Recognition of competitors by male golden hamsters

Golden hamsters, like many animals, form dominant/subordinate relationships after aggressive encounters. We examined whether behavioral responses by males that won or lost fights would differ toward familiar and unfamiliar male stimulus animals. In Experiment 1, male winners or losers of fights explored an arena containing a confined stimulus animal that was either familiar or novel and had either won or lost a fight. Compared to dominant males, losers spent less time in proximity to stimulus males and investigated them less. Losers also displayed higher levels of stretch-attend postures (indicative of risk assessment) than winners, and they showed more escape and locomotion in response to familiar winners than to unfamiliar winners, indicating recognition of the male that they had lost to. In Experiment 2, losers scent marked less to the odors of a familiar winner than to those of an unfamiliar winner. Thus, male hamsters appear to use familiarity with a former opponent's odors to adaptively regulate their responses to variations in social threat.     

A role for the hamster's flank gland in heterosexual communication

It is commonly assumed that the male hamsters' flank glands subserve an agonistic role with other males as the intended recipients of the olfactory message. Because of little evidence to support such a relationship, two studies investigated whether females might selectively respond to male flank odors. In a free choice situation females spent more time in proximity with glandular rubbings obtained from intact males than those obtained from castrates or a clean control. Moreover, these sexually receptive females remained longer near odors from familiar as opposed to novel males. The second experiment demonstrated that receptive females spent more time in proximity with flank odors of dominant males as opposed to their subordinate partners or to a clean control. Females can use flank gland odors to discriminate among males as a function of their hormonal state, familiarity and dominance. If this occurs in the wild, such odors could provide cues which would facilitate intersexual selection.     

A lack of avoidance of flank gland secretions by male Syrian hamsters

It is commonly assumed that sexually dimorphic cutaneous glands are important in supporting inter-male aggression in rodents. Such an intimidation role has been hypothesized as the function of the hamsters' flank gland. Two experiments failed to find avoidance of male flank odors. In one, subordinate males entered and spent comparable amounts of time in clean areas and areas scented with the dominant's flank secretion. In the other study, males were found to respond differentially to flank odors obtained from the subject, a familiar and a novel male. Although the males seemed to have habituated to the odor of the familiar male, there was no suggestion of avoidance of any of these odors. It may be that the males' flank odor is relevant not to other males but to females.     

Social experience, behavior, and stress in guinea pigs

This study examined the effect of varying rearing and testing conditions on guinea pig aggression, courting behavior, endocrine responses and body weight. Pairs of 7-8-month-old males were placed in chronic confrontations for 6-50 days in 2 m2 enclosures. Social behavior was recorded with a total of 882 h observation time. Body weight as well as plasma glucocorticoid, testosterone and norepinephrine titers were determined for each male 20 h before, and 4, 52 and 124 h after, the onset of the chronic encounters. Three experiments were conducted: in Experiment I, 7 pairs of males, each male raised singly with one female (FRM), were confronted in the presence of an unfamiliar female, in Experiment II, 6 pairs of FRM were confronted with no female present, and in Experiment III, 7 pairs of males which were raised in different large colonies were confronted in the presence of an unfamiliar female. In Experiment II and III low levels of aggression, no distinct endocrine changes and no indications of physical injury occurred in winners or losers, whereas in Experiment I high levels of aggression and courting behavior, extreme increases in glucocorticoid titers and distinct decreases in body weights were found in both males. Losers, however, were affected to a much greater extent than winners. These findings suggest that in guinea pigs a causal relationship exists between social rearing conditions, behavior as adults and degree of social stress in chronic encounters.     

Weak winner effect in a less aggressive mammal: correlations with corticosterone but not testosterone

We investigated the existence of the "winner effect" (winning an aggressive encounter following previous victories) and an associated rise in testosterone (T) in the white-footed mouse (Peromyscus leucopus) which generally display low levels of aggression and territoriality. We compared the effect of previously winning three, two, one, or zero resident-intruder encounters on the likelihood of winning a subsequent aggressive encounter. Although 50% of males were removed during training because of peaceful encounters, the winner effect was weak and not significant. We hypothesize that territoriality/aggression may be associated with the strength of the winner effect and discuss whether the slight winner effect exhibited by P. leucopus may become significant when population densities increase and males become more territorial. There was also no associated change in T with winning; however, corticosterone (Cort) changed with experience as winners had low Cort levels compared to losers and controls. Furthermore, low Cort levels in winners were associated with quicker attack latencies. These results contrast with findings of a significant winner effect and increase in T in males of the highly territorial and aggressive California mouse (Peromyscus californicus) using an identical methodology. California mice also attacked their opponents at more caudal regions of the body compared to white-footed mice that attacked their opponents at more rostral regions of the body, possibly related to different levels/types of aggression expressed by the two species.     

Mother and sibling discrimination at a distance by three- to seven-day-old lambs

This study examined some aspects of the developmental phase of social bonding in newborn lambs. In Experiment 1, 3-day-old Merino lambs (singles and twins) were tested in a 2-choice situation for mother discrimination at a distance. In Experiment 2, the ability of twins to discriminate their siblings at a distance was studied 3 and 7 days after birth. When released at 9 m from two postparturient ewes after a latency of 30 s, which allowed the lambs to hear and to see both animals, most lambs chose to run and to stay with their own mothers before the end of the 2-min test (Experiment 1). There was no difference between singles and twins, nor between males and females in their distant discriminative ability. However, twins were initially undecided, slower to reach their mothers, and spent significantly less time with them. When the same testing procedure was used for sibling discrimination, most 3-day-old twins went to a lamb, but the proportion of lambs reaching the sibling first did not differ significantly from random, and the time spent near either lamb was similar (Experiment 2). During these tests, female lambs performed better than males. When tested 7 days after birth, all twins reached a lamb and a high proportion of them chose to go to the sibling first. They also spent most of the time near it. These results demonstrate that mother discrimination from a distance is well established in 3-day-old lambs and sibling discrimination takes longer to develop but has occurred by 7 days of age.     

Shedding facilitates exposure learning in the garter snake (Thamnophis sirtalis)

Young garter snakes were exposed to a specific chemical odor for five consecutive days. Following training, the snakes were individually tested for their preferences for the training odor vs. an unfamiliar odor. Snakes that were trained and tested prior to — or following — their initial shedding displayed no selective responsiveness towards the odors. However, nine snakes which shed for the first time during the training period spent significantly more time near the training odor as compared to the novel odor during subsequent testing. It was concluded that endocrine changes which initiate and accompany shedding may account for the preference for the familiar odor shown by the snakes which shed during training. The possible adaptive value of such exposure learning was also discussed.     

Morphine treatment and reproductive condition alter olfactory preferences for pup and adult male odors in female rats

Administration of morphine sulfate (MS) to pregnancy-terminated and postpartum lactating female rats disrupts both maternal behavior and postpartum aggression. Since the display of these behaviors may be heavily dependent on olfactory cues provided by the stimulus animals (rat pups and adult male rats, respectively), we examined whether MS was affecting the perception of the olfactory stimuli, and whether olfactory perception was modified by reproductive condition. In Experiment 1, lactating rats had their pups removed and were injected with MS (5.0 mg/kg, sc.) or saline. 60 min later they were placed into a two-choice apparatus, one side of which contained bedding soiled by neonates and the other clean bedding. Time spent on each side was recorded for a total of 5 min (300 s; chance = 150 s). Saline-treated mothers spent significantly more time on the pup-odor side, whereas MS-treated females spent significantly less. In Experiment 2, lactating females were treated with MS or saline and exposed to male odors (soiled bedding). MS significantly increased time spent on the side with male odors; when treated with saline, time spent was significantly reduced. Thus, in lactating rats. MS creates an aversion for pup odors while reducing the female's normal aversion toward male odors. In Experiment 3, ovariectomized (ovx) virgin females expressed neither an aversion nor a preference for the odor of pups following saline administration. After MS treatment, however, the virgins showed a distinct preference for pup odors. When exposed to male odors in Experiment 4 ovx virgins showed a marked preference for male odors after MS treatment, and neither a preference nor an aversion after saline. Experiment 5 examined pup odor preferences in intact virgins, early (Day 7), middle (Day 14), late-pregnant (Day 21), and prepartum (Day 22) rats. The pup odor preferences of virgin, Day 7, Day 14, and Day 21 pregnant rats were not different and generally were at chance levels. Day 22 pregnant females exhibited a marked preference for pup odors compared to chance levels, as well as compared to the other four groups. These findings suggest that opiates and endogenous opioids may regulate olfactory preferences and that alterations in this system may underlie normal behavioral changes toward conspecifics prepartum as well as during lactation.     

Female discrimination of male odors correlated with male genotype at the T locus: A response to T-locus or H-2-locus variability?

Female house mice (Mus musculus), derived from several populations of wild-caught mice, were tested for their ability to discriminate between males whose genotype at the T locus was +/+ and those whose genotype was +/t, using odor cues alone. Females spent more time near the odors or +/+ males than near the odors of +/t males. This preference was independent of the T-locus genotype of the female and the particular type of t allele carried by either the male or the female. A female's preference, however, did appear to be related to the genotype of her parents. Females with one +/t parent were more likely to prefer +/t males than were females whose parents were both +/+. In a second experiment 18 females were tested with odors from soiled bedding of recombinant males whose genotype varied at the T locus but who were similar at the H-2 locus. As a control, these 18 females were also tested with bedding of wild-derived +/+ and +/t^{w semilethal} males. Females tested with recombinant males preferred odors of males not carrying lethal t alleles over those of males carrying two lethal t alleles, indicating that T-locus variability, not H-2-locus variability, is responsible for odor differences between +/+ and +/t males. Female responses to odors of recombinanat males did not differ from those to odors of +/+ and +/t^{w semilethal} males. Responses of mice to odor differences associated with T-locus variability may have evolved independently of responses to odor variability associated with the H-2 locus.This work was supported by NIH Grant HD 15997-01.     

Effect of sex steroids and coital experience on ferrets' preference for the smell,sight and sound of conspecifics

Previous research showed that ferrets of both sexes recognize potential opposite-sex mates on the basis of volatile body odors. We compared the ability of estrogen and androgen treatments to activate a preference in gonadectomized male and female ferrets for distal cues (volatile body odors alone or volatile odors+sight+sounds) from male versus female stimulus ferrets using an airtight Y-maze and a "stimulus proximity" or a "discrete trials" testing paradigm. Sexually naive, gonadectomized male and female ferrets that received either testosterone propionate (TP) or estradiol benzoate (EB) spent equal time in proximity to goal boxes that provided either volatile odors alone or odors+sight+sounds of male and female stimulus animals. After they received coital experience, male and female subjects (treated with either EB or TP) showed a significant preference for both types of opposite-sex stimuli. When discrete trials tests were given to these ferrets prior to receiving coital experience, EB-treated females preferred to approach odor only cues, as well as odors+sight+sounds of stimulus males, and this preference was further strengthened after coital experience. Sexually naive, TP-treated males preferred to approach volatile odor cues from stimulus females; however, these animals showed an equal preference for odors+sight+sounds of stimulus females and males. Again, after coital experience, males' preference for both sets of cues from stimulus females was significantly enhanced. Thus, in sexually naive ferrets, discrete trials, but not stimulus proximity tests, revealed a preference for distal cues (body odors with or without concurrent sight and sounds) from opposite-sex conspecifics in subjects of both sexes. Coital experience significantly enhanced these preferences for heterosexual distal cues under both testing paradigms.     

Olfactory alliesthesia in human neonates: Prandial state and stimulus familiarity modulate facial and autonomic responses to milk odors

This study examines the effects of a shift in the motivational state (from hunger to satiety) of human neonates on their behavioral and autonomic responsiveness to artificial and food‐related odors as a function of stimulus familiarity. In Experiment 1, videotaped facial movements and autonomic (respiration rate: RR, heart rate: HR) responses to five olfactory stimuli (familiar regular formula, unfamiliar regular formula, protein hydrolysate formula, vanillin, control) are recorded in 3‐day‐old neonates (n = 14) during episodes of irregular sleep. The infants are tested on average 50 min. before and after bottle feeding. RR discriminates the odor stimuli from the control stimulus, indicating clear olfactory detection. Furthermore, neonates react with higher HR change only when exposed to their familiar formula milk during the postprandial condition. The measurement of facial movements with the Baby‐Facial Action Coding System indicates that disgust and aversive actions are more often evoked by the odor of regular formulas (familiar or unfamiliar) than by the other olfactory stimuli during the postprandial condition. In Experiment 2, untrained adult observers, presented with the videotapes of the infants' facial responses to the odors, are able to decode differential hedonic signals from the sender faces as a function of the infants' motivational states. The present findings are in line with the concept of olfactory alliesthesia as defined in adults. © 1999 John Wiley & Sons, Inc. Dev Psychobiol 35: 3–14, 1999     

Effects of odor familiarity on the development of systematic exploration in the spiny mouse, Acomys cahirinus

Developmental changes in patterns of exploration by infant spiny mice, Acomys cahirinus are described. These were investigated using an unbaited radial maze and under three odor conditions; that is, animals were tested in the presence of either familiar (own) odors, unfamiliar conspecific odors, or no odors (washed floor). Two hypotheses were tested. The first was based on the results of a pilot study, and was that adult animals would explore the radial maze in a systematic and predictable fashion tending to move from one arm to the next sequentially. It was hypothesized that infants would adopt this sequential strategy only gradually, as they matured. The second hypothesis was that such patterns of exploration would depend upon the olfactory environment; the presence of familiar odors might facilitate systematic patterns of exploration in very young Acomys. The results showed that both adults and juveniles were less likely to move sequentially when tested with no conspecific odor present; sequential patterns of movement were most likely in the presence of unfamiliar odor, however. The only significant change with age in infants was found for animals tested with unfamiliar odors; these animals showed a dramatic increase in sequential behavior between 3 and 7 days of age. Two additional experiments are reported, which investigated the preferences of infant Acomys for unfamiliar conspecific odors, and it was found that very young (about 3 days) animals exhibit a preference for odors derived from unfamiliar conspecific litters, even when tested in the physical presence of their own parents. The results are discussed with reference to the use of olfactory information as directional cues for animals exploring the radial maze.     

Amphetamine reverses learning deficits in 6-hydroxydopamine-treated rat pups

At 5 days of age, rat pups were treated with a combination of desmethylimipramine (DMI) and 6-hydroxydopamine (6-OHDA) to selectively deplete brain dopamine (DA) or with vehicle (saline) control solutions. Two days later, all animals received conditioning to a novel odor by pairing the odor with intraoral milk. When the odor was anise, treated pups spent less time near the conditioned stimulus than did controls, but there were no 6-OHDA effects when the stimulus was a lemon odor (Experiment I). The difference in performance between the treated and control animals was not attributable to alterations in activating effects of the reinforcer (Experiment I), changes in olfactory sensitivity or olfactory preference (Experiment II), or sensitization to the stimulus (Experiment III). In Experiment IV, animals received d-amphetamine sulfate (0.5 mg/kg) prior to conditioning, testing, or both conditioning and testing. Amphetamine treatment before conditioning produced an improvement in performance in animals previously treated with 6-OHDA/DMI, but it impaired performance in controls, regardless of the time of injection. The results indicate a role of brain DA in learning in young rats.     

A competitive effect of androgen signaling on male mouse attraction to volatile female mouse odors

Olfaction plays an important role in animal communication. We hypothesized that males recognize the attractive volatile odors attributed to female reproductive ability. We measured the period during which a male mouse spent sniffing volatile odors from a sham-operated female mouse or an ovariectomized mouse without visual or tactile contact. Intact male mice spent more time sniffing volatile odors from proestrous, estrous or metestrous females than from ovariectomized females. There was no difference in castrated male mice. To investigate the involvement of sexual hormone in this behavior, castrated male mice were treated with 17 alpha-estradiol (E), dihydrotestosterone (DHT), or both. E-treatment did not affect sniffing behavior. Regardless of the estrous stages, DHT-treated castrated males spent less time sniffing the volatile odors from sham-operated than from ovariectomized female mice. Both E- and DHT-treated castrated males spent less time sniffing the volatile odors from proestrous or estrous females than from ovariectomized females. These results suggest that neither androgen nor estrogen is sufficient for reproducing male attraction to volatile female mouse odors, and that androgen signaling has a competitive effect against the attraction.     

Responses of male hamsters to the ear gland secretions of conspecifics

The behavioral responses of sexually experienced males toward anesthetized conspecifics were examined. Males directed more mounts and spent significantly more time investigating the ears and genital region of castrated males and diestrous females than intact males. Castrated males were treated in a similar manner to females, suggesting that the absence of an androgen-dependent factor(s) renders an animal more sexually attractive. Males were also administered paired preference tests in which they investigated the odors from the ear glands of diestrous females, castrated males, and intact males. Subjects spent significantly more time investigating the odors originating from ear glands of females and gonadectomized males than intact males. There was no significant difference in the amount of time males spent with the ears of castrated males and diestrous females. These experiments demonstrate that sexually experienced male hamsters can utilize odors associated with the ears to discriminate the gender of conspecifics.     

Lipopolysaccharide facilitates partner preference behaviors in female prairie voles

Exposure to proinflammatory cytokines (e.g., IL-1beta) or lipopolysaccharide (LPS) produces an acute activation of the immune response and results in a repertoire of behavioral patterns collectively termed sickness behaviors. Although nonspecific responses to pathogenic infection have traditionally been viewed as maladaptive effects of infection, sickness behaviors may have significant, adaptive value for the host. One set of adaptive behaviors affected by infection among mammals and birds is mate choice. In Experiment 1, female prairie voles exhibited the expected increase in blood corticosterone concentrations in response to a 0.1 cc i.p. LPS injection (50 microg), indicating activation of the endocrine system. A separate cohort of females was injected with LPS or saline and paired for 6 h with a novel, previously unpaired male. Following the cohabitation period, LPS-injected females spent significantly more time (p < 0.05) with the familiar partner when given a choice between familiar and unfamiliar males in a three-chamber apparatus designed to test partner preferences. Saline-injected females spent significantly more time with the unfamiliar male. In Experiment 2, males injected with LPS or saline spent equal amounts of time with familiar and unfamiliar females following a 6 h cohabitation with a naive female, and therefore, did not exhibit preferences. From a proximate perspective, this study provides evidence that sickness behaviors influence female, but not male, partner preference in prairie voles.     

Attraction of beagles to conspecific urine, vaginal and anal sac secretion odors

Attraction of male and female Beagles to conspecific urine, vaginal and anal sac secretion odors was examined in four experiments. Males spent more relative time investigating female urine odors than odors of vaginal or anal sac secretions. Sexually experienced males, but not sexually inexperienced ones, spent more time investigating estrous than diestrous female urine and vaginal odors. Anal sac secretions from estrous bitches were not more attractive to males than those from diestrous bitches. Estrous females spent no more time than diestrous ones in the investigation of male anal sac secretion and urine odors. Male urine and anal sac secretions elicited little investigation from male conspecifics. Females spent more time investigating female urine odors than female anal sac or vaginal secretion odors, and exhibited a slight general preference for diestrous over estrous stimuli. A positive correlation between the odor investigation times of this study and investigation times of comparable animals to conspecifics in a social situation suggests odor preferences are relatively good indicators of social preferences, and vice versa, in this breed.     

Social stress in guinea pigs

Confrontation studies between eight pairs of two 7-8 months old male guinea pigs, each raised with one female from their 30th day of age, were conducted in an enclosure of 2 m2 in the presence of an unfamiliar female. The opponents were chronically kept together for up to 8 days. When they were not separated after 52 hr, one or (in 1 case) both males fell into a comatose state and died 5-8 days after the onset of the confrontation. Four hr after the onset of the confrontations both opponents showed significantly increased plasma glucocorticoid (CS) titers. At this time as well as 24 hr earlier, prospective winners (PW) and prospective losers (PL) did not yet differ in any physiological parameter measured. Forty-eight hr later, PW were characterized by a low body weight loss, low CS, high testosterone (T) and elevated plasma-catecholamine (CA) values. In contrast, PL showed a high body weight loss, very high CS titers, low T-titers and more elevated CA values than PW. Despite these highly significant differences in physiological parameters, PL and PW did not yet differ in the frequency of any agonistic behavioral element recorded. Seventy-two hr after the onset of the chronic cohabitation, losers began to behave more and more passive, they ceased feeding and drinking and were less and less involved in social interactions. The physiological differences between winners and losers intensified.(ABSTRACT TRUNCATED AT 250 WORDS)     

Group size modifies the patterns and muscle carbohydrate effects of aggression in Betta splendens.

Aggressive encounters of previously isolated individuals were investigated in dyads and groups of five. Fights were longer and more intense when they were performed in dyads compared to fights involving five fishes. During aggressive encounters, an elevation in carbohydrate catabolism was noticed in both dyads and groups. Losing a fight resulted in a reduction in glycogen content and an increase in glycogen synthesis. Similar changes in winners did not appear; thus, the metabolic response in losers was different from that noticed in winners, both in dyads and groups. In dyadic contest winners, a marked increase in the free glucose content and glucose consumption was noticed (without changes in losers). In groups, free glucose content of the winners was not modified, while glucose consumption was enhanced both in winners and losers. Thus, the differences existing between winners and losers were greater in dyads compared to those noticed in groups. The energy cost of aggression seems to be different in dyads compared to groups of five. The rate of glucose oxidation was strongly reduced in dyads (there were no differences between winners and losers in this respect), while in groups, this parameter was not modified.     

Aggressive encounters alter the activation of serotonergic neurons and the expression of 5-HT1A mRNA in the hamster dorsal raphe nucleus

Serotonergic (5-HT) neurons in the dorsal raphe nucleus (DRN) have been implicated in stress-induced changes in behavior. Previous research indicates that stressful stimuli activate 5-HT neurons in select subregions of the DRN. Uncontrollable stress is thought to sensitize 5-HT neurons in the DRN and allow for an exaggerated 5-HT response to future stimuli. In the current study, we tested the hypothesis that following aggressive encounters, losing male Syrian hamsters would exhibit increased c-Fos immunoreactivity in 5-HT DRN neurons compared to winners or controls. In addition, we tested the hypothesis that losers would have decreased 5-HT1A mRNA levels in the DRN compared to winners or controls. We found that a single 15-min aggressive encounter increased c-Fos expression in 5-HT and non-5-HT neurons in losers compared to winners and controls. The increased c-Fos expression in losers was restricted to ventral regions of the rostral DRN. We also found that four 5-min aggressive encounters reduced total 5-HT1A mRNA levels in the DRN in losers compared to winners and controls, and that differences in mRNA levels were not restricted to specific DRN subregions. These results suggest that social defeat activates neurons in select subregions of the DRN and reduces message for DRN 5-HT1A autoreceptors. Our results support the hypothesis that social stress can activate 5-HT neurons in the DRN, reduce 5-HT1A autoreceptor-mediated inhibition, and lead to hyperactivity of 5-HT neurons.