Academic literature on the topic 'Scent marking'

Abstract In wolves Canis lupus, scent marking plays an important role in territory defence. In Europe, studies on patterns of scent marking in wolves have mostly been conducted in mountains or primeval forests, but since these areas are characterised by low human activity, the impact of people on this behaviour has been neglected. We conducted a study that combined genetic methods with an analysis of the spatial distribution of wolf territory markings in lowland managed forests with high human activity. We found that scent markings are deposited by all members of wolf family groups. Wolves most intensively marked crossroads and their vicinity, especially on roads only accessible for four-wheel drive cars. Our study provides further evidence that crossroads of forest roads play a crucial role in wolf scent marking. The results of our study may be useful during inventories of wolf populations based on collecting indirect signs of their presence or non-invasive genetic sampling.

The objectives of this study were to describe the scent-marking behavior of sympatric wolves and coyotes, and determine whether scent marking has an interspecific behavioral significance and, if so, whether it is involved in maintaining ecological separation of the two species. The hypotheses tested were that coyotes travelling within active wolf areas reduce their frequency of marking and avoid wolf scent marks, and that wolves react to sympatric coyotes as transgressing conspecifics, and increase their scent marking. Marking by the two species was similar, both canids using the same scent mounds. Coyotes did not avoid areas marked by wolves, nor did they minimize evidence of their own activity. Coyotes increased their marking significantly in response to wolves. Wolves, however, did not respond to marks by coyotes.

AbstractDifferences in the protein content of individuals may affect their response to signals produced by opposite-sex conspecifics and how opposite-sex conspecifics respond to the individual's own signals. Many terrestrial mammals use over-marks to communicate with potential mates. In this study, we determined whether over-marking behavior is affected by the protein content of the top- and bottom-scent donors of an over-mark. We tested two hypotheses by performing two experiments on meadow voles, Microtus pennsylvanicus, in which the top- or bottom-scent donors of an over-mark were fed a diet containing either 9, 13, or 22% protein; these protein concentrations are similar to those that voles may consume in free-living populations. In experiment 1, we varied the protein content of the top-scent voles but not that of the bottom-scent donor. We tested the hypothesis that top-scent donor voles fed a diet high in protein content deposit more scent marks and more over-marks than do top-scent donor voles fed diets lower in protein content. In experiment 2, the top-scent voles were fed a 22% protein diet but the protein content of the diet of the bottom-scent donor varied. We tested the hypothesis that the top-scent donor will deposit more scent marks if the bottom-scent vole was fed a diet high in protein content than if it was fed a diet lower in protein content. Protein content of the top-scent vole's diet did not affect the number of scent marks and over-marks it deposited. Likewise, the protein content of the bottom-scent vole did not affect the number of scent marks and over-marks deposited by the top-scent vole.

Scent marking food resources is expected to enhance foraging efficiency reducing search time. Many social bees exhibit this behavior, but scent-marking is absent in social wasps, except forVespa mandarinia. We tested for scent marking in the swarm-founding wasp,Polybia occidentalis. This wasp has moderately large colonies and utilizes resources that are concentrated in time and space, making scent marking profitable. Also, this wasp uses chemical markings to lead nestmates to a new nest site during swarm emigration, making it possible that it could use the same behavior to recruit nestmates to a food source. Foragers from 11 colonies were given a choice between a previously visited feeder and an unvisited one, both containing a rich, unscented sucrose solution. There was no difference in the number of visits to the two treatments. However, some individuals chose the feeder on one side more often. We conclude that foragers of this species of wasp do not use odor marks left behind by nestmates to find food, but they do exhibit the tendency, when returning to a food source that has not been depleted, to choose a resource based on its relative position, presumably by using visual cues.

Primates are traditionally considered to have a poor sense of smell. However, olfaction is important for non-human primates as demonstrated by conspicuous scent-marking behaviours in lemurs. We studied two pairs (n = 4) of crowned lemurs (Eulemur coronatus) housed at Colchester and Twycross zoos (UK) by combining behavioural observations and chemical analyses of scent-marks and glandular swabs. We recorded observations of olfactory behaviours for 201 h using instantaneous scan sampling. We investigated the volatile compounds of ano-genital odour secretions (n = 16) using solid-phase microextraction and gas chromatography-mass spectrometry. Males scent-marked most frequently, displaying ano-genital marking for allomarking, head marking for countermarking and wrist marking in specific areas of the enclosure. Females displayed ano-genital marking, predominantly on feeding devices. We detected a total of 38 volatile components in all male ano-genital scent-marks and 26 in all female samples of ano-genital odour secretions, including a series of esters, aldehydes, ketones, alcohols, terpenes, volatile fatty acids and hydrocarbons that have been identified in odour profiles of other primates. In conclusion, we found sexual dimorphism in crowned lemur scent-marking. Male head and wrist marking behaviours might play defensive territorial functions, while ano-genital marking would be related to socio-sexual communication as chemical mate-guarding. Female ano-genital marking might be involved in resource defense.

Identifying the best control method for problematic wildlife is an ever present issue in wildlife management. Popular control methods have ranged from lethal techniques, extirpating the animal, to multiple non-lethal methods focused on deterring undesired behavior. In the past, lethal methods were the preferred choice. However, with increased awareness of the need for biodiversity conservation, new management methods focus on non-lethal control, with emphasis on exploiting aspects of naturally occurring organismal behaviors and ecology. Over the past decade, technological advances in extraction method’s and equipment have also developed new techniques providing a broader range of information about species biology for management use. One of the most well documented conflicts between wildlife and humans is that of the wolf. Using advanced technology and new techniques, we investigated the implication of using chemosensory signals in canid urine to modify behavior as a possible non-lethal alternative in large predator management. Here we used the SBSE method coupled with improved GC/MS equipment to analyze the volatile organic compounds in the urine of four canid species, gray wolf (Canis lupus), red wolf (Canis rufus), wolf-dog hybrids (Canis familiaris) and the domestic dog (Canis familiaris) in order to create working urinary profiles. The extraction method identified several compounds also seen in the urinary profiles of other large predators. In addition, similarities and differences were also noted between taxa and the sexes, and these can be further explored in future studies. Two identified urinary compounds, acetophenone and methyl propyl sulfide, were selected for further behavioral evaluation. We focused on these compounds and their influence as chemosensory signals triggering urine marking events in both the gray wolf and red wolf. Behavioral observations of the effects of these two chemicals indicated they elicited responses from captive wolves. At each of the three study sites, the combination of these chemicals produced urine-marking events along the territory boundary by dominant animals. As a result, the investigation focused on what triggered the urine-marking events, the chemicals themselves, their combination, or the breakdown of the chemicals producing other odorants. It was found that there was no significant degradation of the chemicals over time and environmental conditions produced no significant breakdown of the acetophenone prior to the addition of methyl propyl sulfide. This posed a number of new questions and illustrated the need for additional behavioral studies. The results of this study analyzing chemosensory signals in canid urine, provides biologists with new information to aid in the development of new non-lethal management strategies for handling problematic wildlife as well as providing useful information for future research involving reproduction, predator/prey dynamics, territory maintenance, and a host of other studies focusing on animal ecology in association with chemosensory signaling.

This thesis examines how scent marking in Eurasian beaver (Castor fiber) functions in territorial defence. Beavers usually deposit scent (castoreum and/or anal gland secretion (AGS)) onto small piles of mud and debris, and all age classes and both sexes participate in marking. I hypothesized that scent marking plays an important role in territory defence of free-ranging Eurasian beavers and investigated the following issues. (1) Which factors affect scent-marking behaviour? (2) How are scent marks distributed temporally and spatially during an annual cycle? (3) Is castoreum and/or AGS used in territorial defence? (4) How does this species respond to simulated territorial intruders? (5) Can the Eurasian beaver discriminate between scent from neighbours and strangers, and between scent from its own species and that of the North American beaver (C. canadensis)?

I show that scent marking plays a significant indirect role in territorial defence by the Eurasian beaver. The number of scent marks was density dependent. Beavers with many close neighbours (highly challenged) may need to scent mark more often to be unambiguously recognised as territory owners. However, high-density sites may also be of better quality, providing territory holders with more energy to spend in defence and more reasons to defend. There was a significant positive correlation between the number of scent marks and both the duration of territory occupancy and length of wooded banks. Therefore, residents appear to invest more in scent marking in good quality territories, and when a territory has been occupied for a relatively long time. Theoretically, the greater potential value of the territory for residents, in contrast to intruders, makes it worth fighting harder for.

Territories were scent marked significantly more often in spring when dispersal of 2- years-olds normally occurs and scent marks were concentrated near territorial borders, apparently to maximize the signal effect to potential trespassers on or before entering the territory. Significantly more scent marks were constructed upstream than downstream of the lodge, probably because the movement of dispersing individuals is predominantly downstream. These results support the border maintenance hypothesis.

From January through March castoreum was almost exclusively deposited on scent marks and appears therefore to be the main scent signal used in the defence of Eurasian beaver territories. AGS was rarely deposited and appears to have another function.

Eurasian beaver showed territorial behaviour when an "intruder", in the form of artificially-constructed experimental scent mounds (ESMs containing castoreum from alien adult males, was placed inside the territory. They destroyed the ESMs and overmarked with their own scent in 80% of the trials. Countermarking appears to have been an attempt to mask the odour of alien adult male conspecifics with their own odours. This result therefore gives some support to the scent-masking hypothesis. Scent marks could thus provide a reliable advertisement of an individual’s ability to dominate or defend the area, since only those successfully dominating the area can ensure that their marks both predominate and are more recently deposited than those of any challenging competitors. The countermarking may therefore advertise that the territory is occupied and signal the costs of competition if the threat is ignored. I frequently observed that beavers, after visiting the ESMs, started to patrol the territory. A lack of response to ESMs without castoreum indicated that beavers were responding to the smell of castoreum and not to the sight of the scent mound itself.

Eurasian beavers sniffed both castoreum and AGS from a stranger significantly longer than those from a neighbour. They responded aggressively significantly longer to castoreum, but not to AGS, from a stranger than from a neighbour. When ESMs were allowed to remain overnight and the response measured the following morning, beavers responded significantly stronger to both castoreum and AGS from a stranger than from a neighbour. These findings indicate that Eurasian beavers can use scent to discriminate between neighbours and strangers, thereby supporting existence of the “dear enemy” phenomenon (reduced aggression towards familiar occupants of neighbouring territories).

Eurasian beavers spent significantly longer time responding aggressively to conspecific than to heterospecific (North American beavers) ESMs. They also responded significantly more aggressively to conspecific than to heterospecific ESMs overnight. Gas chromatographic comparisons of castoreum showed that differences between species accounted for 34% of the total variation in compounds detected, while differences between sexes accounted for 13%. For AGS, 49% and 46% of this variation was explained by differences between species and sex, respectively. The results confirm the hypothesis that the Eurasian beaver discriminates between scent marks of the two species, i.e. exhibits species discrimination abilities. This indicates that the Eurasian beaver would regard intrusive scent marks from the North American beaver as a lesser territorial threat than from a conspecific, and would therefore be less likely to spend time and energy countermarking these scent marks.

In conclusion, my study has contributed to a better understanding of the function of territorial scent marking in the Eurasian beaver by demonstrating their capability of transmitting odorous messages efficiently, both temporally and spatially, and their ability to countermark and discriminate ESMs from intruders of different degrees of threat. My results lend support to the idea that the function of territorial scent marking in the Eurasian beaver is to advertise related dominance status, thereby providing opportunities for intruders to assess the presence of the owner, and thus reducing the costs of agonistic conflicts for both the owner and intruder (the status advertisement hypothesis). My results also support the general scent-matching hypothesis, i.e. its predictions 1 (mark where intruders are most likely to encounter marks), 3 (make themselves available for scent matching by intruders) and 4 (remove or replace marks of others) were all supported. However, prediction 2 (mark themselves with the substances used to mark the territory) needs to be clarified. It’s still unclear whether beavers smear castoreum on their pelage, and/or mark themselves with AGS in order to waterproof the fur, and thereby simultaneously function as a “living-scent mark”. The next step should be to clarify these issues. However, the function of scent marking suggested here is not necessarily the only functional mechanism, as one function need not necessarily exclude others. Two other main functions for scent marking in Eurasian beavers that cannot be entirely ruled out are that scent marks may be used to label and thereby defend resources within the territory (the labelling resources hypothesis), and that marking is related to reproduction (e.g. by advertising reproductive status and guarding the mate during the breeding period). My work has emphasized intergroup communication. However, more work is needed to clarify the role of scent marks in intragroup communication.

Denne avhandlingen eksaminerer hvordan duftmarkeringer fungerer i territorieforsvaret hos eurasiatisk bever (Castor fiber). Beveren avsetter vanligvis duft (castoreum fra bevergjelpungene og/eller sekret fra analkjertlene (AKS)) på små hauger av leire og planterester, og alle aldersgrupper og begge kjønn deltar i markeringen. Jeg satte fram hypotesen at duftmarkeringer spiller en viktig rolle i territorieforsvaret til fritt levende eurasiatisk bever, og undersøkte følgende spørsmål. (1) Hvilke faktorer virker inn på duftmarkeringsatferden? (2) Hvordan er duftmarkeringer fordelt i tid og rom i løpet av et helt år? (3) Er castoreum og/eller AKS brukt i forsvaret av territoriet? (4) Hvordan reagerer denne arten på simulerte inntrengere i territoriet? (5) Kan den eurasiatiske beveren diskriminere mellom duft fra naboer og fremmede, og mellom duft fra sin egen art og den nordamerikanske beveren (C. canadensis)?

Jeg viste at duftmarkeringer spiller en signifikant, indirekte rolle i forsvaret av territoriet hos den eurasiatiske beveren. Antall duftmarkeringer var tetthetsavhengig. Bever med mange nære naboer (høyt utfordret) trenger sannsynligvis å duftmarkere oftere for å bli utvetydig gjenkjent som eier av territoriet. Plasser med høy tetthet er imidlertid kanskje også av bedre kvalitet, noe som gir eierne av territoriet mer energi å bruke på forsvaret, og flere grunner for å forsvare. Det var en signifikant positiv korrelasjon mellom antall duftmarkeringer og varigheten av okkupasjonen av territoriet samt lengden av banker med trær. Bofaste ser derfor ut til å investere mer i duftmarkeringer i territorier med god kvalitet og når et territorium har vært okkupert for relativ lang tid. Teoretisk, jo store potensiell verdi territoriet har for de bofaste, i kontrast til inntrengere, desto hardere bør eieren slåss for å beholde det territoriet.

Territoriet ble duftmarkert signifikant oftere om våren når spredningen av 2-åringer normalt skjer, og duftmarkeringene ble konsentrert nær grensene til territoriet, tilsynelatende for å maksimere signaleffektiviteten til potensielle inntrengere før de entrer territoriet. Signifikant flere duftmarkeringer ble konstruert oppstrøms i forhold til nedstrøms av hytta, sannsynligvis fordi bevegelsen av individer på vandring hovedsakelig er nedstrøms. Disse resultatene støtter hypotesen om grenseopprettholdelse.

Castoreum ble nesten utelukkende avsatt på duftmarkeringer fra januar til ut mars og ser ut til å være hovedlukten brukt i forsvar av eurasiatiske beverterritorier. AKS ble sjelden avsatt og har muligens en annen funksjon.

Den eurasiatiske beveren viste territorial atferd når en ”inntrenger”, i form av kunstig konstruerte eksperimentelle duftmarkeringshauger (EDH’er) med castoreum fra fremmede voksne hanner, ble plassert inne i territoriet. De ødela EDH’ene og overmarkerte med sin egen lukt i 80% av forsøkene. Overmarkeringen ser ut til å ha vært et forsøk på å maskere duften fra de fremmede voksne hannene med sin egen duft. Disse resultatene gir dermed noe støtte til duftmaskeringhypotesen. Duftmarkeringer kan derfor sørge for en troverdig annonsering av et individs evne til å dominere eller forsvare et område, siden bare de som suksessfullt dominerer et område kan sikre at deres markeringer både dominerer og er nyligere avsatt enn de fra en utfordrende konkurrent. Overmarkeringen annonserer derfor muligens at territoriet er opptatt og signaliserer kostnaden av konkurransen hvis trusselen ignoreres. Jeg observerte at beverne ofte startet å patruljere territoriet etter å ha besøkt EDH’ene. En mangel på respons på EDH’er uten castoreum indikerer at beveren reagerte på duften av castoreum og ikke på synet av duftmarkeringshaugen.

De eurasiatiske beverne snuste på castoreum og AKS fra en fremmed, signifikant lenger enn fra en nabo. De reagerte aggressivt, signifikant lenger på castoreum, men ikke på AKS, fra en fremmed enn fra en nabo. Når EDH’ene forble ute over natta og responsen ble målt den påfølgende morgenen, reagerte beverne signifikant sterkere på både castoreum og AKS fra en fremmed enn fra en nabo. Disse resultatene indikerer at den eurasiatiske beveren kan bruke duft for å diskriminere mellom naboer og fremmede, og gir dermed støtte til tilstedeværelsen av ”kjære fiende” fenomenet (redusert aggresjon mot kjente okkupanter på naboterritoriene).

De eurasiatiske beverne tilbrakte signifikant lenger tid på å reagere aggressivt på artsfrenders enn ikke-artsfrenders (nordamerikanske bevere) EDH’er. De reagerte også signifikant mer aggressivt på artsfrenders enn ikke-artsfrenders EDH’er over natt. Sammenligninger av castoreum gasskromatogram viste at forskjeller mellom artene forklarte 34% av den totale variasjonen i forbindelsene oppdaget, mens forskjeller mellom kjønnene forklarte 13%. For AKS, var henholdsvis 49% og 46% av denne variasjonen forklart av forskjeller mellom arter og kjønn. Disse resultatene bekrefter hypotesen at den eurasiatiske beveren diskriminerer mellom duftmarkeringer fra de to artene, med andre ord utøver arts diskrimineringsevner. Dette indikerer at den eurasiatiske beveren vil anse påtrengende duftmarkeringer fra den nordamerikanske beveren å utgjøre en mindre territoriell trussel enn fra en artsfrende, og vil derfor mindre sannsynlig bruke tid og energi på å overmarkere disse duftmarkeringene.

Jeg konkluderer med at mitt studium har bidratt til en bedre forståelse av funksjonen av duftmarkering i territoriet til den eurasiatiske beveren ved å demonstrere deres evne til å overføre duftbeskjeder effektivt, både i tid og rom, og deres evne til å overmarkere og diskriminere EDH’er fra inntrengere som utgjør ulik grad av trussel. Mine resultater gav støtte til ideen at funksjonen for duftmarkering av territoriet hos eurasiatisk bever er å annonsere dominans status, og dermed sørge for muligheter for inntrengere til å vurdere tilstedeværelsen av eieren som vil redusere kostnadene av de agonistiske konfliktene for både eier og inntrenger (statusannonseringshypotesen). Mine resultater støtter også den generelle duftssammenligningshypotesen, med andre ord dens prediksjoner 1 (duftmarker hvor inntrengere er mest sannsynlig å møte disse), 3 (gjør seg tilgjengelig for duftsammenligning av inntrenger) og 4 (fjern eller erstatt duftmarkeringer av andre) ble alle støttet. Prediksjon 2 (duftmarker seg selv med duften brukt til å markere territoriet) trenger imidlertid å klargjøres. Det er fortsatt uklart om beveren smører castoreum på pelsen, og/eller markerer seg selv med AKS for å gjøre pelsen vanntett og dermed fungere samtidig som en ”levende duftmarkering”. Det neste steget bør være å redegjøre for disse spørsmålene. Funksjonen til duftmarkering som er foreslått her er imidlertid nødvendigvis ikke den eneste funksjonelle mekanismen, siden en funksjon ikke trenger å utelukke andre. To andre hovedfunksjoner for duftmarkering hos eurasiatisk bever som ikke helt kan utelukkes er at duftmarkeringer kan bli brukt til å merke og dermed forsvare ressurser innen territoriet (hypotesen om ressurs merking), og at duftmarkeringen er relatert til reproduksjonen (for eksempel ved å annonsere reproduktiv status og bevoktning av maken i løpet av paringstiden). Mitt arbeid har lagt vekt på kommunikasjonen mellom familiegrupper. Mer arbeid trengs imidlertid for å klargjøre duftmarkeringens rolle i kommunikasjonen innen familiegrupper.

Paper V reprinted with kind permission of Elsevier, sciencedirect.com

defend feeding territories, and that the spatinl distribution of 'patches' of earthworms (their major prey) determines the size of badger territories. Such a food-based model has been widely accepted, but has a number of equivocal assumptions which are discussed. Recently Roper, Shepherdson & Davies (1986) proposed an alternative model of territorial organisation, based on the seasonal pattern of territory marking with faeces and anal gland secretion, which suggested that territoriality in badgers may be more related to defence of oestrus females by resident males than to defence of food resources. This hypothesis relies on an un-quantified correlation between seasonal patterns of territory marking, mating and road mortality. Here, I attempt to test the strength of the association between these distributions and to test predictions about the relative contribution of the two sexes to territorial defence. To do this I present further data on the deposition of faeces and anal secretion at latrines, together with new data on the seasonality of road mortality and bite-wounding in badgers. In addition, I report the results of experiments in which latrines were continually monitored in an attempt to assess the relative contribution of the two sexes to territory marking and patrolling. Finally, I report the results of a chemical investigation (using gas chromatography) of the scent profile of anal gland secretion. My results confirm the bimodal seasonal pattern of deposition of faeces and anal secretion at latrines, but whilst the two distributions were similar, there were differences which suggested that the two territorial markers may have different functions. The seasonal pattern of deposition of anal secretion showed essentially the same distribution as data on mating, testis weight, bite wounding and road mortality and I concluded that my results were consistent with the anti-kleptogamy hypothesis. However, the seasonal pattern of deposition of faeces at latrines could more easily be explained by seasonal changes in food availability. My results showed that the incidence of bite wounding and of patrolling at latrines, was higher in males than females, which is consistent with predictions derived from the anti-kleptogamy model, but not with food-based models which predict territory defence to be shared equally amongst group members. Finally, chemical analysis of anal secretion revealed that whilst it probably did not signal the sex or identity of it's producer it may carry information about group membership. In addition, the secretion was found to be of low volatility and composed of long-chain fatty acids some fifteen of which were identified. These results are consistent with the idea that anal secretion acts as a long-term territory marker. In conclusion I suggest that my results for the deposition of anal gland secretion at latrines are consistent with the hypothesis proposed by Roper et al., (1986) that territoriality in badgers may at least be partly linked to the defence of oestrus females by resident males. By contrast seasonal variation in defecation at latrines may at least be partly explained by seasonal changes in food availability. Given that food and mates are the most important resources for the survival of an animal, in the short-term and long-term respectively, it is likely that models of the territorial spacing pattern of badgers would have to take both resources into account.

Thesis (PH.D.)--Michigan State University. Zoology and Ecology, Evolutionary Biology and Behavior, 2008.
Title from PDF t.p. (viewed on Aug. 11, 2009) Includes bibliographical references (p. 156-178). Also issued in print.

Home ranges of males (1204 km2) and females (1510 km2) were similar. Female home range size was positively related to the dispersion of prey and generally, but not exclusively, they displayed home range fidelity. Overlap between female home ranges was extensive, although they rarely met up. Male home ranges overlapped extensively and there was no difference in size between coalition and single males. Males overcame the problem of scent marking a large home range by concentrating scent marks in core areas. Generally female cheetah home range size is affected by resource productivity, although where prey are migratory, or in fenced reserves where movements are constricted, and areas where disturbance is severe, this may be different. Southern Kalahari males apparently need large home ranges to increase the likelihood of locating wide-ranging and sporadically receptive females. Mean dispersal distance for subadult males (96 km) was further than for females (39 km).

This chapter examines the scene as a segment of dramatic action marked at the beginning and end by an empty stage, scene as big effect, and scene as fictional setting. It begins with a discussion of the scene to which Robert Greene alludes inGreenes, groats-vvorth of witte(1592). It then considers the folio text ofHenry VI, Part Threeto show how acts and scenes were marked in plays printed between 1590 and 1630. It also discusses two ways that scenes can be registered verbally in scripts: marking and remarking. Finally, it explores the connection between physical means and theatrical ends in early modern usage in the context of scene, along with senses of ‘scene’ that may be increasingly remote from theatres as physical structures but that nonetheless maintain an important relationship with theatrical performance as a way of framing and understanding human experience.

The sense of smell has a great importance in our daily life. In recent years, smells have been used for marketing purposes with the aim of improving the person’s mood and of communicating information about products as household cleaners and food. However, the scent design discipline can be also applied to any kind of products to communicate their features to customers. In the area of Virtual Reality several researches have focused on integrating smells in virtual environments. The research questions addressed in this work concern whether Virtual Prototypes, including the sense of smell, can be used for evaluating products as effectively as studies performed in real environments, and also whether smells can contribute to increase the users’ sense of presence in the virtual environment. For this purpose, a Virtual Reality experimental framework including a prototype of a wearable olfactory display has been set up, and experimental tests have been performed.

Phytophagous insects respond to chemicals from their host plants in a number of ways, including orientation or attraction in response to volatiles produced by plants. Orientation to odors from host plants may occur in order to locate food, mates, or oviposition sites. A detailed understanding of these behaviors are the chemical stimuli evoking them may provide useful means for attracting and trapping insect pests of crop plants. Heliothine moths (Helicoverpa and Heliothis herein) include a number of major pests of cultivated crop plants throughout the world. In North America, these include Heliothis virescens and Helicoverpa zea. In much of Eurasia (including Israel) Australia, and Africa, these include Helicoverpa armigera and Helicoverpa peltigera. These 4 species of concern all are attracted to odorants from host plants (Tingle and Mitchell 2992, Mitchell et al 1991, 1992 BARD feasibility study report). Host plant chemicals also play a role in the sexual behavior of Helicoverpa species. Synthesis and possibly release of sex pheromone in H. zea and H. phloxiphaga is stimulated by kairomones from hosts plants (Raina 1988, 1992). Pheromona scent marking on host plants also occurs in H. virescens and H. zea. Studies of several other insects, including the cabbage looper Trichoplusia ni, have a variety of other behaviors may occur in association with host plants, including the use of plants as sexual rendesvous sites and of direct involvement of plant chemicals in sexual behavior. Some pest species of moths also may use host plants as adult food sources. These studies were undertaken to develop a more thorough understanding of how Heliothis/Helicoverpa moths use host plant odorants to locate and select foods, mates, and oviposition sites. We used Heliothis virescens and Helicoverpa zea in Florida, and Helicoverpa armigera and Helicoverpa peltigera in Israel as objects of study because of their pest status. It is hoped that such an understanding will provide direction for work to discover and develop novel means to control these pests through behavioral manipulation. The specific objectives of the proposal were to 1) identify host odor affects on known Heliothine sexual behavior, 2) identify novel sexual behavior that is how dependent, 3) isolate and characterize host kairomones important to pest Heliothine host and mate-location behavior, and 4) investigate female attraction to males.