Behavioural and electrophysiological responses by reproductive female Neogobius melanostomus to odours released by conspecific males

Behavioural and electrophysiological responses of reproductive and non‐reproductive female round gobies Neogobius melanostomus to water previously occupied by male round gobies (reproductive male water) were compared. Reproductive females spent more time than non‐reproductive females in a tank near the input of water conditioned from reproductive males. Also, reproductive females swam significantly faster than non‐reproductive females, suggesting that reproductive male odour may have activated spawning behaviour. Olfactory epithelial field potential measurement (electro‐olfactogram, EOG) showed that reproductive male water was a potent olfactory stimulus to reproductive females, but not to non‐reproductive females. Reproductive females responded significantly more than non‐reproductive females to solid‐phase (octadecylsilane) extracts of reproductive male water. Also, when these extracts were separated on reverse phase high performance liquid chromatography (HPLC), reproductive females showed noticeably greater responses than non‐reproductive females to the fractions that eluted between 30 and 40 min. The behavioural data support the hypothesis that reproductive male round gobies release compounds into the water that attract potential mates. The EOG data indicate that these compounds can be quantitatively extracted from the water and be partially purified by HPLC. The evidence is not sufficient to indicate whether or not the compounds are steroids. The relatively early elution time on HPLC, however, suggests that if these compounds are steroids, then it is more likely that they will be conjugated rather than free steroids.     

Evidence of a Male Sex Pheromone in the Round Goby (Neogobius melanostomus)

The reproductive success of the round goby (Neogobius melanostomus), an invasive fish, may be mediated by the use of pheromones. We hypothesized that reproductive male (RM) round gobies release sex pheromone to which reproductive females (RF) respond. In this study, we compared behavioural and electrophysiological responses of reproductive and non-reproductive female round gobies to conspeci fic males. Results of behavioural experiments in the laboratory showed that RF spent significantly more time near the source of the male odour compared with odours from control water. However, RF did not distinguish between odours from non-reproductive male (non-RM) water and control water. Non-reproductive females (non-RF) were not attracted to odours released from RM or non-RM water. Results of electro-olfactogram (EOG) responses showed that both RF and non-RF discriminated between HPLC fractionated RM and non-RM odours. However, the EOG responses of RF were about eight-fold higher than non-RF exposed to RM odours. These findings confirm that RM round gobies release a pheromone signal that attracts RF. The results of this research may be useful in developing control strategy using natural pheromones to disrupt the reproductive behaviour of the invasive round goby and to curtail its effects on native species. An erratum to this article is available at .     

Male competition is disruptive to courtship in the Pacific blue‐eye

The present study examined the effects of competition on male courtship in the Pacific blue‐eye Pseudomugil signifer , a species of fish where females have previously been shown to use courtship, but not male fighting prowess, as an important mate choice cue. Courtship bouts directed to a stimulus female were shortest when two males were allowed to freely interact (contact treatment) and longest when there was only one male (non‐interaction). Courtship length in trials where one of two males was confined to a clear cylinder (visual) was intermediate between the other two treatments. Courtship in visual and contact treatments was constantly disrupted. The percentage of interrupted courtships was higher for the contact compared to visual treatment where aggressive interactions were also longer in duration and took place more frequently. Within contact trials, dominant males courted longer than subordinates despite both males experiencing comparable rates of courtship disruption. These results suggest that male‐male competition can have important implications for adaptive female choice particularly in circumstances where the benefits being sought by females are unrelated to male fighting ability.     

Does the reproductive status of male round gobies (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) influence their response to conspecific odours?

While pheromone communication has been well documented in the female round goby, male responses have not been examined. We determined if male round gobies responded to odours of their conspecifics, and whether that response varied with their reproductive status (reproductive and non-reproductive). No significant differences were observed for any treatment (dechlorinated water, or water conditioned by reproductive males (RM) and non-reproductive males (NRM) and females), although RM spent more time inside shelter than NRM (p < 0.05). Morphologically, length and weight did not vary, but gonadosomatic index, seminal vesicle somatic index and relative head width did differ between the two male groups (p = 0.000). Round goby males guard nests in cavities, and so a preference for shelter with increasing reproductive status is expected. Also, a lack of male responses to conspecific odours may be adaptive in this species, since males nest together in high densities, where increased aggression might prove detrimental for nesting males.     

Spider sex pheromones: emission, reception, structures, and functions

Spiders and their mating systems are useful study subjects with which to investigate questions of widespread interest about sexual selection, pre- and post-copulatory mate choice, sperm competition, mating strategies, and sexual conflict. Conclusions drawn from such studies are broadly applicable to a range of taxa, but rely on accurate understanding of spider sexual interactions. Extensive behavioural experimentation demonstrates the presence of sex pheromones in many spider species, and recent major advances in the identification of spider sex pheromones merit review. Synthesised here are the emission, transmission, structures, and functions of spider sex pheromones, with emphasis on the crucial and dynamic role of sex pheromones in female and male mating strategies generally. Techniques for behavioural, chemical and electrophysiological study are summarised, and I aim to provide guidelines for incorporating sex pheromones into future studies of spider mating. In the spiders, pheromones are generally emitted by females and received by males, but this pattern is not universal. Female spiders emit cuticular and/or silk-based sex pheromones, which can be airborne or received via contact with chemoreceptors on male pedipalps. Airborne pheromones primarily attract males or elicit male searching behaviour. Contact pheromones stimulate male courtship behaviour and provide specific information about the emitter's identity. Male spiders are generally choosy and are often most attracted to adult virgin females and juvenile females prior to their final moult. This suggests the first male to mate with a female has significant advantages, perhaps due to sperm priority patterns, or mated female disinterest. Both sexes may attempt to control female pheromone emission, and thus dictate the frequency and timing of female mating, reflecting the potentially different costs of female signalling and/or polyandry to both sexes. Spider sex pheromones are likely to be lipids or lipid soluble, may be closely related to primary metabolites, and are not necessarily species specific, although they can still assist with species recognition. Newer electrophysiological techniques coupled with chemical analyses assist with the identification of sex pheromone compounds. This provides opportunities for more targeted behavioural experimentation, perhaps with synthetic pheromones, and for theorising about the biosynthesis and evolution of chemical signals generally. Given the intriguing biology of spiders, and the critical role of chemical signals for spiders and many other animal taxa, a deeper understanding of spider sex pheromones should prove productive.     

Foraging and mate-finding in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae) under the risk of predation

Animal foraging and reproductive behaviour is influenced by other simultaneous demands such as predator avoidance. The trade-offs between these demands may depend on sex or mating experience. This study demonstrates that the olfactory-mediated foraging and mate-seeking behaviours in the silver Y moths, Autographa gamma , are affected by auditory cues mimicking their bat predators. Both males and females changed their foraging behaviour under simulated predation risk. Fewer moths reached the odour source following sound stimulation and the time to find the odour source increased by up to 250%. However, there were no significant differences between male and female ability to reach the plant odour source or the duration of the flight towards the source when stimulated with ultrasound. Hence females are not more cautious than males when observed in the same behavioural context. Risk-taking in males was independent of whether they were flying toward a flower odour or sex pheromones having equal attractive value. This indicates that the trade-off between olfactory and acoustic cues is independent the type of odour. Mated females were not as strongly affected by sound as non-mated, indicating that flower odours have a higher adaptive value for mated females, suggesting that some processes following mating experience influence the trade-off between flower odours and simulated bat sounds.     

Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

Turbidity affects association behaviour in male Poecilia latipinna

Male association preferences in a bisexual‐unisexual species complex were studied in clear and turbid environments. In south and central Texas, where the gynogenetic sexual‐parasite Poecilia formosa lives syntopically with Poecilia latipinna as its sexual host species, association times of P. latipinna males with conspecific sexual and heterospecific asexual females in clear and turbid water were measured sequentially. Turbidity had an influence on male mate association behaviour. Males spent less time with any kind of female stimulus in turbid water. There was no preference for conspecific sexual females, either in turbid water or under clear conditions. Also, origin of males and acclimatization to turbid water had no effect. How turbidity as a source of visual noise might affect communication among individuals and how this environmental factor might contribute to the stability of this sexual‐asexual mating complex in nature are discussed.     

Wake up and smell the conflict: odour signals in female competition

Odour signals used in competitive and aggressive interactions between males are well studied in the context of sexual selection. By contrast, relatively little is known about comparable signals used by females, despite current interest in the evolution of female ornaments and weaponry. Available evidence suggests that odour signals are important in competitive interactions between female mammals, with reductions or reversals of male-biased sexual dimorphism in signalling where female competition is intense. Scent marking is often associated with conflict between females over access to resources or reproductive opportunities. Female scent marks may therefore provide reliable signals of competitive ability that could be used both by competitors and potential mates. Consistent with this hypothesis, we report that aggressive behaviour of female house mice is correlated with the amount of major urinary protein (MUP) excreted in their urine, a polymorphic set of proteins that are used in scent mark signalling. Under semi-natural conditions, females with high MUP output are more likely to produce offspring sired by males that have high reproductive success, and less likely to produce offspring by multiple different sires, suggesting that females with strong MUP signals are monopolized by males of particularly high quality. We conclude that odour signals are worthy of more detailed investigation as mediators of female competition.     

Contact chemoreceptive sex recognition in the male cricket, Teleogryllus commodus

ABSTRACT. A behavioural continuum ranging from extreme aggression to overt sexual behaviour was used to measure male crickets' reactions to being touched in a standardized manner on their antennae with freshly severed antennae of male and female conspecifics. Sexually receptive males responded primarily with the aggressive song to male antennae and with the courtship song to female antennae. The ontogeny of the antennal effectiveness and the males' reaction was also determined. Because the ability of the antennae to elicit a reaction could be eliminated by treating them with chloroform, and since there were no morphological differences between male and female antennae, separate male and female sex recognition pheromones must be implicated. The chemo-tactile nature of these substances is indicated by the inability to obtain either behavioural or EAG responses to air carrying the odour of males or females.     

Sexual dimorphism of arrestment and gregariousness in the bed bug (Cimex lectularius) in response to cuticular extracts from nymphal exuviae

Releaser pheromones have direct behavioural effects to arrest, attract or disperse insects, whereas interactions within groups of social insects are often influenced by primer pheromones. The behaviour of insects displaying intermediate levels of sociality is largely unexplored in this context. In the present study, both the gregariousness and arrestment (settling near the odour source) of bed bugs Cimex lectularius L. (Hemiptera: Cimicidae) in response to conspecific exuvial extracts are described. Adult males are arrested on filter papers with extracts derived from exuviae of fifth‐instar nymphs. Adult females and nymphs display no significant evidence for such behaviour. Adults of both sexes show no preference for extracts of male versus female fifth‐instar exuviae. Arrestment of adult males does not occur on papers treated with fourth‐instar exuvial extracts. Because the insects are assayed behaviourally in groups, an index is calculated describing how far bugs are away from being located independently of one another, as a measure of gregariousness. Adult males have lower values for this index (i.e. locations are closer to independence). Adult females, nymph cohorts and mixed age groups all have higher values for this index, which tend to increase over time. Females exhibit a clear increasing dose‐dependent relationship for this index. It is concluded that the extracts of fifth‐instar nymphal exuvia arrest males on refuges that possess the odour source. However, gregariousness is induced in females, without evidence of a tendency to assemble near the odour source.     

Male traits influence odour‐based mate choice in the three‐spined stickleback

Female three‐spined sticklebacks Gasterosteus aculeatus preferred odour stemming from males with a nest over 'neutral' water and odour from males without any nest in an experimental flow channel. Males without any nest were preferred over females when either having the same major histocompatibility complex (MHC) class lI genotype or being unattractive in respect to MHC class II alleles.     

Friend or foe: behavioral responses to conspecifics in the northern scorpion, <Emphasis Type="Italic">Paruroctonus boreus</Emphasis> (Scorpionida: Vaejovidae)

We investigated the behavioral response of male Paruroctonus boreus to male and female pheromones of conspecifics. Adult male scorpions actively search for females during the mating season, but lack visual cues to direct their movement. Chemical cues (pheromones) are thought to be responsible for directing movement. In a series of laboratory experiments, in Y-mazes, males were exposed to areas that had contained males and females from the same population and areas that had contained conspecifics from different populations. Males spent significantly more time in areas that had contained females from the same population but did not show a preference for areas exposed to females from different populations. Males also significantly avoided areas exposed to males from the same population. Our results suggest that P. boreus has a sex-specific response to pheromones and that this response may be population-specific.     

Differential lifetime success and performance of native and non‐native Atlantic salmon examined under communal natural conditions

The lifetime success and performance characteristics of communally reared offspring of wild native Burrishoole (native), ranched native (ranched) and non‐native (non‐native) Atlantic salmon Salmo salar from the adjacent Owenmore River were compared. Non‐native 0+ year parr showed a substantial downstream migration, which was not shown by native and ranched parr. This appears to have been an active migration rather than competitive displacement and may reflect an adaptation to environmental or physiographic conditions within the Owenmore River catchment where the main nursery habitat is downstream of the spawning area. There were no differences between native and ranched in smolt output or adult return. Both of these measures, however, were significantly lower for the non‐native group. A greater proportion of the non‐native Atlantic salmon was taken in the coastal drift nets compared to the return to the Burrishoole system, probably as a result of the greater size of the non‐native fish. The overall lifetime success of the non‐native group, from fertilized egg to returning adult, was some 35% of native and ranched. The ranched group showed a significantly greater male parr maturity, a greater proportion of 1+ year smolts, and differences in sex ratio and timing of freshwater entry of returning adults compared to native, which may have fitness implications under specific conditions.     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.     

Shoaling and mate choice of wild-type Tanichthys albonubes in the presence of the red fluorescent transgenic conspecifics

Shoaling and sexual behaviour of wild-type male and female white cloud mountain minnow Tanichthys albonubes were measured in the presence of the red fluorescent transgenic conspecifics under laboratory conditions. Wild-type female test fish showed no significant preference, whereas wild-type male test fish preferred to be near a shoal of red transgenic fish rather than wild-type fish. When placed in a potentially reproductive context, wild-type males had a higher competitive ability over transgenic males; wild-type females spent more time with wild-type males in visually mediated experiments, but wild-type males performed more courtship displays towards transgenic females. These results suggest that the red body colouration does not appear to disturb signal communication between wild-type and transgenic T. albonubes in shoaling behaviour; transgenic males have no mating advantage over wild-type males, but the red body colouration of transgenic females may affect mate choice of wild-type males.     

Sex differences in pectoral muscles but not in pectoral fins in the three‐spined stickleback Gasterosteus aculeatus

The pectoral muscle index ( I _PM)( I _PM = 100 M _PM M⁻¹, where M _PM and M are the pectoral muscle and body masses, respectively) fin‐area and fin ray length were studied over a year in male and female three‐spined stickleback Gasterosteus aculeatus from a marine population (Öresund, Sweden) kept under simulated natural light and temperature conditions. A castration‐replacement experiment was used to test androgen effects on the I _PM, fin‐area and fin ray length. Non‐breeding males were castrated or sham‐operated in winter ( i.e . the fish had low levels of androgens). Castrated control and sham‐operated fish were implanted with empty Silastic capsules and castrated groups with capsules containing the androgens testosterone or 11‐ketoandrostenedione into the abdominal cavity. The experiment was terminated after 41 days, when the controls had matured. No morphological differences were found in pectoral fins between sexes during the year, except during the peak breeding season (May), where females showed larger fin‐area and longer fin ray in length compared to males. No effects of androgens treatment or of castration on pectoral fin‐area or fin ray length was observed. Breeding and non‐breeding males showed higher I _PM compared to females. The lower I _PM in females than in males could not be explained by the larger gonads in the former alone, as a sex difference in I _PM was still present after deduction of the ovaries from the female body mass. The I _PM was higher in sham‐operated compared to castrated fish. No effects of androgens treatment on I _PM was observed.     

Behavioural development of conspecific odour preferences in bank voles,Clethrionomys glareolus

Biological odours of conspecifics are known to have strong influences on behavioural interaction in bank voles Clethrionomys glareolus. This experiment tested two hypotheses. (1) Olfactory cues from familiar and unfamiliar mature opposite-sex conspecifics differ in their attractiveness to males and females, and their behavioural reactions change with age. (2) A genetically based mechanism is involved in female recognition of kin.In a two-choice preference test, prepubertal males and females were more attracted to familiar than to unfamiliar odours of opposite-sex conspecifics, as manifested by more time spent sniffing familiar voles. As the young reached sexual maturity they shifted their odour preferences. Mature males and females preferred the novel odour of unrelated opposite-sex conspecifics to that of relatives. The results of experiments testing the second hypothesis indicate that females use a genetically based mechanism to recognise their kin. Young and mature females were able to recognise the odour of their biological but socially unknown fathers, and showed the same pattern of behaviour as females in previous experiments.The possible biological functions of kin recognition in bank voles are discussed.     

Variability in diet and foraging behaviour between sexes and ploidy forms of the hybridogenetic Squalius alburnoides complex (Cyprinidae) in the Guadiana River basin, Portugal

Samples of the Squalius alburnoides complex, a hybridogenetic Iberian minnow composed of diploid, triploid and, less commonly, tetraploid forms, were collected from a tributary of the Ardila River (Guadiana River basin) between February 1999 and January 2001. Seasonal variation in diet was evident, and was probably linked to prey availability. Distinct foraging behaviours between ploidy forms were found towards several prey items, suggesting that diploid adult males fed mostly near the surface, whereas diploid adult females tended to feed near the bottom and submerged vegetation. Triploid females exhibited an intermediate foraging behaviour, although there was greater affinity towards diploid male feeding behaviour. Diploid males which, in contrast to diploid and triploid females, have non‐hybrid genomes in the Guadiana drainage, exhibited a higher specialization for food. Despite considerable dietary overlap, there appeared to be spatial segregation of feeding niches between the three forms, especially during dry periods when prey availability was lower, which may be a strategy for diminishing competition for food. Therefore, considering asexual generalist and specialist hypotheses, it appears that the different ploidy levels are generalist, opportunistic feeders that partition the resources when limited.     

Methyl-testosterone induces male-typical ventilatory behavior in response to putative steroidal pheromones in female round gobies (Neogobius melanostomus)

Male round gobies (Neogobius melanostomus) transiently increase their rate of ventilation when exposed to nanomolar concentrations of three putative steroidal pheromones (etiocholanolone, ETIO; estrone, E1; estradiol-glucuronide, E2-3g), whereas females exhibit this response only to ETIO. In this study we implanted females with Silastic capsules that were empty (Blank) or filled with methyl-testosterone (MT) to determine whether androgen induces a male-typical pattern of ventilatory response. As with untreated females in our previous studies, Blank females increased ventilation only in response to ETIO. In contrast, MT females tested 2-3 weeks postimplant responded not only to ETIO, but also to E1 and E2-3g. These results provide the first demonstration of an androgen inducing a male-typical behavioral response to a steroidal pheromone in adult female fish. Together with our previous findings of sexually isomorphic sensory responses to steroid odorants, the present results also suggest that a central action of androgen is responsible for sexually dimorphic behavioral responses to putative pheromonal steroids in Neogobius.