombyx mori (L.). The elucidation of the structure spanned 20 years and required a half million female abdomens. A few years later, (Z)-7-dodecenyl acetate was identified as the sex pheromone of the cabbage looper, Trichoplusia ni (Berger, 1966). By 1970, following the pioneering work by Silverstein on bark beetles, in which three terpenes were identified as a synergistic pheromone blend for Ips paraconfusus (Silverstein et al., 1966), it became recognized that most insect pheromones consisted of multicomponent blends. This has since been shown to be true for most insects, and single-component pheromones are rare. Over the past four decades, extensive research on insect pheromones has resulted in the chemical and or behavioral elucidation of pheromone components from several thousand species of insects, with much of the work concentrating on sex...

Insects And Humans

Chemosterilization is an alternative to sterilization by irradiation in situations where the latter is not practical. For example, it may prove to be useful for species that cannot easily be mass-reared that is, the pest has to be treated directly in the field. It may be possible to combine the chemosterilants with sex pheromones on female mimics to which males would be attracted and attempt to mate, the physical contact thus leading to a male receiving a dose of the chemical. However, the use of chemosterilants has been limited, principally because of concerns about environmental safety.

Box 43 Reception of communication molecules

Aphrodisiacs Danaus

Sex pheromones Male and female conspecific insects often communicate with chemical sex pheromones. Mate location and courtship may involve chemicals in two stages, with sex attraction pheromones acting at a distance, followed by close-up courtship pheromones employed prior to mating. The sex pheromones involved in attraction often differ from those used in courtship. Production and release of sex attractant pheromones tends to be restricted to the female, although there are lepidopterans and scorpionflies in which males are the releasers of distance attractants that lure females. The producer releases volatile pheromones that stimulate characteristic behavior in those members of the opposite sex within range of the odorous plume. An aroused recipient raises the antennae, orientates towards the source and walks or flies upwind to the source, often in a zig-zag track (Fig. 4.7) based on ability to respond rapidly to minor changes in pheromone concentration A spectacular example of...

Box 42 Reception of communication molecules

Moth Releasing Pheromones Diagram

Sex pheromones Male and female conspecific insects often communicate with chemical sex pheromones. Mate location and courtship may involve chemicals in two stages, with sex attraction pheromones acting at a distance, followed by close-up courtship pheromones employed prior to mating. The sex pheromones involved in attraction often differ from those used in courtship. Production and release of sex attractant pheromones tends to be restricted to the female, although there are lepidopterans and scorpionflies in which males are the releasers of distance attractants that lure females. The producer releases volatile pheromones that stimulate characteristic behavior in those members of the opposite sex within range of the odorous plume. An aroused recipient raises the antennae, orientates towards the source and walks or flies upwind to the source, often in a zig-zag track (Fig. 4.7) based on ability to respond rapidly to minor changes in pheromone concentration by direction change (Box 4.2)....

Box 52 Nuptial feeding and other gifts

Courtship displays may be complex or absent, depending on species, but generally the female attracts the male via sex pheromones and visual cues. Typically, the male approaches the female cautiously, arresting movement if she turns her head towards him, and then he leaps onto her back from beyond her strike reach. Once mounted, he crouches to elude his partner's grasp. Copulation usually lasts at least half an hour and may continue for several hours, during which sperm are transferred from the male to the female in a spermatophore. After mating, the male retreats hastily. If the male were in no danger of becoming the female's meal, his distinctive behavior in the presence of the female would be inexplicable. Furthermore, suggestions of gains in reproductive fitness of the male via indirect nutritional benefits to his offspring are negated by the obvious unwillingness of the male to participate in the ultimate nuptial sacrifice - his own life

Relevance to the Sterile Insect Technique SIT

Ulation that are releasing their own sex pheromones. By using sterile insects as living dispensers of pheromone, the level of mating in the natural population is reduced, and when matings do occur they are likely to be between sterile males and females of the natural population.

Solid Phase Micro Extraction

Solid phase micro extraction is a technique invented in 1990 and ever since then it has been used very extensively for many different purposes. The extraction is based on the affinity for compounds to absorb on an absorbent (a liquid stationary phase) (Pawliszyn 1997). It was first introduced by Pawliszyn and co-workers (Arthur and Pawliszyn 1990) for the extraction of organic compounds from an aqueous medium. However, many people have seen the benefits of this technique and now it is used for the analysis of headspace samples to identify volatiles emitted by living plants (Vercammen et al. 2000), identification of sex pheromones (BorgKarlson and Mozuraitis 1996 Rochat et al. 2000) as well as sampling solid samples (Lommelen et al. 2006). The major benefit of SPME is that it is very quick and can be used in connection with a gas chromatograph (GC, see below) or a liquid chromatograph. During sampling the compounds of interest are absorbed on the stationary phase and after a certain...

Spiders

Figure 3.35 Three female-produced sex pheromones of spiders. The first is the dimer of the simple metabolite 3-hydroxy butyric acid and the second is from the primary metabolite citric acid. The third, 8-methyl-2-nonanone, is rather more like some insect pheromones Figure 3.35 Three female-produced sex pheromones of spiders. The first is the dimer of the simple metabolite 3-hydroxy butyric acid and the second is from the primary metabolite citric acid. The third, 8-methyl-2-nonanone, is rather more like some insect pheromones

Iridoids

In 1949 Pavan isolated the substance he called iridomyrmecin from the Argentine ant Iridomyrmex humilis (now Linepithema humile). Iridomyrmecin, dolichodial and other monoterpenes with a methylcyclopen-tanoid structure are called iridoids (Figure 6.12). They act as defensive compounds in ants, stick insects, rove beetles and leaf beetle larvae. Some of the group (e.g. the isomer of nepetalactone shown in Figure 6.12) are also sex pheromones in aphids. Reaction of iridomyrmecin with

Other Functions

The cuticular waxes may have important roles in preventing the entry of microorganisms and in chemical communication (i.e., they serve as semiochemicals). It has been suggested that the waxes may prevent adhesion of microorganisms or may be toxic to them. Cuticular hydrocarbons are also known to serve as contact sex pheromones, for example, in female Diptera and Blattella, attracting or inducing copulatory behavior in males, or serving as an aphrodisiac to keep the male in position until insemination has occurred (Schal et al., 1998). In termites, the cuticular hydrocarbon blend is highly specific and serves as a species-and or caste-recognition pheromone. (See also Chapter 13, Section 4.1.2.) Interestingly, some beetles that live in termite colonies produce the same hydrocarbon profile as the host, enabling them to remain unmolested in the nest. The species-specific nature of the lipids has been turned to advantage by some parasitic Hymenoptera who use these chemical cues (known as...

Alkenes

Many other alkenes are found to be involved in sexual communication (Howard and Blomquist 1982), for example, the stable fly, Stomoxys calcitrans uses (Z9)-C311 and (Z9)-C331 as sex pheromones (Sonnet et al. 1979). Among the social insects, honeybees and wasps are much more responsive to alkenes than alkanes (Dani et al. 2001 Dani et al. 2005 Chaline et al. 2005). Furthermore, the proportions of alkenes in three termite (Macrotermes falciger) phenotypes were associated with inter-group aggression levels (Kaib et al. 2002). Alkenes are often found on the cuticle of ants and correlation studies have shown that they may be used as a fertility signal (Monnin 2006). However, there is now direct evidence using bioassays with synthetic compounds that the (Z9)-C231-(Z9)-C291 and (Z9)-C251-(Z9)-C331 alkenes found in Formica japonica (Akino et al. 2004) and F. exsecta (Martin et al. 2008c) ants respectively, act as nest-mate recognition cues in these species.

Diterpenes

Diterpenes are less volatile because of their greater molecular mass, and cyclized ones are generally solids, but some diterpenes are found in insect glands where they probably act as pheromones. Uncyclized diterpenes, like isomers of springene (Figure 7.1), simple alcohols such as geranylgeraniol, geranyllinalool, and geranylcitronellol and their esters are particularly found in Hymenoptera, but also in termites. These compounds and uncyclized sesquiterpenes are also found in mammalian scent glands. Geranylgeraniol has been found in the labial glands of male bumblebees, in the Dufour glands of the stingless bee Nannotrigona testaceicornis, the ant Ectatomma ruidum, in the female sex pheromones of click beetles Agriotes, Sinapus and Melanotus species, Coleoptera,

Chemical Stimuli

The major olfactory role comes from multiporous sensilla, which are hair- or peg-like setae, with many round pores or slits in the thin walls, leading into a chamber known as the pore kettle. This is richly endowed with pore tubules, which run inwards to meet multibranched dendrites (Box 4.2). Development of an electroantennogram (Box 4.3) allowed revelation of the specificity of chemoreception by the antenna. Used in conjunction with the scanning electron microscope, micro-electrophysiology and modern molecular techniques have extended our understanding of the ability of insects to detect and respond to very weak chemical signals (Box 4.2). Great sensitivity is achieved by spreading very many receptors over as great an area as possible, and allowing the maximum volume of air to flow across the receptors. Thus, the antennae of many male moths are large and frequently the surface area is enlarged by pectinations that form a sieve-like basket (Fig. 4.6). Each antenna of the male...

Semiochemicals

Semiochemicals are chemicals that mediate interactions between organisms (Nordlund, 1981). They are divided into two major groups pheromones, which mediate in intraspecific interactions and allelochemi-cals for interspecific interactions. Each of these groups can be subdivided further allelochemicals as allomones, kairomones, synomones and apneumones (Table 7.7) and pheromones as sex pheromones, alarm pheromones and epideitic or aggregation pheromones. While the majority of research and development has concentrated on pheromones, particularly those of

Mass trapping

Mass trapping, as its name implies, is the use of large numbers of pheromone traps to catch a large proportion of the pest population. This technique has proved unsuccessful for a whole range of lepidopteran and coleopteran pests (Campion, 1989 Jones and Langley, 1998). Lepidopteran sex pheromones only attract the males, hence to be successful, highly efficient traps are needed in order to catch a high enough proportion of the male population to prevent mating with females. Mass trapping is more appropriate with aggregation pheromones, since these attract both males and females. As part of an IPM programme, mass trapping has proved useful in the control of the cotton boll weevil (Anthonomus grandis Dickerson, 1986) and the spruce bark beetle Ips typographus (Raty et al., 1995 Gr goire et al., 1997). In a large mass trapping programme aimed at the control of I. typographus in Norway (1979-1980), the aggregation pheromone component 2-methyl-3-buten-1-ol was dispensed in over 600,000...

Searching For Mates

Chemical signals are usually volatile and provide long-distance information about the signaller's general, but not precise, location. However, virgin females of some species lay down trails of sex pheromones Aphelinus asychis (Aphelini-dae), a solitary parasitoid of aphids (Fauvergue et al., 1995, 1998 Kazmer et al., 1996), Trichogramma brassicae (Trichogrammatidae), a gregarious parasitoid of pyralid moth eggs (Pompanon et al., 1997 Fauvergue et al., 1998a), and Ascogaster reticulatus (Braconidae) an egg-larval parasitoid of tortricid moths (Kamano et al., 1989). In A. asychis, males respond to an encounter with the pheromone by intensively searching in or near the marked area. Because the pheromone declines in activity within less than 24 hours it provides a reasonably precise cue to the location of virgin females. Fauvergue et al. (1998a) compared the response to trails and patches of pheromones deposited on the substrate by males of A. asychis and T. brassicae.

Host Finding

Other parasitoids are attracted from longer distances directly to hosts. Some parasitoid females respond to pheromones produced by their host. Parasitoids of the European elm bark beetle, Scolytus multistriatus, are attracted to multilure, the aggregation pheromone of adult beetles. Aphytis spp. (Hymenoptera Aphelinidae) are attracted to the sex pheromone produced by their host, California red scale (Aonidiella aurantii). A number of true bugs produce sex pheromones that are attractive to a variety of fly (Tachinidae) and to hymenopteran parasitoids. Some parasitoids are drawn from a distance to chemicals produced by plants in response to damage caused by host herbivores. For example, the braconid larval parasitoid Cotesia marginiventris responds to volatile terpenoids released from corn seedlings as a result of eating damage caused by host Spodoptera caterpillars. These chemicals may be components of the induced resistance that plants have developed against pathogens and herbivores....

Behavior And Ecology

Mating in cockroaches generally is preceded by courtship behavior facilitated by sex pheromones. In some species a blend of volatile compounds is produced by virgin females to attract and orient males (e.g., Periplaneta species and the brownbanded cockroach). In the German cockroach, the sex pheromone is a blend of nonvolatile and volatile cuticular components that elicits courtship by males following palpation of the female's integument by the male's antennae. Once courtship is initiated in the male, he turns away from the female and raises his wings to expose dorsal tergal glands the female feeds on pheromones from these glands as the male grasps her genitalia with his pair of caudal claspers. Most species copulate in an end-to-end position. During the hour or so that follows, a spermatophore is formed and passed from the male into the genital chamber of the female. Only about 20 of females mate again after the first gonotrophic cycle.

Pheromones

Pheromones may be arranged in rather broad, sometimes overlapping, categories based on their functions. There are sex pheromones, caste-regulating pheromones, aggregation pheromones, alarm pheromones, trail-marking pheromones, and spacing pheromones. 4.1.1. Sex Pheromones In the term sex pheromones are included chemicals that (1) excite and or attract members of the opposite sex (sex attractants), (2) act as aphrodisiacs, (3) accelerate or retard sexual maturation (in either the opposite and or same sex), or (4) enhance fecundity and or reduce receptivity in the female following their transfer during copulation.

Attractants

The function of pheromones in the biology of many stored-product and fabric pests follows two general patterns. These are sex pheromones for the species that have short-lived adults, and aggregation pheromones for the species that have long-lived adults. The short-lived adults usually do not feed, and mating and oviposition are the chief activities of the adults. Soon after emergence, females of these species usually produce a strong sex pheromone to lure males for mating. The long-lived adult males and females feed, and males generally produce an aggregation pheromone to attract other males or both males and females. Females of these species often produce sex pheromones.

Pyralidae

Of most pyralids are negatively phototactic until they are about to pupate. They are also sensitive to crowding and this condition can delay development and result in small caterpillars and adults. Most of the stored-food pyralids produce sex pheromones. The chief component of the female pheromone of several species is cis-9, trans-12-tetradecadien-i-yl acetate. Within 24 h of emergence females assume the calling posture, in which the abdomen is lifted between the wings and the scent glands are extruded. Males locate females by flying against air currents and orient using the concentration of the pheromone in the air stream. Sex pheromones have been extracted from the wing glands of male Ephestia elutella and Plodia interpunctella. The rapid wing beating of males prior to mating may disperse these chemicals, which appear to be important in the final stage ofcourtship. The pheromones released from male P. interpunctella produce a turning response in receptive females.

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