Effect of social condition on behavioral development during early adult phase in <Emphasis Type="Italic">Drosophila prolongata</Emphasis>

Behavioral changes during early development provide useful insights into the internal mechanisms that generate complex behavior expressed by mature individuals. At the same time, social conditions during early adult phase can influence behavior in later stages of development even in holometabolous insects. In this study, age-dependent changes in courtship behavior and the effect of social conditions were examined in a fruit fly, Drosophila prolongata. Younger males showed lower mating activity and simpler courtship behavior. Mating activity reached a maximum level by 5 days after eclosion, whereas expression of complex courtship behavior was not yet fully developed at that time, suggesting that they are controlled by different mechanisms. When two males were maintained in the same vial, not only mating activity but also courtship complexity was reduced, demonstrating for the first time that preceding social experience, not current social conditions, influenced the complexity of male courtship. The effect of social experience was completely erased by 1 day of isolation, however, showing that social experience did not suppress or promote behavioral development itself. Rather, these results suggest that the observed effect of social experience was a plastic response of males that reduced investment in courtship effort by anticipating increased male–male competition.     

Computer-manipulated stimuli as a research tool in Mozambique tilapia <Emphasis Type="Italic">Oreochromis mossambicus</Emphasis>

Multimodal communication is essential in social interactions in cichlid fish, including conspecifics’ recognition, agonistic interactions and courtship behaviour. Computer-manipulated image stimuli and sound playback offer powerful tools to assess the relative relevance of visual and acoustic stimuli in fish behavioural studies, but these techniques require validation for each taxon. The aim of the present study was to investigate whether Mozambique tilapia Oreochromis mossambicus responds to computer-manipulated visual stimuli and acoustic playback. Six experiments were conducted: computer animation playback, video playback, interaction with a mirror, presentation of a live male in a jar alone and combined with courting sound playback or with white noise playback. Individual agonistic interactions (lateral displays, up and down swimming, butting) and courting behaviours (tilting leading, digging) were tallied for each experiment. Our results suggest that non-interactive computer-manipulated visual stimuli is not a suitable tool in behavioural research with Mozambique tilapia. In contrast, interaction with a live male in a jar seems to remain the best visual research instrument inducing significant strong behavioural responses. Although none or only a few agonistic interactions were observed towards video playbacks or computer animations, such interactions significantly increased towards a male in jar and were modulated by courtship sound playback, suggesting the additional relevance of sound playback as a tool in behavioural research with Mozambique tilapia, including the study of multimodal signalling.     

The role of auditory stimuli in the courtship of Drosophila melanogaster

Simulated courtship song of male Drosophila melanogaster was played to males or females of this species. Upon receiving the song males increase their locomotor activity and start courting each other, whereas females reduce their locomotor activity. In wingless males the locomotor activity difference between the silent control and the experimental sound situation is much larger than in winged males, due to the inactivity of wingless males in the control situation. Males which had been kept singly up to the time of the experiment exhibit higher locomotor and sexual activity than group housed males. A second component of the male courtship song ‘sine song’ is described, together with experiments which investigate the sensory basis of the effect male courtship song has on males.     

Abrupt population changes along smooth environmental gradients

Populations often exhibit abrupt changes in abundance associated with a smooth, continuous change in some component of their environment, with the abruptness usually attributed to inter-specific interactions or physical extremes. This paper presents a spatially explicit single-species population model in which intra-specific interactions alone are responsible for such an abrupt change. The essential mechanism involves cooperation in both colonization (through enhanced recruitment near other individuals) and mortality (protection through a “safety-in-numbers” interaction). Large fluctuations in population density would likely be observable near the transition region.     

Court and spark: electric signals in the courtship and mating of gymnotoid fish

By mimicking tropical rainy season conditions in aquaria, we stimulated two species of gymnotoid electric fish, Eigenmannia virescens and Apteronotus leptorhynchus, to spawn in captivity. Their courtship activity, breeding behaviour and electric social communication were monitored in several groups over 2 years. Groups of both species established dominance hierarchies correlated with electric organ discharge frequency, aggressiveness and size. Spawning was preceded by several nights of courtship during which the male modulated its electric organ discharge to produce ‘chirps’. Continual bouts of chirping lasted for hours on evenings prior to spawning. These electrical signals play a significant role in courtship and spawning, as gravid E. virescens females could be stimulated to spawn by playing back into the tank a tape recording of male courtship chirps. In both species the chirp invovves a slight increase in frequency followed by a cessation of the dominant frequency. This suggests a common mode of signal production in these two different genera of fish. Chirps are short and abrupt during aggressive encounters, but assume a softer and more raspy quality during courtship.     

Population size,social behaviour,and dispersal in house mice: A quantitative investigation

An experimental study was conduced to examine some relationships between population size, social behaviour, and dispersal utilizing wild genotype Mus musculus in seminatural enclosures. Groups of male and female mice were used to simulate natural populations. The results indicate that male territoriality occurred significantly more often in populations of greater size but equal density, and these groups demonstrated a significantly greater rate of agonistic interactions per animal when compared to smaller populations. No significant differences were observed for other categories of social interaction. The percentage of animals emigrating and the population density following emigration were the same in small and large groups.     

Intraspecific interactions in a high‐density leopard population

Although less studied than interspecific interactions, interactions among members of the same species can influence space use and temporal activity. Using techniques commonly applied to the analysis of interspecific interactions—multispecies occupancy modeling and the analysis of temporal activity patterns—we studied intraspecific interactions within a high‐density population of Persian leopards (Panthera pardus saxicolor) in Tandoureh National Park, northeastern Iran. Using camera‐trap data, we investigated spatiotemporal interactions between male leopards, lone female leopards, and families (cubs/females with cubs). While we hypothesized that male and female leopards would display different temporal activity patterns, we did not predict spatial avoidance between these groups. We also predicted that leopard families would exhibit spatiotemporal avoidance from male leopards due to the risk of infanticide. Contrary to our expectations, we did not find any evidence for spatial or temporal avoidance between leopard families and adult male leopards. Male and lone female leopards exhibited positive pairwise co‐occurrence, consistent with reports of high overlap between male and female leopard home ranges. While a high level of overlap in temporal activity patterns was found between males/lone females and males/families, there was evidence for variation in the proportion of time each leopard group was active in particular periods of the diel cycle. Male leopards showed cathemeral activity, while lone females and families were more active during daylight hours. The application of these techniques to interactions within a species has improved understanding of the ecology and behavior of this endangered solitary carnivore.     

No Evidence for Activity Adjustment in Response to Increased Density in Daphnia magna

Increased population density may lead to a decrease in energy available for growth and reproduction via effects on the activity level of individuals. Whilst this may be of particular importance for organisms that compete for defendable resources and/or have a high frequency of social interactions, it is less obvious how individual activity should covary with population density when food resources are not defendable or direct interactions among individuals are negligible. Based on observations that there is a general negative relationship between population density and metabolism it has been suggested that organisms actively reduce activity under increased density to accommodate an expected decrease in food availability. However, in the absence of direct activity measurements the validity of this hypothesis is unclear. Here we test for such anticipatory adjustments of activity levels in the planktonic cladoceran Daphnia magna Straus, a filter feeder whose food resources are not defendable, meaning that density responses can be evaluated in the absence of direct interactions. We tested for changes in activity in response to two separate density cues, one being the direct physical and visual stimuli resulting from being in the vicinity of conspecifics (‘direct density experiment’), and the other being the detection of olfactory cues in their environment (‘olfactory cue experiment’). Ten genetically distinct clones were used to evaluate the potential for genetic variation in these responses. Our measures of activity were highly repeatable, and there was significant variation in activity among clones. Furthermore, this clonal variation was consistent in the ‘direct density’ and ‘olfactory cue’ experiments. The estimated evolvability of the trait (1.3–3.2%) was within the range typically observed in behavioural traits. However, there was no indication that the activity level of individuals respond to population density, either directly to actual density or to olfactory cues representing high density. In this case, the energetic influence of density on population dynamics is sufficiently described by effects on per capita resource availability.     

Density-Dependent Behavior of Shrews (Sorex araneus, S. caecutiens, and S. minutus) under Natural and Experimental Conditions

The behavior of three common shrew species was compared in a test of pairwise placement in a neutral territory (10 min) for various population densities in natural habitats and for animals kept isolated or in groups in open-air cages. The behavior of the animals captured in nature was regarded as the norm. We analyzed the total number of social contacts (as an index of social activity), as well as the frequencies of direct and ritualized aggressive interactions, rejection of contacts, and friendly interactions (as parameters characterizing the structure of interactions). Isolation of the shrews proved to decrease their social activity. The highest frequency of interactions was observed in the animals kept in groups in open-air cages. The level of sociality and changes in behavior related to differences in population density in the three shrew species can be more significant than the species-specific differences in their behavior.     

Involvement of different mesotocin (oxytocin homologue) populations in sexual and aggressive behaviours of the brown anole

The oxytocin (OT) family of neuropeptides are known to modulate social behaviours and anxiety in mammals and birds. We investigated cell numbers and neural activity, assessed as Fos induction, within magnocellular and parvocellular populations of neurons producing the OT homologue mesotocin (MT, Ile⁸-oxytocin). This was conducted within the male brown anole lizard, Anolis sagrei, following agonistic or courtship encounters with a conspecific. Both neurons colocalizing and not colocalizing corticotropin-releasing factor (CRF) were examined. Parvocellular neurons of the paraventricular nucleus exhibited a positive correlation between courtship frequency and Fos colocalization, regardless of whether they produce just MT or MT + CRF. Magnocellular populations showed only trends towards positive relationships with courtship and no cell populations showed aggression-related Fos induction. These findings are novel because they demonstrate the involvement of MT neurons in male social behaviour, especially in reptiles for whom the involvement of MT in social behaviour was previously unknown.     

Negative frequency-dependent selection is intensified at higher population densities in protist populations

Natural populations of free-living protists often exhibit high-levels of intraspecific diversity, yet this is puzzling as classic evolutionary theory predicts dominance by genotypes with high fitness, particularly in large populations where selection is efficient. Here, we test whether negative frequency-dependent selection (NFDS) plays a role in the maintenance of diversity in the marine flagellate Oxyrrhis marina using competition experiments between multiple pairs of strains. We observed strain-specific responses to frequency and density, but an overall signature of NFDS that was intensified at higher population densities. Because our strains were not selected a priori on the basis of particular traits expected to exhibit NFDS, these data represent a relatively unbiased estimate of the role for NFDS in maintaining diversity in protist populations. These findings could help to explain how bloom-forming plankton, which periodically achieve exceptionally high population densities, maintain substantial intraspecific diversity.     

Sex before or after blood feeding: Mating activities of Aedes aegypti males under conditions of different densities and female blood feeding opportunities

Blood feeding and mating are critical behaviors that regulate both mosquito population maintenance and disease transmission. However, our understanding of mosquito mating systems remains incomplete. One of the most critical issues is a lack of understanding regarding how and where males and females encounter one another. This study was performed to investigate changes in key mating behaviors of Ae. aegypti relative to female blood feeding opportunities, taking into account male density. We compared courtship latency and copulation activity between single and pooled males in a range of assays performed in the presence or absence of a blood source and after blood feeding. The time taken by grouped males to initiate courtship in the presence of a host was much shorter than that in single males. There was no significant difference in courtship latency between pooled and single males in the absence of a blood source or after blood feeding. At low male density, the presence of the host and blood meal ingestion provided better conditions for copulation. At high male density, however, copulation activity was decreased after blood feeding, but remained high regardless of the presence or absence of the host. In addition to providing insight into the mating ecology of Aedes aegypti, this study indicated that the presence of a blood source influences how males encounter and copulate with females. The observation that copulation activity decreases after blood feeding when males are numerous provides new avenues for improving mass release programs of sterile mosquitoes.     

Increased density and male-male interactions reduce male longevity in the medfly, Ceratitis capitata

We investigated the effect of adult density on male and female longevity and behavioural interactions in a mass-reared strain of the medfly. Male survival decreased significantly, and male-male behavioural interactions increased significantly, with increasing male density (males were kept at 1, 2, 6 and 10 flies per 285 cm²in pots of 5.5 cm in diameter and 12 cm high). No such effects were observed in females over the same density range. This suggests that male survival decreased in response to the deleterious effects of male-male behavioural interactions. At the highest density, courtship wing fanning directed towards other males decreased significantly over time. In addition, as either density or male-biased sex ratio increased, males were increasingly likely to be engaged in courtship wing fanning that was not directed towards females. These results suggest that high adult density may significantly reduce male reproductive success, by decreasing survival and mating success. The results highlight the potentially high survival costs of male-male interactions and indicate differences in the nature of reproductive costs in the two sexes.     

Courtship interactions stimulate rapid changes in GnRH synthesis in male ring doves

Many birds and mammals show changes in the hypothalamo-pituitary-gonadal (HPG) axis in response to social or sexual interactions between breeding partners. While alterations in GnRH neuronal activity play an important role in stimulating these changes, it remains unclear if acute behaviorally-induced alterations in GnRH release are accompanied by parallel changes in GnRH synthesis. To investigate this relationship, we examined changes in the activity of GnRH neurons in the brains of male ring doves following brief periods of courtship interactions with females. Such interactions have been previously shown to increase plasma LH in courting male doves at 24 h, but not at 1 h, after pairing with females. In the first study, males allowed to court females for 2 h had 60% more cells that showed immunocytochemical labeling for GnRH-I in the preoptic area (POA) of the hypothalamus than did control males that remained isolated from females. To determine whether an increase in GnRH gene expression preceded this increase in GnRH immunoreactivity in the POA, changes in the number of cells with detectable GnRH-I mRNA in the POA were measured by in situ hybridization following a 1 h period of courtship interactions with females. In this second study, courting males exhibited 40% more cells with GnRH-I in this region than did isolated control males. GnRH-immunoreactive neurons in two other diencephalic regions failed to show these courtship-induced changes. Plasma LH was not elevated after 1 or 2 h of courtship. These results demonstrate that the release of GnRH-I in the POA that is presumably responsible for courtship-induced pituitary and gonadal activation is accompanied by a rapid increase in GnRH synthesis that occurs before plasma LH levels increase. We suggest that this increase in GnRH synthesis is necessary to support the extended period of HPG axis activation that is seen in this species during the 5–10 day period of courtship and nest building activity.     

Impact of male alternative reproductive tactics on female costs of sexual conflict under variation in operational sex ratio and population density

Sexual conflict over mating rate is both pervasive and evolutionarily costly. For females, the lifetime reproductive fitness costs that arise through interactions with potential mates will be influenced by the frequency of such interactions, and the fitness cost of each interaction. Both of these factors are likely to be influenced by variation in operational sex ratio (OSR) and population density. Variation in OSR‐ and density‐dependent male alternative reproductive tactics (ARTs) may be particularly important if the fitness costs that females experience vary with the reproductive tactics that males express. Using a simple model, we consider several examples of OSR‐ and/or density‐dependent variation in male ARTs and the frequency of male–female interactions, and find that variation in the expression of male ARTs has the potential to augment or diminish the costs of frequent male interactions for females. Accurately documenting variation in the expression of male ARTs and associated female fitness costs will benefit future work in this area.     

Male swordtails court with an audience in mind

Females are usually considered to be the target of male courtship behaviour. In nature, however, social interactions rarely occur without other observers; thus, it is conceivable that some male courtship behaviours are directed not towards females, but rather towards male rivals. The northern swordtail, Xiphophorus birchmanni, is a freshwater fish found in high densities in natural streams. Males court by swimming close to and in parallel with the female, raising their large sail-like dorsal fin, and quivering briefly. Here, we show that females prefer males that display small dorsal fins to those with large ones, and that males are less aggressive to other males with large dorsal fins. Male swordtails also raise their dorsal fins more frequently when courting in the presence of other males. These results suggest that, despite female avoidance of large dorsal fins, males that raise their fin during courtship benefit by intimidating potential competitors; the intended receivers of this signal are thus males, not females. Intrasexual selection can therefore offset the forces of intersexual selection, even in a courtship display.     

Variable patterns of density-dependent survival in social bacteria

In numerous species of social animals and social microorganisms,fitness is positively dependent on population density, at leastin some environments and over some density ranges. This "Alleeeffect" is observed in the cooperative bacterium Myxococcusxanthus during multicellular fruiting body development, duringwhich the standard laboratory genotype sporulates less efficientlyat lower population densities and produces no spores below aminimum threshold density. Here we demonstrate significant quantitativevariation in Allee patterns among distinct natural isolatesof M. xanthus. Isolates with similar developmental performanceat intermediate population densities exhibit stark variationin performance at both very low and very high densities. Suchvariation has implications for evolutionary performance underfluctuating natural environments. It also suggests that distinctintraspecific populations of social animals and other socialmicrobes with different selective histories may vary in theeffects of density on social fitness.     

Parasite transmission in social interacting hosts: monogenean epidemics in guppies

Background

Infection incidence increases with the average number of contacts between susceptible and infected individuals. Contact rates are normally assumed to increase linearly with host density. However, social species seek out each other at low density and saturate their contact rates at high densities. Although predicting epidemic behaviour requires knowing how contact rates scale with host density, few empirical studies have investigated the effect of host density. Also, most theory assumes each host has an equal probability of transmitting parasites, even though individual parasite load and infection duration can vary. To our knowledge, the relative importance of characteristics of the primary infected host vs. the susceptible population has never been tested experimentally.

Methodology/Principal Findings

Here, we examine epidemics using a common ectoparasite, Gyrodactylus turnbulli infecting its guppy host (Poecilia reticulata). Hosts were maintained at different densities (3, 6, 12 and 24 fish in 40 L aquaria), and we monitored gyrodactylids both at a population and individual host level. Although parasite population size increased with host density, the probability of an epidemic did not. Epidemics were more likely when the primary infected fish had a high mean intensity and duration of infection. Epidemics only occurred if the primary infected host experienced more than 23 worm days. Female guppies contracted infections sooner than males, probably because females have a higher propensity for shoaling.

Conclusions/Significance

These findings suggest that in social hosts like guppies, the frequency of social contact largely governs disease epidemics independent of host density.     

Eavesdropping and signal matching in visual courtship displays of spiders

Eavesdropping on communication is widespread among animals, e.g. bystanders observing male-male contests, female mate choice copying and predator detection of prey cues. Some animals also exhibit signal matching, e.g. overlapping of competitors' acoustic signals in aggressive interactions. Fewer studies have examined male eavesdropping on conspecific courtship, although males could increase mating success by attending to others' behaviour and displaying whenever courtship is detected. In this study, we show that field-experienced male Schizocosa ocreata wolf spiders exhibit eavesdropping and signal matching when exposed to video playback of courting male conspecifics. Male spiders had longer bouts of interaction with a courting male stimulus, and more bouts of courtship signalling during and after the presence of a male on the video screen. Rates of courtship (leg tapping) displayed by individual focal males were correlated with the rates of the video exemplar to which they were exposed. These findings suggest male wolf spiders might gain information by eavesdropping on conspecific courtship and adjust performance to match that of rivals. This represents a novel finding, as these behaviours have previously been seen primarily among vertebrates.     

Modification of the response to high density conditions in the guppy, Poecilia reticulata (Peters)

An examination was made of some of the factors which influence the response to density in the guppy. Water from high density fish was given to fish populations kept at high and low densities. This induced high density behaviour in low density fish by increasing aggression and decreasing courtship. But giving 'fresh' water to high density fish failed to alleviate the response to density. 'High density water' slightly reduced the numbers of young found in low density groups. High density fish kept in the dark had more young than high density groups in the light; and the number of young found was decreased when 'high density water' was given to low density fish. The number of ovarian stages was decreased in the dark in high density and was not overridden by the type of water. Giving 'high density water' to low density fish decreased the number of oocytes stage III, but the interaction between water and 'light' was confounded by the total numbers of ovarian stages. Increasing the 'visual space' of high density fish with mirrors partially decreased aggression and increased courtship and number of young found; but had little affect upon the number of ovarian stages. Decreasing the number of physical contacts by keeping the fish chronically tranquillized reduced the activity of the fish. This reduced the aggression of the high density fish and also reduced the differences between the high and low density fish on the basis of their courtship scores, number of young found and number of stages in the ovary. Fish from differing population sizes were given a choice between varying fish densities. The fish from a 'normal' sized population and with an extended visual field, tended to make a choice conforming to the overall selections compared with fish from populations of extreme size and with a reduced visual field.