Postpartum aggression in mice: Experiential and environmental factors

Six experiments were conducted to assess the influence of duration of lactation, the presence of young, and the stimulus characteristics of intruder animals upon postpartum aggression of mice. The first experiment showed that postpartum aggression toward conspecifics was highest between Day 3 and Day 8, declined between Day 9 and Day 14, and was present toward males but absent toward females between Day 15 and Day 21 of the lactation period. Experiment 2 showed that lactating mice rarely attacked conspecifics to which they had been previously exposed but would readily attack strangers. Experiment 3 and 4 demonstrated that lactating animals never attacked intruders when tested 5 hr after pup removal. However, placement of young behind a wire partition in the home-cage for 5 hr or replacement of the offspring for as little as 5 min following 5 hr of separation restored postpartum aggression. The fifth experiment showed that 1- and 10-day old intruders were seldom attacked while intense aggression was directed against 14- and 20-day old intruders. Finally, Experiment 6 demonstrated that 14-day old intruders whose hair was removed were rarely attacked.     

Mammary stimulation and maternal aggression in rodents: thelectomy fails to reduce pre- or postpartum aggression in rats

In mice, tactile stimulation of the nipples appears to be critical for the onset of postpartum maternal aggression. Surgical removal of the nipples (thelectomy) blocks aggression if performed prior to parturition. In rats, indirect evidence suggests a similar role for nipple stimulation in maternal aggression. Two experiments were undertaken to determine whether thelectomy prior to mating reduces pregnancy-induced and/or postpartum aggression in this species. In the first, thelectomized and sham-thelectomized females were subjected to home cage tests (pups, if any, present) with unfamiliar male intruders on Gestation Days 18 and 21 and Lactation Days 3 and 5. Additional groups of thelectomized females were tested one time only on either Lactation Day 5 or 12. Thelectomized and control females were equally aggressive; postpartum, nearly all females in both groups attacked. Experiment 2 used females that were hysterectomized-ovariectomized (HO) on Gestation Day 16. Such females are not aggressive prior to initiating maternal behavior, but become highly aggressive (over 80% attacking) after commencing maternal care. Females again were thelectomized or sham-thelectomized prior to mating. On Day 16 HO was performed, and 48 hr later continuous exposure to pups was begun. After the females had displayed maternal behavior for 1.5-2 days, intruder tests were conducted. All females attacked at least once, with no differences between treatment groups. Thus thelectomy does not reduce maternal aggression in the rat. This finding, however, does not preclude a role for tactile ventral stimulation in mediating maternal aggression.     

Further evidence for the role of nitric oxide in maternal aggression: effects of L-NAME on maternal aggression towards female intruders in Wistar rats

It has been shown that nitric oxide (NO) increases aggression in male mice, whereas it decreases aggression in lactating female mice and prairie voles. It is also known that aggression can be exhibited at different levels in rodent species, strain or subtypes. The aims of this study were to investigate the proportion of aggressiveness in Wistar rats, the effect of intraperitoneally administered nonspecific nitric oxide synthase (NOS) inhibitor L-NAME (NG-nitro L-arginine methyl ester) on maternal aggression towards female intruders, and whether these effects are due to NO production or not. Rats were given saline intraperitoneally on the postpartum Day 2 and aggression levels were recorded. The same rats were given 60 mg/kg L-NAME or D-NAME (NG-nitro D-arginine methyl ester) on the postpartum Day 3 and their effects on aggression levels were compared to saline. While L-NAME administration did not cause any differences in the total number of aggressive behavior, aggression duration and aggression intensity, it reduced the proportion of animals showing aggressive behavior. In addition, the latency of the first aggression was significantly increased by L-NAME. In the D-NAME group, however, no significant change was found. Our results have shown that L-NAME reduces maternal aggression towards female intruders in Wistar rats through inhibition of NO production. These results suggest that the role of NO in offensive and defensive maternal aggression shares neural mechanisms.     

Aggression in the female golden hamster: effects of reproductive state and social isolation

To gain information on possible hormonal correlates, the aggressive behavior of intact female hamsters towards males was observed at various times during the estrous cycle, pseudopregnancy, pregnancy, and lactation. For methodological information, estrous cycle females also were tested after varying periods of social isolation. It was found that pregnant and especially lactating hamsters were more aggressive than pseudopregnant or estrous cycling females. Comparisons of days within each reproductive condition showed that aggression tended to be higher on certain days: the day preceding behavioral estrus of the estrous cycle, Day 10 of pregnancy, and the first 5 days of lactation. Except for pseudopregnancy, sexual behavior unaccompanied by aggression occurred at some time during all reproductive conditions, and both sexual behavior and aggression were found to occur together on Day 10 of pregnancy and Day 1 of lactation. The changes in aggressive behavior associated with reproductive states were attributed to increased male interest, inhibition by ovarian hormones, and facilitation by prolactin. Increasing periods of social isolation also were found to be associated with increased aggression. It was suggested that this effect, too. might have been due to increased prolactin levels.     

Effects of resource distribution on activity and territory defence in house mice, Mus domesticus

How much an animal invests in defending a territory depends, in part, on the quality, quantity and distribution of resources, but do animals target their investment in defence within the territory according to the location of different resources? We examined whether wild-caught male and female house mice increased aggression towards intruders of the same or opposite sex when in areas containing their food/water or nest site. Pairs of resident mice were established in enclosures consisting of two areas. In split enclosures, food and water were in one area and a nest site in the other, while in clumped enclosures all resources were in one area with no resources in the other. Residents of both sexes were quicker to attack intruders, and were more aggressive, in areas containing resources than in an area containing no resources. There was no difference in aggression between areas containing food/water, a nest site or both. The effect of resources on territorial defence was due largely to changes in aggression against intruders of the opposite sex. Residents, especially males, attacked intruders of their own sex more than those of the opposite sex. This bias was much stronger when no resources were present, owing to low aggression against opposite-sex intruders in the absence of resources. There were no differences in intruder aggression, and the relative size of the intruder (heavier or lighter than the resident) had no effect on resident aggression. Residents also spent less time out of their nest in clumped enclosures owing to low activity in the nonresource area.     

Intermale aggression of subordinate resident long-evans rats

Dominant male rats were separated from their colonies and subordinate residents were tested for aggression after a two week period. Subordinate animals showed significant increases in body weight and aggressive behavior toward intruders. Replacement of the previously dominant male led to rank reversals in three of six colonies. To assess whether two weeks of separation from the dominant male was necessary for subordinate males to exhibit aggression towards conspecifics, another intruder test was given following an additional two weeks of social housing. At the conclusion of this test, the more aggressive resident was removed and a naive intruder was immediately introduced into the colony with the subordinate male resident and again at 1, 5, and 10 day intervals. Aggressive posturing by the subordinate resident increased immediately following the removal of the dominant male, and remained at a high level throughout the subsequent intruder tests. In contrast, biting was initially low but increased steadily throughout the period of separation. These findings demonstrated that social or situational changes within a colony can greatly influence the aggression of subordinate males.     

The role of progesterone in pregnancy-induced aggression in mice

A series of six experiments was performed in order to explore the potential involvement of progesterone (P) in pregnancy-induced aggression (PIA) displayed by Rockland-Swiss mice toward adult male intruders. In Experiment 1, circulating levels of P and aggression were low on gestation Days 6 and 10 while both the behavior and the steroid reached peak levels by gestation Day 14. By gestation Day 18 (the day prior to parturition), serum P was at its lowest level yet aggressive behavior was still intense. Also, individual differences in the display of fighting behavior by pregnant females were not related to circulating P. Experiments 2 and 3 showed that supplemental P treatment to early pregnant female mice did not advance the onset of aggression. Experiment 4 showed that P treatment promoted the onset and elevated the incidence of aggression in virgin mice, but only in those females with intact ovaries. Experiment 5 showed that the aggressive behavior of P-stimulated virgin females was qualitatively and quantitatively different from that exhibited by pregnant mice in that the former exhibited fewer attacks and lunges than the latter. Finally, Experiment 6 showed that the removal of P from aggressive, P-stimulated virgins dramatically attenuated levels of the behavior. This contrasts sharply with the continued fighting behavior observed in late pregnant P-deficient mice. Thus, although P augments aggression in female mice it apparently is not a sufficient stimulus for producing pregnancy-like aggressive behavior.     

Mice: progesterone stimulates aggression in pregnancy-terminated females

Three experiments were conducted in order to assess the role of progesterone (P) in the aggressive behavior displayed by late pregnant Rockland-Swiss mice toward adult male intruders. In Experiment 1, hysterectomy on the 15th day of gestation reduced the aggressive behavior normally displayed by pregnant mice toward male intruders. In Experiment 2, Silastic implants of P stimulated aggression in hysterectomized mice but did not fully restore the behavior to the level of fighting normally observed in pregnant animals. In Experiment 3, aggressive behavior in P-treated hysterectomized animals was inhibited by simultaneous exposure to estradiol (E). Also, treatment with E alone did not stimulate aggression in hysterectomized mice. While pregnancy-induced aggression is promoted by P, other neuroendocrine factors may act in concert with the steroid to fully stimulate aggression in gravid mice.     

Territory Defence in Black Redstarts,Phoenicurus ochruros: Effects of Intruder and Owner Age?

European black redstarts, Phoenicurus ochruros, vigorously defend all-purpose territories and exhibit delayed plumage maturation, most subadult males looking exactly female-like in their first breeding season. We tested the hypotheses that such dull subadult male plumages are beneficial in order to reduce aggression of adult males either by deception or by honest signalling of subordinance status, and that, in turn, conspicuous (adult) plumage colorations are able to intimidate contenders because they act as a signal of fighting ability and aggressive motivation. Adult and dull yearling black redstart territory owners were confronted with intruders mimicked by stuffed mounts of either a conspicuous adult or a dull subadult male. Our results do not support the hypotheses tested: dull plumages of young intruders did not reduce aggression from adult territory owners and aggressiveness towards intruders was significantly higher in yearling territory owners as compared with adult owners. Conspicuous intruders did not deter dull territory owners and we found no indications that conspicuous male coloration is a signal of fighting ability or aggressive motivation in this species. Instead, the amount of aggressive response to intruders showed considerable individual variance within age classes regardless of the plumage of the intruder and the status of the owner.     

Effects of ergocornine and prolactin on aggression in the postpartum golden hamster

The effects on aggressive behavior of prolactin (PRL) and ergocornine hydrogen maleate, an inhibitor of PRL secretion, were investigated in the female golden hamster. Because high aggression and PRL levels are associated with lactation in hamsters, postpartum females were used as subjects. In the first experiment, three groups of ovariectomized and hysterectomized females were compared: normally lactating, ergocornine-treated, and ergocornine plus replacement PRL treated. Normally lactating mothers were typically aggressive towards males in an arena, whereas females given ergocornine were not. Females given both ergocornine and PRL showed an intermediate level of aggression. Although ergocornine suppressed aggression towards adult males, attacks on pups increased. A second experiment sought to determine if ergocornine would depress aggression when PRL involvement was unlikely. At least 30 days following pup removal, females from the first experiment were “trained” to attack home-cage intruders consistently. After ergocornine administration, home-cage attacks by these experienced females were not diminished. Since PRL levels were probably low in these animals, it was concluded that the effects of ergocornine on aggression were limited to instances in which PRL was involved, and that PRL probably can facilitate aggression.     

Aggression persists after ovariectomy in female rats

Aggression occurs not only in males but also in females, however, under different sex-specific stimulus and endocrine conditions. After being housed with males, female rats exhibit frequent and intense aggressive behavior toward unfamiliar rats. However, the female residents primarily attack female intruder rats, while the male residents attack males and not females. Altering the hormonal condition of the intruders can modify the behavior that they provoke from the residents. Castration of the male intruders reduces aggression from male residents, but ovariectomy of the female intruders does not alter the behavior of the female residents. Treatment of the gonadectomized intruders with gonadal steroids significantly alters the response of the male residents. Resident-intruder aggressive behavior depends on the presence of the testes in the male residents but not on the ovaries or on lactation in the female residents. Even 7 weeks after ovariectomy the female residents continue to show aggressive behavior toward female intruders. In the same time period the castrated male residents show a marked decrease in aggressive and sexual behavior.     

Information on Resource Quality Mediates Aggression between Male Madagascar Hissing Cockroaches, Gromphadorhina portentosa (Dictyoptera: Blaberidae)

Male Madagascar hissing cockroaches, Gromphadorhina portentosa Schaum (Dictyoptera: Blaberidae) have a well‐defined dominance hierarchy that has been assumed to explain the outcome of most competitive interactions. We studied whether males of this species would alter their level of aggression towards unfamiliar rivals as a function of changing resource availability and value – two factors that are key to aggression levels in non‐hierarchical species. We quantified male aggression as three variables (aggressive state – behaviours measured by their duration; aggressive act – behaviours measured by their frequency of occurrence; aggressive latency – the latency to first aggressive behaviour, either state or act) and tested for any context‐specific variation within each by manipulating both territorial status (males were either residents or intruders) and access to mates (female present or absent). Both the presence of a female and territorial status affected male aggression towards rivals as measured by duration of aggressive state. Highest levels of aggression were displayed by residents when a female was present. These results show that inter‐male aggression in G. portentosa is tuned to the immediate expected payoff from fighting, and not exclusively aimed at establishing dominance relationships (which can affect future payoffs).     

Daily changes in concentrations of prolactin in serum of prepubertal bulls exposed to short- or long-day photoperiods

Early temporal changes in concentrations of prolactin (PRL) in serum after a sudden change in photoperiod and daily responsiveness to PRL-releasing and inhibiting factors were investigated in prepubertal Holstein bull calves exposed to different photoperiods. In calves switched from 8-hr light: 16-hr dark to 16-hr light:8-hr dark, there was no observable change in the daily pattern of serum concentrations of PRL after 1, 2, or 4 days. On the other hand, in animals switched from 16-hr light:8-hr dark to 8-hr light:16-hr dark, there was a consistent increase in serum PRL from 33.4 ng/ml on Day 0 to maximum values of 57.3, 62.7, and 78.9 ng/ml between 14 and 18 hr after onset of light on Days 1, 2, and 4, respectively. Thus, absence of light allowed expression of a daily rhythm in serum concentrations of PRL that persisted for at least 4 days after the photoperiod switch. There were no differences in L-dopa inhibition of PRL release in animals exposed to 16-hr light:8-hr dark at 3 or 15 hr after onset of light. However, thyrotropin-releasing hormone-induced release of PRL was greater 3 hr after onset of light (11 hr after onset of dark) compared with release at 9, 15, and 21 hr after onset of light in animals exposed to 16-hr light:8-hr dark, but not in bulls exposed to 8-hr light:16-hr dark. The results provide evidence that the cue for the putative photosensitive period of PRL secretion in cattle may be more closely associated with onset of dark, not onset of light.     

Teratogenic, biochemical, and histological studies with mice prenatally exposed to 2.45-GHz microwave radiation

Pregnant CD-1 mice were exposed to 2.45-GHz continuous wave microwave radiation at an incident power density of 30 mW/cm2. The local specific absorption rate near the uterine area (deep colonic location), as determined from time-temperature profiles measured with a Vitek thermistor probe, was 40.2 mW/g. Groups of mice were exposed 8 hr per day through Days 1-6 or 6-15 of pregnancy. Other groups of animals were exposed to an elevated ambient temperature of 31 degrees C which increased the colonic temperature 2.3 degrees C, the same as that produced by the microwaves. Sham-irradiated groups of animals were treated exactly the same as the microwave-exposed animals. For the two conditions, temperature exposed and sham exposed, two groups of animals were used. One group was handled in the same manner as the microwave-irradiated group and the other group was not handled so as to evaluate the effects of stressing the animals by handling. Eleven groups of animals were used in the complete study: five groups for gestational Days 1-6, five groups for gestational Days 6-15, and one group of cage control animals. On Day 18 of gestation the dams of all experimental groups were sacrificed and their reproductive status was determined. The fetuses were examined for visceral and skeletal alterations. Brain cholinesterase activity and histology were evaluated in the groups exposed on Days 6-15. The results show that microwave radiation increases embryo lethality at the early stages of gestation (exposure Days 1-6). Fetal toxicity and teratogenicity were not significantly increased by exposure to microwaves on either Days 1-6 or 6-15 of gestation. Cholinesterase activity and histology of the brain of 18-day-old fetuses were not adversely affected.     

Parity-associated reductions in behavioral sensitivity to opiates

Behavioral and physiological responses differ between primiparous and multiparous female rodents. Specifically, multiparous females respond with the full repertoire of maternal behaviors much more rapidly and with greater intensity than their primiparous counterparts. Since opiates inhibit the expression of maternal behavior in postpartum rats and can be reversed by means of the opiate antagonist naloxone, we investigated whether multiparous females would be resistant to the inhibitory effects of opiates on maternal behavior, relative to primiparous females. In Experiment 1 we evaluated the effects of a range of doses of morphine sulfate (MS; 0.625, 1.25, 2.5, 5.0, and 10.0 mg/kg or saline) on maternal behavior in primiparous females on Days 5-6 of lactation. The 5.0 and 10.0 mg/kg doses effectively disrupted maternal behavior, whereas the lower doses were ineffective or only marginally disruptive. In Experiment 2, age-matched female rats were timed-mated and tested for maternal behavior from Day 5 to 13 of lactation, after daily injections of the 5.0 mg/kg dose of MS. On Day 5 of lactation, this morphine treatment eliminated full maternal behavior in 87% of the primiparous animals, but only 37% of the multiparous animals were affected. By Day 10 of lactation, 100% of the multiparous females displayed full maternal behavior after MS treatment, whereas only 69% of primiparous females were responsive. In Experiment 3, analgesic responses were measured both in rats experiencing their initial or second pregnancy, and in postpartum, lactating rats after MS (5.0 mg/kg) administration. Using a tail-flick apparatus to measure analgesia, we found multigravid females to be significantly less analgesic prepartum than primigravid females, suggesting less sensitivity to endogenous opioids.(ABSTRACT TRUNCATED AT 250 WORDS)     

Dominance and aggression in social groups of male and female rats

Two experiments were performed to examine aggression and dominance in domestic male and female $\text{Rattus norvegicus}$ living in small mixed-sex (3 males and 3 females) groups. Experiment 1 examined the development of aggression in females. A single female (alpha) within each of the six colonies tested showed the preponderance of attacks on male intruders placed into the home-cage when male colony residents were absent. Over 12 weeks of intruder-aggression training female alphas showed only a mild nonsignificant elevation of aggressive behavior. A comparison of aggression of male and female colony alphas tested with opponents of each sex revealed that aggression was mainly directed at like-sex opponents, and that female attack was more defensive in character than male attack regardless of opponent sex. The highest intensity of aggression occurred when male alphas confronted male intruders. Although intruders never showed offense toward male residents, 61% of intruding males showed offense in response to attack by females.Experiment 2 investigated the relationship between aggressive dominance and competitive measures of dominance within each of 10 mixed-sex colonies. Alpha stat s of male and female colony residents did not reliably predict priority of access to food or water in tests of direct resource competition with like-sex colony members. When colony males were simultaneously tested for copulation, the copulatory behavior of alpha males was significantly greater than that of other colony males. Results are discussed in relation to the role of aggression in the reproductive strategy of male and female $\text{Rattus norvegicus}$.     

Induction of ovarian follicular cysts in the pregnant rat by human chorionic gonadotropin.

Prolonged stimulation by human chorionic gonadotropin (hCG) induces ovarian follicular cysts in progesterone-synchronized immature rats [Bogovich, Endocrinology 1989; 124:1646-1653]. To determine if unabated stimulation by hCG has a similar effect on follicular development in adult ovaries, pregnant rats were given either 0 (control), 1, or 3 IU hCG twice daily for 9 days beginning on Day 13 of pregnancy. By Day 22 of pregnancy, rats treated with 1 IU hCG possessed large antral follicles at least 1 mm in diameter: approximately 33% larger than the diameters of preovulatory follicles observed in control rats (0 IU hCG). In contrast, rats treated with 3 IU hCG displayed ovarian follicular cysts up to 5 mm in diameter, with well-developed thecae and just a remnant of granulosa cells. Progesterone, androstenedione, and estradiol accumulation was greater in follicular incubates from hCG-treated rats than in incubates from control rats. Progesterone increased in response to cAMP in incubates from all treatment groups on all days tested. Androstenedione increased in response to cAMP on Day 22 of pregnancy for follicles from control animals, on all days tested for follicles from rats treated with 1 IU hCG, and on Days 15-19 for follicles from rats treated with 3 IU hCG. Androstenedione production in the presence of 300 ng of exogenous testosterone was significantly greater in follicular incubates from animals treated with 1 and 3 IU hCG than incubates from control animals on Days 19-22 of pregnancy.(ABSTRACT TRUNCATED AT 250 WORDS)     

Food restricting young hamsters (Mesocricetus auratus) affects sex ratio and growth of subsequent offspring

Trivers and Willard (1973) predicted that stressed adult female mammals may enhance their fitness by skewing offspring sex ratios and maternal investment to favor daughters. The present study investigated whether stressing young hamsters might also influence sex ratio and growth of subsequent offspring. Control females received food ad libitum (A) on Days 1-50 postpartum (AA). Experimental females were food-restricted (R) either on Days 1-25 (RA), Days 26-50 (AR), or Days 1-50 (RR) postpartum. Subjects were mated when 91-95 days old. Litter sizes and survivorship (= % litters within a treatment that contained at least one pup), sex ratio (= % males), and pup weights in the next generation were recorded every fifth day from parturition until Day 25 postpartum. Control litters contained significantly more offspring at birth than did RR litters. Sex ratio was significantly higher at birth for AA litters than for the other treatments. Postpartum sex ratio within each group remained similar to that recorded at birth. RR litters contained significantly fewer pups compared to the other three treatments from Days 5-25. RR female pups weighed significantly more at birth than their counterparts in the other treatments. Weights of males at birth were similar in all treatments. By Day 25, both male and female RR pups weighed significantly less than control, AR, and RA pups. Food restriction early in life may have long-term consequences on sex ratio and pup growth in golden hamsters.     

Effect of suppression of plasma prolactin on ovulation, plasma gonadotrophins and corpus luteum function in LH-RH-treated anoestrous ewes.

Nineteen Scottish Blackface ewes were given LH-RH (3 X 30 micrograms i.v., 90-min intervals) during anoestrus when prolactin levels were elevated. Plasma levels of prolactin were suppressed with CB 154 (twice daily, i.m.) on Days -5 to 0 (N = 5), 0 to +5 (N = 5) or -5 to +5 (N = 5) around the day of LH-RH treatment (Day 0). Control animals (N = 4) received saline on Days -5 to +5. Nine animals ovulated forming corpora lutea as judged by laparoscopy on Day +7. No difference in FSH or LH levels was found between treatments and ovulations occurred equally in all treatment groups. Progesterone levels were less than ng/ml in all animals up to Day 14. It is concluded that short-term suppression of prolactin does not affect the incidence of ovulation or corpus luteum progesterone production in LH-RH-treated anoestrous ewes.     

Maternal aggression as a deterrent to infanticide in Peromyscus leucopus and P. maniculatus

Adult male and female white-footed mice (Peromyscus leucopus) and deermice (P. maniculatus) from sympatric populations commit infanticide on neonates of either species. A laboratory experiment was conducted to determine whether aggression in maternal females could effectively deter infanticidal intruders. Females of both species exhibited post-partum aggression and dominated intruders of either sex or species in 83 of 88 trials. In the absence of the mother, neonates were attacked by intruders in 82 out of 84 trials. Maternal females were considerably more aggressive than anoestrous females. The implications for female territorially as a mechanism to prevent infanticide are discussed.