The evolution of violence in men: the function of central cholesterol and serotonin.

Numerous studies point to central serotonin as an important modulator of maladaptive behaviors. In men, for instance, low concentrations of this neurotransmitter are related to hostile aggression. A key player in serotonin metabolism seems to be central cholesterol. It plays a fundamental role in maintaining the soundness of neuron membranes, especially in the exocytosis transport of serotonin vesicles into the synaptic cleft. In this review, we attempt an evolutionary approach to the neurobiological basis of human male violence. Hominid evolution was shaped by periods of starvation but also by energy demands of an increasingly complex brain. A lack of food resources reduces uptake of glucose and results in a decreased energy-supply for autonomous brain cholesterol synthesis. Consequently, concentrations of neuromembrane cholesterol decrease, which lead to a failure of the presynaptic re-uptake mechanism of serotonin and ultimately to low central serotonin. We propose that starvation might have affected the larger male brains earlier than those of females. Furthermore, this neurophysiological process diminished the threshold for hostile aggression, which in effect represented a prerequisite for being a successful hunter or scavenger. In a Darwinian sense, the odds to acquire reliable energetic resources made those males to attractive spouses in terms of paternal care and mate support. To underpin these mechanisms, a hypothetical four-stage model of synaptic membrane destabilization effected by a prolonged shortage of high-energy, cholesterol-containing food is illustrated.

Fight or flight? Effects of vaginal oestrus on cortisol, testosterone, and behaviour in guinea pig female-female interaction.

It is accepted that social stress in relation to confrontation and competition can elicit behavioural and hormonal changes in social mammals. These effects have, however, been less frequently studied among female-female interactions. In the present study female-female confrontation experiments were carried out to monitor socio-positive and agonistic behaviour by controlling for the oestrus cycles of 12 individuals. Additionally, plasma cortisol (CORT) and testosterone (T) levels were determined before and after the experiments. During non-oestrus conditions a significant increase in CORT levels from pre- to post confrontation was registered and females spent more time to sit side by side. During vaginal oestrus the confrontation experiments revealed avoiding of a conspecific female by showing increased flight behaviour. However, during that period no changes in CORT levels were found. But, a non-significant increase in T was measured from pre- to post confrontation in both cycle phases, while no differences in the display of aggressive behaviours were found. These findings indicate considerable influences of different oestrus cycle phases on social stress-induced CORT secretion and the modulation of socio-positive and agonistic behaviour in female guinea pigs.

Non-lactating versus lactating females: a comparison of sex steroids, sexual coloration, and sexual behavior in Japanese macaques.

Female Japanese macaques are seasonal breeders distinguished by their red-colored hindquarters, face, and nipple skin areas. Intensity of coloration seems to be associated with sexual attractiveness, behavior, and fluctuating sex steroids. Our aim was to investigate whether the color intensity of these regions differed between lactating (LA) and non-lactating (NLA) females during sexually inactive (SI) and active (SA) phases. Coloration scores of 19 adult females were classified using color tables. Estrogen and progesterone metabolites were determined in fecal samples. Weekly comparison between both groups revealed significantly increased coloration of the hindquarters area from week 13 (SI) until the end of the observation period, and for the nipple skin throughout the SI and SA periods. Face coloration differed marginally. Hormonally, NLA females showed significantly increased excretion rates of sex steroids at the end of the SI phase and throughout the whole SA period. Logistic regression analyses between elevated fecal steroids and nipple coloration disclosed a significant relationship for NLA females during the SI period. This connection persisted and included hindquarter coloration during the SA period. NLA females showed increased intromission with ejaculation, but no difference was found for intromission without ejaculation. In conclusion, results demonstrate increased endocrine excretion rates for NLA females during the whole observation period, paralleled by an enhanced, fertility-signaling sexual attractiveness.

Spatial learning and memory differs between single and cohabitated guinea pigs.

In socially-living animals, social enrichment enhances spatial learning and memory while separation from conspecifics can severely impair these abilities. In the present work, guinea pigs were kept in isolation or cohabitated in heterosexual pairs and then subjected to a labyrinth task. Latency-time to bait, error-rate, amount of movement and pre- and post-experimental cortisol (CORT) were registered. During a 5d-acquisition phase, single animals (N=19) showed a more efficient encoding of spatial information, with significantly decreased latency-time and error-rate over the time course. In contrast, cohabitated animals (N=19) did not show a significant improvement. Three days after acquisition, memory was tested in a retention test, under the same conditions. With regard to behavioral performance, there was no significant difference between cohabitated and single animals. Pre-experimental CORT was significantly higher in cohabitated animals when compared to single ones. Post-experimentally, CORT increased significantly in singles but not in cohabitated animals when compared to pre-experimental values. Thus, both groups did not differ from each other at that point. Social condition seemed to be an important modulator, in that learning and memory were more impaired in paired animals than in single ones. The failure of cohabitated animals to encode spatial memory more quickly may have been caused by a more chronically up-regulated HPA-axis. The post-experimental CORT increase of singles may be due to more efficient handling of short-term stress exposure.

Non-invasive measurement of adrenocortical and gonadal activity in male and female guinea pigs (Cavia aperea f. porcellus).

Taking blood samples is a common method in biomedical and biological research using guinea pigs. However, most blood sampling techniques are complicated and highly invasive and may therefore not be appropriate for certain research topics concerning stress and reproduction. Thus, a non-invasive method to measure steroid hormones is critically needed. The aim of this study was the biological validation of corresponding enzyme immunoassays for the measurement of fecal cortisol, progesterone, estrogen, and testosterone metabolites in guinea pigs. We examined the effect of subcutaneous injections of ACTH or saline on fecal cortisol metabolites to investigate the suitability of fecal samples to monitor adrenocortical activity. Furthermore, we investigated whether fecal sex steroid metabolites accurately reflected endocrine changes observed in plasma samples during female estrous cycles and male puberty, respectively. In addition, we compared fecal testosterone metabolites of intact males, castrated males, and females to investigate the reliability of fecal samples in discriminating gonadal status of males. Concentrations of fecal cortisol metabolites were significantly increased following ACTH challenge, indicating that adrenocortical activity can be monitored via fecal samples. Secondly, in females, plasma and fecal gonadal steroids were significantly correlated in most subjects. The assay for testosterone metabolites, on the other hand, could not clearly discriminate between test groups. From these findings we conclude that fecal samples can be used for the non-invasive assessment of adrenocortical and female reproductive status in guinea pigs. Testosterone metabolism seems to be more complex and further investigations are needed to establish a more suitable assay.

Urinary androgens and cortisol metabolites in field-sampled bonobos (Pan paniscus).

Urinary metabolites of androgens and cortisol were measured in free-living male and female bonobos. Sex differences and correlations between adrenal and gonadal steroid excretion were investigated. The immunoreactive concentrations of androgens were measured with two different androgen assays. One assay used a testosterone (T) antibody raised with a 17beta-hydroxy group, and the other employed an antibody raised against a reduced form, 5alpha-androstane-17alpha-ol-3-one-CM (17alpha) with cross reactivity for epitestosterone and 5alpha-androstanedione. Both assays have been used in bonobo and chimpanzee studies where non-invasive techniques were employed. The levels of 17alpha-androgen metabolites were 1.7- and 3-fold higher than those of T-metabolites in males and females. The two androgen assay results correlated in males but not females. There was a sex difference in the T-metabolites measured. Male levels were significantly higher. Levels of 17alpha in the two sexes were similar. Cortisol metabolite levels (CORT) were similar between the sexes. The T-metabolites were significantly correlated with CORT in males but not in females. In females, the 17alpha-androgen metabolites correlated with CORT. This suggests that either androgen secretion or metabolism differs between the sexes. A parsimonious interpretation of the androgen assay cortisol/androgen correlation differences would be that larger components of dehydroepiandrosterone (DHEA), androstenedione or epitestosterone from the adrenal androgens were being excreted and measured in the females. The CORT/T metabolite interactions in males may reflect male-specific social or metabolic endocrine conditions.

Impact of social environment on female chimpanzee reproductive cycles.

The socio-sexual environment of a female is known to affect ovarian function. Increased male contact can enhance menstrual cycle regularity. Conversely, social deprivation constitutes a form of stress that often alters cyclicity and the secretion of reproductive hormones. The present study was carried out on captive female chimpanzees to examine possible interactions among housing conditions, menstrual cycle length, morphological changes in secondary sexual character expression and endocrine release patterns related to follicular and luteal function. Animals were housed over a period of 2 years either with a male conspecific or singly. Blood samples were collected over three cycles, and anogenital swelling changes registered to define menstrual cycle phases. Fecal sampling techniques were used to monitor cortisol as a measure of stress-load. Male presence seemed to affect female cyclicity. Females housed with a male had shorter and more regular cycles than singly housed females. Prolactin, gonadotropins and estradiol levels were generally higher in paired females during specific cycle phases. Group variation was not always significant. No differences were found in progesterone. Sexually cohabited females tended to have lower fecal cortisol metabolites immediately before and after maximum tumescence. We suggest that the close behavioral, physical and olfactory contact with a male conspecific can act as a sort of zeitgeber to modulate ovarian function by stabilizing the female cycle and, perhaps, enhancing folliculogenesis and ovulation.

Morphometric and hormonal changes during the chimpanzee menstrual cycle.

BACKGROUND: Sex steroids affect many peripheral tissue sites in female mammals. Receptors for these hormones have been found in skin, fat, and bone. In women, these tissues can show morphological changes during the menstrual cycle that may be directly related to steroid secretion. METHODS: The present study was done on chimpanzees to document morphometric markers associated with these tissues (anogenital swelling volume, skin fold thickness as indicator of subcutaneous fat, bony diameters of mandible, wrist, and elbow) and to compare them with cyclic patterns of estradiol, progesterone, testosterone, gonadotropins, and prolactin. RESULTS: Swelling volume changed significantly over the menstrual cycle. All other morphometric parameters showed variation without statistical significance. Skin folds were thickest during the luteal phase. Bony diameters displayed similar but less distinctive changes. Testosterone correlated positively with diameter sites, inversely with subcutaneous fat. No relationships with either estradiol or progesterone were found. We assume that subcutaneous fat and morphometric bone parameters exhibit cycle-dependent changes that may be caused by changes in steroid secretion.