Social olfaction in male brown lemmings (Lemmus sibiricus = trimucronatus) and collared lemmings (Dicrostonyx groenlandicus): II. Discrimination of mated and unmated females

When tested in a Y-maze olfactometer, male brown and collared lemmings (Lemmus sibiricus = trimucronatus and Dicrostonyx groenlandicus) preferred the odor of unmated receptive females to the odor of females with which they had just copulated. Similarly, sexually satiated males preferred the odor of an unmated receptive female to that of a strange female that had recently copulated with another male. Sexually experienced males without recent copulatory experience also demonstrated this preference, but sexually naive males did not. Sexually satiated collared lemmings preferred the odor of bedding from a novel estrous female to bedding from the female with which they had just copulated even when the bedding was collected before mating occurred. These results suggest that discrimination between prior mates and unmated females may be based on individual recognition as well as recognition of subclasses of females (i.e., mated vs. unmated, familiar vs. unfamiliar).     

Olfactory discrimination of estrous condition by the male golden hamster (Mesocricetus auratus)

After repeated exposure to receptive and nonreceptive females, male golden hamsters were tested for olfactory preferences in a four-choice olfactometer. Males discriminated between the odor of anesthetized females in different stages of their estrous cycle when the airstreams carrying the stimulus odors were diluted. Previous failures to demonstrate such an ability were probably due to ceiling effects resulting from laboratory testing. Males preferred the odor of females on the day before receptivity (a day coincident with maximal scent marking by the female) and least preferred the odor of females on diestrus-1 (the day on which females attacked and chased males during pretesting encounters). Detection and quick response to an impending estrus would be especially important for males of a solitary and promiscuous species in which there is a first male mating advantage.     

Effect of anosmia on reproduction in male and female wolves (Canis lupus)

Anosmia was produced in two female and three male wolves by transection of the olfactory peduncle and was confirmed by their inability to detect meat, urine, feces, anal-gland secretions, and fish emulsion. All operated animals continued to investigate the environment with their noses, to interact normally with other pack members, and to feed at levels which maintained presurgical body weights. No effect was found on reproductive physiology (females: estradiol or progesterone concentrations, ovulation, pregnancy or parturition; males: testosterone, testicular recrudescence or sperm numbers, motility or maturation). One anosmic female became dominant and although she urine-marked with a flexed leg, the rate was lower than typical for dominant females and perhaps contributed to her failure to pair-bond with the dominant male. One anosmic male raised-leg-urinated while competing for pack dominance and when kenneled away from other males. Precopulatory, copulatory, and maternal behavior were observed for one anosmic female and appeared normal. However, neither male that was sexually naive before surgery showed interest in proestrous or estrous females. The possibility that secondary degeneration of brain regions mediating sexual behavior was responsible for the failure of these males to respond was not supported. Not only was the lack of male sexual response the only serious deficit following transection, but the male which was sexually experienced prior to surgery did copulate successfully during his second postoperative breeding season despite continued anosmia. Chemosensory priming from female urine during the protracted proestrous phase, as well as urinary and vaginal odors during estrus, appear to be critical for induction of full sexual potency in sexually naive males. The importance of urine and vaginal secretions in the sexual response of experienced males is uncertain.     

Sex differences in urinary odors produced by young laboratory rats (Rattus norvegicus)

Responses of adult female rats to male and female pup urine were examined in three experiments. Investigatory sniffing of male and female pup urine deposits by maternal rats was compared in a series of simultaneous choice tests given at 3-day intervals between Day 2 and Day 17 postpartum. Male urine was consistently preferred. Introduction of male, but not female, pup urine to the nest was also found to significantly elevate maternal licking of anogenital regions of pups. Thus, urine from pups of various ages contains sex-identifying odors that differentially elicit spontaneous maternal interest. The odor of male urine may provide a sufficient stimulus to account for the greater anogenital licking that males of this species normally receive. Nonmaternal, naive, adult females behaved like maternal rats, preferring male urine in the choice test. This indicates that the maternal condition of a dam is not necessary either for the olfactory discrimination or for the male odor preference.     

Sexual experience and preference for males or females in the female rat

The influence of sexual experience on preference behavior was investigated in adult female rats. In the first experiment, preference behavior for sexually active males versus estrous females was investigated in female rats who were experienced with mounting behavior. Preference of these mount experienced females was compared to preference of females who were naive in this respect. Mount experience with estrous females induced a female-directed preference in ovariectomized female rats, irrespective of whether the females were treated with OIL or with testosterone-propionate (500 micrograms TP, injected once 48 h prior to testing). Sexually naive, OIL-treated females did not show a preference for males or females, but TP induced a preference for a male. Individual differences in mount frequency were not correlated with preference behavior. In the second experiment, the influence of experience with feminine sexual responses on preference behavior was investigated. OIL-treated and TP-treated females oriented equally toward males and females when they had been given the opportunity to copulate with males prior to the preference tests. The results strongly suggest that prior sexual experience is an important determinant of preference behavior. The female's behavior during sexual interactions was, however, not predictive of later preference.     

Responsiveness to estrous chemostimuli in male rats (Rattus norvegicus) of different ages

Previous findings indicate that Norway rats (Rattus norvegicus) respond differentially to estrous chemostimuli in early infancy and that adult male sexual responsiveness to such chemostimuli depends on infantile experience with suckling odors (Fillion & Blass, in press). The development of behavioral responsiveness to estrous chemostimuli in male rats from late infancy to adulthood was examined in this study. Preweanling, adolescent, and adult males were observed for 20 min in the presence of an anesthetized estrous or an anesthetized diestrous female. This was, for adolescent and adult subjects, the first heterosexual encounter since weaning. Rats in each age-group responded differentially according to female state, but in distinct, age-related ways. Preweanling spent more time investigating the female's body if she was estrous. Adults expressed the same affinity but by more focused investigation directed perivaginally. Adolescents showed no female-directed bias but groomed themselves more if the female was estrous. These data identify developmental continuities and discontinuities in male rat behavior toward sexually receptive females.     

Modulation of male mouse sniff, attack, and mount behaviors by estrous cycle-dependent urinary cues

Modulation of male urogenital sniff, attack, and mount behaviors by urinary cues from female mice (SJL/J) was assessed in two experiments. Experiment 1 examined the efficacy of urine from ovariectomized and intact females in modulating the behavior of sexually experienced and inexperienced males vis-à-vis the control substance, water. It was found that urine from ovariectomized females showed no behavior-modulating properties, whereas urine from intact females decreased attack but increased sniff and mount behaviors in male mice. These findings are indicative of ovarian-dependent chemosignals contained in the urine of only intact females. Prior sexual experience had no effect on behavior of males when exposed to urine from ovariectomized females. Sniff and mount behaviors were, however, increased when sexually experienced males were exposed to urine from intact females. Using sexually inexperienced males, Experiment 2 assessed the chemosignal activity of urine from females in each of four estrous phases. It was determined that urine from proestrous and estrous females increased sniff and mount behaviors and decreased attack behavior. Urine from metaestrous females was shown to possess only sniff-inducing and mount-reducing chemosignal properties. Urine from females in diestrous lacked attack- and mount-modulating chemocues but may possess a sniff-inducing chemosignal.     

Life-span changes in the copulatory behavior of male rats (Rattus norvegicus)

The copulatory behavior of 5-, 10-, 15-, and 20-month-old sexually naive male rats (Long-Evans) was studied during 2-hr tests with receptive females. There was no apparent change in sexual arousal as measured by latency to initiate copulation across age, with subjects from all groups exhibiting comparable latencies to first mount and first mount with intromission. The numbers of ejaculations achieved were also similar across ages. Significant age differences were found for frequency of mounts, with 20-month-olds having the highest mean frequency. The persistent mounting by older males appeared to account for significant group differences in interintromission interval and ejaculation latency. It is suggested that motor deficits may impair the ability of older males to achieve intromission, increasing the number of mount bouts as well as the number of mounts per bout, thus extending the length of each copulatory series.     

Scent marking by male dwarf hamsters (Phodopus sungorus campbelli) in response to conspecific odors

Male hamsters (Phodopus sungorus campbelli) scent marked at greater frequencies in the presence of conspecific odor, either male or female, than in clean areas. Marking rates were not different between male and female odor conditions. In contrast, the pattern of investigation in response to conspecific odor did differ between the male and the female odor conditions. When male odors were used as stimuli, investigative sniffing increased at the scent marking sites but when female odors were used as stimuli investigative sniffing increased throughout the arena. Thus, male dwarf hamsters direct their scent marks at both male and female conspecifics, discriminate between male and female odor, and investigate in a different manner areas scent marked by males and females.     

Avoidance of relatively aggressive male Japanese Quail (Coturnix japonica) by sexually experienced conspecific females

Sexually experienced female Japanese quail (Coturnix japonica) that are offered a choice between 2 conspecific males previously observed engaging in an aggressive encounter prefer to affiliate with the less aggressive male. The authors determined whether this apparent preference for less aggressive males results from females approaching less aggressive individuals or avoiding more aggressive individuals. The authors found that females that had seen 2 males fight before choosing, in counterbalanced order, between each of them and a neutral stimulus were indifferent to less aggressive males but avoided more aggressive males. The results are consistent with the view that in species in which male courtship and mating are potentially harmful to females, females keep away from relatively aggressive males in order to avoid the physical punishment that can result from contact with them.     

Social chemosignals in five Belontiidae (Pisces) species

Approach behaviors toward conspecific chemical stimuli of the opposite sex were examined in five Belontiidae species: Betta splendens, Macropodus opercularis, Colisa labiosa, C. lalia, and Trichogaster trichopterus. Approach was measured by (a) preference for section 1 of a three-section tank, which contained a vertical tube that introduced the stimulus water, and (b) occupancy of the tube. Experiments 1A and 1B showed that (a) approach behaviors were displayed by the isolated male Betta only to ripe-female stimulus water, (b) group-housed males of the remaining four species were not attracted to female-conditioned water, and (c) socially isolated males of these four species preferred section 1 during presentation of either ripe- or nonripe-female-inhabited water but occupied the tube only during exposure to ripe-female-conditioned water. The findings of Experiment 2A were that (a) the female Betta, regarless of physiological state, showed approach behaviors to male-inhabited water and (b) only ripe females of the remaining species indicated a preference for section 1 during male-water exposure but performed no tube entries. Results of Experiment 2B indicated that social isolation of the females, especially ripe females, facilitated their approach behaviors.     

Unconditioned sexual incentive motivation in the male Norway rat (Rattus norvegicus)

Sexual incentive motivation was evaluated in a procedure consisting of a large open field where incentive animals were confined behind wire mesh openings. When sexually inexperienced male rats (Rattus norvegicus) were exposed to the receptive female-male incentives, they spent more time close to the female. If the incentives were receptive female-nonreceptive female, the receptive female was preferred. However, when the alternatives were nonreceptive female-male, no preference was obtained. Castration abolished preference for the receptive female, and treatment with testosterone propionate restored it. Estradiol plus oil is as efficient as estradiol plus progesterone for giving the ovariectomized female incentive properties. The living female can be replaced with female odor. Sexual experience did not have any long-term effects on the female's incentive value, but immediately preceding limited sexual activity enhanced it     

Odor cue determinants of urine marking in male rats (Rattus norvegicus)

This study examined the spatial distribution of urine marking by sexually inexperienced male rats under six odor conditions: minimum odor (baseline), conspecific male urine, conspecific female urine, an animal's own urine, male gerbil urine, and amyl acetate. The gerbil urine and amyl acetate conditions were included to test the theory that marking, rather than being elicited by and directed as a display to appropriate conspecifics, occurred to the introduction of any novel or disruptive odor. The subject's own urine was used to test the theory that the animal marked to maintain an optimum level of its own odor in the environment. The results showed that marking was directed predominantly at objects in the environment over which the animal could readily pass. All odor conditions except "own odor" sharply increased marking over baseline levels. Marking decreased with repeated exposure to particular stimuli. The results strongly supported the theory that urine marking, at least in sexually inexperienced male rats, is produced by novel odor stimuli introduced into a familiar environment.     

Sex differences in investigatory and grooming behaviors of laboratory rats (Rattus norvegicus) following exposure to novelty

Prior research suggested that during exposure to novel stimuli, rodent investigation and self-grooming behaviors may be sexually dimorphic and interact with ambient illumination. To test this notion we compared the behavior of adult male and female groups of Long-Evans hooded rats in normal room lighting (860 lx) and in very dim, red light (0.2 lx) following exposure to a novel juvenile conspecific. Illuminance level had little or no effect, but investigatory and subsequent self-grooming behaviors of males were substantially greater than those of females, and females engaged in greater ambulatory activity than did males. In a second experiment adult males and females were exposed to a novel inanimate object. No reliable sex differences were observed. We conclude that social novelty, as provided by exposure to a juvenile conspecific, stimulates greater investigation and postinvestigatory self-grooming than exposure to a novel inanimate object and that exposure to novel conspecifics presents a useful method for the investigation of sex differences, gonadal hormone effects, and interactions of hormones with neurotransmitter systems governing motor control systems.     

Effects of pre- and postpubertal social experience on sexual and social behavior in male and female brown lemmings (Lemmus sibiricus)

The effects of prepubertal social contact and postpubertal mating experience on sexual behavior were examined in male and female brown lemmings (Lemmus sibiricus). Subjects of both sexes were either isolated at weaning (18 days) or then housed with siblings for another 10 days before being isolated. Before testing for sexual behavior at the age of 75 days, half of each group of males received a series of exposures to estrous females and half of each group of females was housed with a stud male. The results of mating tests indicated that whereas prior sexual experience facilitated subsequent copulatory behavior in both sexes, prepubertal interactions subsequently facilitated contact social and sexual behavior in males but did not increase the behavioral scores of females, which engaged in high levels of sexual behavior regardless of prepubertal experience. The effect of prior copulatory activity on male sexual performance could not be attributed to increasing age (i.e., maturational processes) because the behavior of males tested once but at different ages did not differ appreciably. Prepubertal social interaction appeared to predispose males to benefit from sexual experience later in life.     

Mechanisms and evolution of synchronous chorusing: emergent properties and adaptive functions in Neoconocephalus katydids (Orthoptera: Tettigoniidae)

Synchronous interactions arise in various animal species that rhythmically broadcast acoustic, vibratory, and visual signals. These interactions are characterized by a coincidence in both rate and phase of the rhythms of neighboring signalers. Theory predicts several ways in which synchronized rhythms may specifically benefit the interacting signalers. However, synchrony may also arise as an emergent property, a default phenomenon that is neither preferred by conspecific receivers evaluating the signals nor advantageous to the signalers themselves. Here, we examine several well-studied cases of acoustic synchrony in Neoconocephalus katydids (Orthoptera: Tettigoniidae), a New World genus wherein males broadcast loud advertisement songs. We report that call synchrony found in N. spiza and N. nebrascensis results from two rather different mechanisms of rhythm adjustment. Moreover, synchrony in the former species appears to represent an incidental byproduct of signal competition between evenly matched males, whereas in the latter species synchrony functions as a specific adaptation in which cooperating males ensure that critical call features can be perceived by females. We discuss the separate evolutionary trajectories that may have led to similar outcomes, synchronous chorusing by advertising males, in these closely related species.     

Attack and defense in conspecific fighting in tree shrews (Tupaia belangeri)

Principles of conspecific defense have been analyzed for rodents, in which specific target sites for biting by attackers on defenders serve as an important determinant of the actions involved in both attacker and defender behavior. In an effort to determine the generality of these principles, attack and defensive behaviors and target sites for biting attack were evaluated in a nonrodent species, the tree shrew (Tupaia belangeri). Brief daily and repeated conspecific dyadic encounters between adult, socially experienced males (dominants, attackers), and adult, socially naive males (subordinates, defenders) that had been transferred into the territory of the dominants, produced a polarization of attack and defense. The dominant males showed chase, chase attack, jump attack, and biting behaviors, while the subordinates displayed flight and freezing. The vast majority of bites, as well as wounds and bruises, were on the subordinates’ backs. These patterns are very similar to those previously found in rats and mice and suggest that the organization of fighting, with targets of biting (or other painful) attack serving as an important determinant of both attacker (dominant) and defender (subordinate) behavior, may show considerable generality across nonrodent as well as rodent species. Although relatively few wounds were found after 28 days of repeated and daily encounters, the subordinate tree shrews show a variety of behavioral, neuroendocrine, and central nervous changes, indicating that they are stressed by these encounters per se. Aggr. Behav. 27:139–148, 2001. © 2001 Wiley‐Liss, Inc.     

Effects of fluprazine (DU 27716) upon aggressive and sexual behavior of testosterone-treated female rats

Female rats, chronically treated with Testosterone Propionate (TP), were injected with Fluprazine (DU 27716) or saline and tested for social aggression, masculine and feminine sexual behavior. Fluprazine-treated females were less aggressive than saline-treated females, as indicated by a shift from offensive to less offensive parameters of aggression. At the same time, mounting in Fluprazine-treated females was almost totally abolished, both in aggression tests and in tests for sexual behavior. Feminine sexual responses increased during aggressive encounters but were slightly inhibited when females were confronted with sexually active males. Females treated with Fluprazine and tested for mounting with a receptive female showed a substantial increase of offensive aggression directed at the receptive stimulus female. It is concluded that Fluprazine does not selectively inhibit offensive aggression in TP-treated female rats.     

Effects of selective breeding for high and low aggressiveness and of fighting experience on odor discrimination in mice

The present study was designed to determine whether male and female mice, selectively bred for high (TA) and low (TNA) aggressiveness, react in different ways when exposed to odors from males of the same strains. The effects of losing or winning fights on odor preferences in the TA males were also studied. The low aggressive TNA males were found to avoid areas soiled by the highly aggressive TA males. The TA males, on the other hand, were most interested in the areas soiled by other highly aggressive TA males. Along with decreasing aggressiveness, losing fights radically changed the odor preferences of the TA males, the soiled bedding from winner TA males now having a strong aversive effect. TNA females preferred the odors of TA males, while the TA females preferred the odors of TNA males. The genotype of the receiver of odor cues is important for the preference choices; this effect can, however, be changed through experience.     

Male hamster preference for odors of female hamster vaginal discharges: studies of experiential and hormonal determinants.

Male hamsters approach sources of odors from female hamster vaginal discharges and spend significantly more time around these odor sources than around control locations in the test box. This preference for female hamster vaginal odors appears in sexually inexperienced as well as experienced males, even in individuals isolated from females since the time of weaning. Castration significantly reduces the sex odor preference, and treatment with testosterone propionate partially restores it.