Oestrous State Research Articles

A hypothetico-deductive approach to assessing the social function of chemical signalling in a non-territorial solitary carnivore.

The function of chemical signalling in non-territorial solitary carnivores is still relatively unclear. Studies on territorial solitary and social carnivores have highlighted odour capability and utility, however the social function of chemical signalling in wild carnivore populations operating dominance hierarchy social systems has received little attention. We monitored scent marking and investigatory behaviour of wild brown bears Ursus arctos, to test multiple hypotheses relating to the social function of chemical signalling. Camera traps were stationed facing bear ‘marking trees’ to document behaviour by different age sex classes in different seasons. We found evidence to support the hypothesis that adult males utilise chemical signalling to communicate dominance to other males throughout the non-denning period. Adult females did not appear to utilise marking trees to advertise oestrous state during the breeding season. The function of marking by subadult bears is somewhat unclear, but may be related to the behaviour of adult males. Subadults investigated trees more often than they scent marked during the breeding season, which could be a result of an increased risk from adult males. Females with young showed an increase in marking and investigation of trees outside of the breeding season. We propose the hypothesis that females engage their dependent young with marking trees from a young age, at a relatively ‘safe’ time of year. Memory, experience, and learning at a young age, may all contribute towards odour capabilities in adult bears.

Atrazine and Breast Cancer: A Framework Assessment of the Toxicological and Epidemiological Evidence

The causal relationship between atrazine exposure and the occurrence of breast cancer in women was evaluated using the framework developed by Adami et al. (2011) wherein biological plausibility and epidemiological evidence were combined to conclude that a causal relationship between atrazine exposure and breast cancer is “unlikely”. Carcinogenicity studies in female Sprague-Dawley (SD) but not Fischer-344 rats indicate that high doses of atrazine caused a decreased latency and an increased incidence of combined adenocarcinoma and fibroadenoma mammary tumors. There were no effects of atrazine on any other tumor type in male or female SD or Fischer-344 rats or in three strains of mice. Seven key events that precede tumor expression in female SD rats were identified. Atrazine induces mammary tumors in aging female SD rats by suppressing the luteinizing hormone surge, thereby supporting a state of persistent estrus and prolonged exposure to endogenous estrogen and prolactin. This endocrine mode of action has low biological plausibility for women because women who undergo reproductive senescence have low rather than elevated levels of estrogen and prolactin. Four alternative modes of action (genotoxicity, estrogenicity, upregulation of aromatase gene expression or delayed mammary gland development) were considered and none could account for the tumor response in SD rats. Epidemiological studies provide no support for a causal relationship between atrazine exposure and breast cancer. This conclusion is consistent with International Agency for Research on Cancer’s classification of atrazine as “unclassifiable as to carcinogenicity” and the United States Environmental Protection Agency's classification of atrazine as “not likely to be carcinogenic.”

Expression of estrogen receptors in the hypothalamo-pituitary-ovarian axis in middle-aged rats after re-instatement of estrus cyclicity

During reproductive aging female rats enter an anovulatory state of persistent estrus (PE). In an animal model of re-instatement of estrus cyclicity in middle-aged PE rats we injected the animals with progesterone (0.5 mg progesterone/kg body weight) at 12:00 for 4 days whereas control animals received corn oil injections. After the last injection animals were analyzed at 13:00 and 17:00. Young regular cycling rats served as positive controls and were assessed at 13:00 and 17:00 on proestrus. Progesterone treatment of middle-aged PE rats led to occurrence of luteinizing hormone (LH), follicle stimulating hormone (FSH), and prolactin surges in a subset of animals that were denoted as responders. Responding middle-aged rats displayed a reduction of ER-β mRNA in the preoptic area which was similar to the effect in young rats. Within the mediobasal hypothalamus, only young rats showed a decline of ER-α mRNA expression. A decrease of ER-α mRNA levels in the pituitary was observed in progesterone-responsive rats and in young animals. ER-β mRNA expression was reduced in young regular cycling rats. ER-β mRNA levels in the ovary were reduced following progesterone treatment in PE rats and in young rats. Taken together our data show that cyclic administration of progesterone reinstates ovulatory cycles in intact aging females which have already lost their ability to display spontaneous cyclicity. This treatment leads to the occurrence of preovulatory LH, FSH and prolactin surges which are accompanied by differential modulation of ERs in the hypothalamus, the pituitary and the ovary.

Increases in Plasma Concentration of Progesterone by Protease-Sensitive Urinary Pheromones in Female Rats

Plasma progesterone concentrations in female Wistar rats after exposure to urine preparations with and without protease-treatment were measured to explore the effects of protease-sensitive pheromones on the endocrine state. Exposure to crude urine excreted from male rats induced an increase in the plasma progesterone concentration in female rats. The progesterone concentration of oestrous females increased with an increase in the protein concentration in urine samples. Exposure of females in the oestrous state to urine preparations treated with protease did not induce increases in plasma progesterone. These results suggest that the presence of a protease-sensitive component in male urine exerts an influence on the endocrine state of oestrous females.

Can female–female stimulation of breeding condition occur in dairy goats?

The social stimulation of oestrus by oestrous females (i.e., the ‘female–female effect’) was investigated in dairy goats under typical pasture-grazing conditions in New Zealand. Five anoestrous does were grouped with either one doe treated with hormones to induce oestrus ( n=10 groups), or one doe which received a sham hormone treatment ( n=10 groups). The occurrence and frequency of three activities (vocalising, tail wagging and urinating), that reveal an oestrous state when performed frequently, were recorded in each group. Progesterone concentrations in milk and vulval swelling were also measured to determine whether a female–female effect had occurred. No female–female effect occurred in this study. However, vocalisations and tail wagging were accurate oestrus indicators, and vocalisations associated with induced oestrus were socially facilitated within groups. Therefore, group-mates were attentive to changes in the behaviour of oestrous females, which in turn changed their behaviour, but the change did not appear to influence the groups’ reproductive state.

Social regulation of reproduction in the female mound-builder mouse ( Mus spicilegus)

Social environment influences the reproductive physiology and sexual behaviour of the female house mouse Mus musculus. An all-female environment tends to suppress the oestrous cycles, whereas the presence of a male induces and synchronises sexual receptivity. However, reproductive responses to social environment may differ among the various species of rodents. In mound-builder female mice, Mus spicilegus, periods of sexual receptivity are interrupted by periods where adult females display a vaginal closure. We investigated the influence of different social environments on the vaginal opening and oestrous state of adult female M. spicilegus. Result showed that when females were grouped their vaginas were generally closed but that vaginal opening occurred when they were isolated or housed with a sexual partner. Females became sexually receptive when housed with a male, but when isolated their cervical smears did not reach characteristics of the oestrus. In female M. spicilegus, male presence thus has a stimulating effect on oestrous induction. Furthermore, cohabitation with females has an inhibiting effect on vaginal opening.

Oestrous state dynamics in chemical communication by captive female Asian elephants

In many mammals, reproductive status is revealed through chemical cues in urine. The reproductive status of receivers may influence their interest in such signals. For social mammals that live in matrilineal groups, females may benefit by detecting the reproductive condition of herdmates. Responses to urine during oestrous cycles of senders and receivers are potential indicators of signal functions. We examined the chemosensory responses, first by four captive female Asian elephants,Elephas maximus , over their oestrous cycles to familiar follicular and luteal phase urine and second by 14 different female Asian elephants to unfamiliar conspecific follicular and luteal phase urine. We asked whether females could distinguish the reproductive state of another female as measured by their differential response to luteal- and follicular-phase urine. We further examined the influence of the receiver's reproductive status on response levels. Females responded more with specific tactolfactory trunk behaviours to follicular- than to luteal-phase urine, but only when the receiving female was in her follicular phase. Like their male conspecifics, Asian elephant females can detect changes in the reproductive state of conspecifics. The functional significance of this ability has yet to be determined but may be related more to the resource holding power of females in follicular phase than to a means for females to synchronize oestrous cycles. Such female–female communication may have important effects on social group dynamics. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Sexual aggression in mice: effects of male strain and of female estrous state.

We have observed that 50% of FVB/NtacfBR ("FVB") males were aggressive toward females in tests of mating behavior. We decided to gather basic evidence for an effect of genotype on this behavior by testing for strain differences between FVB and C57BL/6J ("B6") male mice. Also, hypotheses developed from theoretical work on sexually coercive behavior suggest there should be a cycling effect on sexually aggressive behavior (Smuts and Smuts, 1993). We tested for an estrous cycling effect by using stimulus females in a state of either estrus or diestrus. Both strains modulated mating behavior to female cycling state. Cycling state did not modulate B6 aggression. FVB males were more aggressive than B6 males, and cycling state modulated their behavior. We conclude that sexually aggressive male mice modulate aggressive behavior to female social signals and indicate how this data can be used to study the genetics of sexual aggression.

Nocturnal home ranges and social interactions of the brushtailed rock-wallaby Petrogale penicillata at Hurdle Creek, Queensland.

The nocturnal ranges of six male and 15 female adult brush-tailed rock-wallabies (Petrogale penicillata) were calculated from November 2001 to March 2002 at Hurdle Creek, Queensland. Social interactions were recorded during the same period. Nocturnal range data were collected by walking transects with a spotlight, and identifying individuals from their colour-coded reflective eartags. Males? nocturnal home ranges averaged 2.84 + 0.32 ha, while those of females averaged 2.01 + 0.20 ha, using the 100% minimum convex polygon method. Home ranges at this site were thus smaller than those described for this species at other sites. The nocturnal home ranges of males and females overlapped with those of several other individuals of both sexes. There appeared to be three groups within the population who emerged from their diurnal refuges along separate lengths of cliffs, had nocturnal ranges that overlapped highly with those of their own group and associated more often with members of their own group than with those of others, indicating social structuring within the population. Males and females associated with and had sexual interactions with several different partners, and there was no indication of long-term guarding of females by males at night. However, males may have monitored the oestrus state of females during the day and guarded females at night only during their oestrus periods.

Site-specific decrease of progesterone receptor mRNA expression in the hypothalamus of middle-aged persistently estrus rats

Middle-aged females gradually become acyclic and spontaneously develop a persistently estrus (PE) state. PE rats, acyclic for 30 days (early PE), are unresponsive to the positive feedback action of estrogen, but respond to a progesterone challenge with a luteinizing hormone (LH) surge and ovulation; unlike long-term PE rats, acyclic for 90 days, neither estrogen nor estrogen plus progesterone will elicit an LH surge [10th International Congress of Endocrinology, San Francisco, P3 (1996) 1061]. We hypothesize that the PE state may develop due to a diminished level of estrogen-induced progesterone receptor (PR) expression in the hypothalamus that prevents progesterone from stimulating LH regulating circuits. To test this hypothesis, PR mRNA levels were measured in hypothalamic regions of young, proestrus (2–3 months of age), early PE (10–12 months) and long-term PE (13–15 months) rats. The anteroventral periventricular nucleus (AVPV), an important regulatory site of the LH surge, had decreased PR mRNA levels in early and long-term PE rats compared with proestrus rats. However, PR mRNA levels were reduced only in long-term PE rats in the ventromedial nucleus (VMH) and arcuate nucleus (ARH). In the medial preoptic nucleus (MPN), levels of PR mRNA did not change. A previous report showed that exogenous progesterone stimulates an LH surge in young and early PE animals, indicating that the expression of PR mRNA demonstrated in this study is sufficient to mediate progesterone facilitation of the LH surge in early PE rats. In acyclic, long-term PE rats, diminished estrogen-induced expression of progesterone receptors is correlated with a previously shown inability to respond to exogenous progesterone.

Comparative estrogenic effects of p-nonylphenol by 3-day uterotrophic assay and female pubertal onset assay

Nonylphenol (NP) is widely used as a component of detergents, paints, pesticides, and many other formulated products. Several studies have demonstrated that NP is estrogenic in fish, avian, and mammalian cells. NP also competitively inhibits the binding of 17β-estradiol (E2) to the estrogen receptor (ER). However, there are relatively few in vivo data related to this issue in mammals. The aim of this study was to investigate the estrogenic activity of NP in animal models. We performed a 3-day uterotrophic assay using immature female rats for comparison with other endpoints of Tier I screening including vaginal opening (VO) in prepubertal intact female rats. For the uterotrophic assay, diethylstilbestrol (DES) (0.2 and 1.0 μg/kg) and p-NP (10, 25, 50, 100, and 200 mg/kg) were administered subcutaneously to immature Sprague–Dawley female rats for 3 consecutive days (postnatal days (PND) 20, 21, and 22). For the female pubertal onset assay, DES (0.2, 1.0, and 5.0 μg/kg) and p-NP (10, 50, and 100 mg/kg) were administered daily by oral gavage from 21 days of age for 20 days. In the uterotrophic assay, statistically significant increases in uterine wet weight were observed at doses of 100 and 200 mg/kg p-NP. DES (0.2 and 1.0 μg/kg) also significantly increased uterine weight compared to the vehicle control. In the female pubertal onset assay, the age of VO was advanced following oral exposure to DES (1.0 and 5.0 μg/kg) and p-NP (50 and 100 mg/kg). Estrous cyclicity was monitored in prepubertal rats from the day of VO to the day of necropsy. Irregular estrous cycles were observed in the groups treated with DES (5.0 μg/kg) and p-NP (50 and 100 mg/kg). High-dose DES (5.0 μg/kg) produced a persistent estrus state, whereas p-NP (50 and 100 mg/kg) increased the number of days in diestrus. Serum thyroxine (T 4) concentrations were decreased in a dose-dependent manner by DES and p-NP treatment. A significant decrease in serum T 4 level was observed at high-dose DES (5.0 μg/kg) and p-NP (100 mg/kg). Serum TSH level was significantly increased by DES (5.0 μg/kg) treatment. Statistically significant decreases in ovarian weight were observed in female rats treated with DES (5.0 μg/kg) and p-NP (100 mg/kg). Our data demonstrate that p-NP can accelerate the onset of puberty and alter estrous cyclicity in prepubertal female rats at oral doses lower than the subcutaneous doses typically used in the uterotrophic assay. We therefore suggest that the female pubertal onset assay may be used as a sensitive testing method to detect environmental agents with weak estrogenic activity, but requires further research.

Evaluation of the 20-day pubertal female assay in Sprague-Dawley rats treated with DES, tamoxifen, testosterone, and flutamide.

The Endocrine Disrupter Screening and Testing Advisory Committee (EDSTAC) has recommended the rodent pubertal female assay as a Tier I test to detect potential endocrine disrupters (EDs). This assay is designed to screen estrogenic activity in immature rats exposed to chemicals during sexual maturation. The aim of this study was to evaluate whether this assay can detect the EDs with effects brought about through various mechanisms. Immature Sprague-Dawley female rats (21 days of age) were dosed daily for 20 days by oral gavage (DES, tamoxifen, and flutamide) or sc injection (testosterone). The mean age at vaginal opening (VO) was 32.3 +/- 0.5 days in control rats. Although VO was unaffected by DES at doses of 0.2 and 1.0 microg/kg, a high dose of DES (5.0 microg/kg) significantly advanced the age at VO to 24 days. Both tamoxifen (50 and 200 microg/kg) and flutamide (25 mg/kg) also significantly accelerated VO to 27.8 +/- 0.5, 25.1 +/- 0.1, and 26.1 +/- 0.1, respectively. However, testosterone dose-dependently delayed VO (exposure to 1.0 mg/kg extended VO to 37.3 +/- 0.8 days, and VO did not occur in 2 of 10 animals by the time of necropsy at 41 days of age). Estrous cyclicity was monitored in rats from VO to necropsy. Irregular cycles were observed in the groups treated with DES (5.0 microg/kg), tamoxifen (200 microg/kg), testosterone (1.0 mg/kg), and flutamide (25 mg/kg). High dose of DES showed a persistent estrus state throughout the entire observation period. In addition, the number of days in diestrus was increased by tamoxifen (200 microg/kg) and flutamide (25 mg/kg) treatments. Significant decreases in ovarian weight were observed in 5.0 microg/kg DES (64% of control), 25 mg/kg flutamide (76% of control), and 200 microg/kg tamoxifen (47% of control). Testosterone also significantly decreased the ovarian weights in all treatment groups. Uterine weights were also decreased significantly at high doses of tamoxifen (200 microg/kg, 39% of control) or testosterone (1.0 mg/kg, 47% of control). In hormone analysis, tamoxifen significantly increased serum E(2) levels at 50 microg/kg. The mean serum levels of TSH were significantly increased in tamoxifen (10 and 50 microg/kg), testosterone (0.2 mg/kg), and flutamide (1.0 and 25 mg/kg) treatment groups compared with the control. However, serum T(4) levels were significantly reduced by testosterone. Furthermore, serum T(3) levels were significantly increased in DES, tamoxifen (10 and 50 microg/kg), testosterone (1.0 mg/kg), and flutamide (1.0 and 5 mg/kg). Our data demonstrate that the rodent pubertal female assay is useful for identifying potential EDs having not only estrogenic/antiestrogenic but also androgenic/antiandrogenic activities. However, further validation study is necessary to identify chemicals that operate through other action mechanisms, including steroid biosynthesis inhibitors and thyroid inhibitors. Moreover, additional data on other compounds with weak endocrine disrupting activity will be required to further characterize the sensitivity of the female pubertal assay.

Male–female interactions and socio-sexual isolation of Mus spicilegus toward two other species of east-European mice, M. macedonicus and M. m. musculus

Social and sexual interactions were analysed as potential behavioural processes assessing reproductive isolation between the mound-building mouse, Mus spicilegus, and two other wild species, M. macedonicus and M. musculus. Dyadic encounters among strange M. spicilegus individuals and between M. spicilegus and the two other species members were conducted with females in an oestrus state. In intraspecific M. spicilegus encounters, there was a high incidence of close contact behaviours initiated by the females. In interspecific encounters M. musculus males and females and M. macedonicus females showed social interest toward their M. spicilegus protagonists, whereas the later behaved in an unfriendly or aggressive manner. Thus socio-sexual interactions shown by the different species seem to be sufficient to prevent any attempt to mate interspecifically. Arguments in favour of recognition against reinforcement hypothesis are discussed.

The influence of maturation and gender on the anti-arrhythmic effect of ischaemic preconditioning in rats.

The aim of this study was to investigate the influence of maturation and gender on the anti-arrhythmic effect of myocardial ischaemic preconditioning in rats. Coronary artery occlusion was carried out in either rats anaesthetised with sodium pentobarbitone or in rat isolated hearts. Cardiac arrhythmias occurring in the 30 min post-occlusion period were assessed. In anaesthetised 3 month (m) old male rats ischaemic preconditioning, with a 3 min temporary coronary artery occlusion, significantly reduced the total number of ventricular ectopic beats (VEBs) from 2074 +/- 206 to 490 +/- 139 and the incidence of ventricular fibrillation (VF) from 40 to 0% during a subsequent 30 min occlusion (P < 0.05). In middle-aged male rats (16 m) the anti-arrhythmic effect of preconditioning was unaltered (VEBs were reduced from 1958 +/- 121 to 245 +/- 66 and VF from 70 to 0%). In 3 m old anaesthetised female rats the effect of ischaemic preconditioning was also evident (VEBs reduced from 961 +/- 170 to 154 +/- 48; P < 0.05). In non-preconditioned age-matched female animals the total number of VEBs (961 +/- 170), VF (0%) and mortality (0%) were significantly (P < 0.05) lower than in respective male animals. In female rats, attenuation of ischaemia-induced arrhythmic severity was most pronounced in the oestrus state. In hearts isolated from weight-matched male and female rats the incidence of ventricular tachycardia (81 vs 25%) and the total number of VEBs (351 +/- 73 vs 81 +/- 50) were significantly (P < 0.05) different. It is concluded that in rats neither maturation nor gender influence the anti-arrhythmic effect of ischaemic preconditioning. However, female rats exhibit a lower level of arrhythmic activity during sustained coronary artery occlusion than male rats both in vivo and in vitro.

Potential adverse effects of phytoestrogens.

Evaluation of the potential benefits and risks offered by naturally occurring plant estrogens requires investigation of their potency and sites of action when consumed at natural dietary concentrations. Our investigations have examined the effects of a range of natural dietary concentrations of the most potent plant isoflavonoid, coumestrol, using a rat model and a variety of estrogen-dependent tissues and endpoints. Treatments of immature females demonstrated agonistic action in the reproductive tract, brain, and pituitary at natural dietary concentrations. Experiments designed to test for estrogen antagonism demonstrated that coumestrol did not conform to the picture of a classic antiestrogen. However, coumestrol did suppress estrous cycles in adult females. Developmental actions were examined by neonatal exposure of pups through milk of rat dams fed a coumestrol, control, or commercial soy-based diet during the critical period of the first 10 postnatal days or throughout the 21 days of lactation. The 10-day treatment did not significantly alter adult estrous cyclicity, but the 21-day treatment produced in a persistent estrus state in coumestrol-treated females by 132 days of age. In contrast, the 10-day coumestrol treatments produced significant deficits in the sexual behavior of male offspring. These findings illustrate the broad range of actions of these natural estrogens and the variability in potency across endpoints. This variability argues for the importance of fully characterizing each phytoestrogen in terms of its sites of action, balance of agonistic and antagonistic properties, natural potency, and short-term and long-term effects.

Ovarian Carbonyl Reductase Delayed Onset of Persistent Estrus in the Offspring of Parathyroidectomized Mother Rats

We investigated the effects of maternal parathyroidectomy on day 5 of pregnancy on the ovarian carbonyl reductase (CR) in the offspring of rats. Changes in the ovarian CR of the offspring were examined at 4 and 8 months of age. In 8-month-old female offspring, the ovarian weight and the ovarian CR activity were significantly higher in rats from parathyroidectomized mothers than in rats from sham-operated mothers, and the offspring of the parathyroidectomized mothers showed the regular 4-day estrous cycles at 8 months of age, while the offspring of the sham-operated mothers were in the state of persistent estrus. Furthermore, intense immunostaining was found in the theca interna cells and the interstitial gland cells in the ovary of rats from the parathyroidectomized mothers, whereas in the ovary of the age-matched normal rats, immunostaining was faint. These results suggest that the maternal parathyroidectomy affect the activity and localization of CR in the ovary of female offspring.

Changes in plasma lipoprotein during the oestrous cycle of the bitch

Five bitches were kept under controlled conditions of diet and exercise for up to four months. They were monitored at regular time intervals for fasting plasma concentrations of cholesterol, triacylglycerols and apolipoproteins Al and B. Lipoprotein cholesterol and triacylglycerol concentrations were also determined in the fasting plasma. Vaginal cytology and plasma progesterone were monitored at weekly intervals to determine the oestrous state of the bitches. Lipoprotein and apolipoprotein concentrations remained steady during anoestrus but large increases and wide fluctuations were shown in the concentrations of both cholesterol and triacylglycerols during metoestrus. Metabolic studies involving the measurement of lipids in canine blood must take into account the stage of oestrus of any bitches involved.

Control of Candida albicans vaginitis in mice by short-duration butoconazole treatment in situ.

A short-duration treatment for candidal vaginitis applying butoconazole in situ was tested in an experimental mouse model. One week after artificial induction of an oestrus state (by oestradiol benzoate injection), mice were inoculated intravaginally with 1.5 x 10(7) blastospores of Candida albicans (strain ATCC 44858). Treatment consisted of butoconazole solutions (1%, 2.5% and 5%) administered intravaginally. The development of the infection was monitored daily for 12 days and then three times a week by local samples plated on BiGGY agar. In parallel, twice a week, the vaginas of three mice in each group (control and treated groups) were isolated in order to estimate organ invasion by C. albicans. This was assessed by anatomopathological studies on a fixed and stained part of the organ and by serial dilutions of a homogenate of the remaining part plated on Sabouraud glucose chloramphenicol agar. After a 48-h incubation period the number of colony forming units per gram of tissue was counted. For each treatment, a remission of 10-15 days was observed, but was followed by a recurrence for the lower dose of butoconazole; only the higher butoconazole concentrations tested (2.5% and 5%) gave an apparent full cure for most of the mice tested. The use of the mouse model of candidal vaginitis confirms that short-duration treatment is possible when elevated doses of butoconazole are used.

Social modulation of behavioral reproductive senescence in female rats.

During aging, the lordosis reflex of female rats changes from a state of cyclic lordosis, when the lordosis reflex can be elicited by manual palpation only on proestrus, to a state of constant lordosis (CL), when it can be elicited daily. Social signals from other females altered this pattern of behavioral senescence. Group living decreased the lordosis reflex intensity in both old and young rats and delayed the onset of CL. Long-term group living delayed CL until late in the life span, while temporary group living delayed it only until rats returned to isolation. Long-term group living also changed the temporal relationship between CL and the acyclic, anovulatory state of estrus. Finally, the lordosis reflex intensity, measured by manual palpation, predicted the old rats' receptivity to a male, indicating that constant lordosis is a biomarker for other aspects of behavioral reproductive senescence.

Evolution of permanent estrus displays in hominids

Accounts of the reality of “concealed ovulation" (i.e., concealed estrus) in humans do not stand up to tests supplied by the facts of morphology and physiology when these are viewed in a fully Darwinian perspective of descent. The origin and transformation of morphology and physiology occur together with behaviors. The latter cannot be decoupled from the somatic attributes of organisms. Although there is natural selection of organisms, this success is largely the result of selection for survival strategies, the latter being directly causal to differential reproduction. Reproductive strategies are thus constrained by feeding and locomotion. Historical and functional attributes of the quadrupedal ape ancestor with a temporary, visually oriented, estrus display (TED) channelled the transformation of the sexual signal system of the newly bipedal protohominid. Mechanical constraints of bipedality ushered in this system based on the attractivity of the ancestral vulva-focused estrus swellings, but in a permanent form. Permanent estrus displays (PED) of hominids are a continuation from a chimp-like ancestry of swollen and naked perineal signals. PED, realized when bipedality evolved, occurred through a phyletic automimickry (of the antecedent condition in the lineage) of the swollen labia of estrus by the fatty buttocks and by the spreading of hairless skin. A zoologically unique sexually dimorphic deposition of fat altered the buttocks and female body contours emphasizing the rear, and thus making permanent the signal paraphernalia of the ancestral estrus state. Further spread of such skin onto thighs and torso, and the development of pendulous breasts, in automimicry of the swollen perineal region of the ancestry, and the hominid buttocks, are the results of continuous sexual selection based on a successful somatic and behavioral female survival and reproductive strategy. In this example of inseparable sexual and natural selection, the continuous attractivity of PED females resulted in both feeding advantages obtained through the males, and probable anti-infanticidal strategy against males. This further fueled sexual selection of males as a result of both intermale and female-male selection. Male competition within a complex reconciliatory hierarchy of the group would have increasingly selected for phenotypes with a behavioral ability to deal with tensions created by the PED. Male hair reduction is the hormonally-mediated epiphenomenon of successful female traits. We explore explanations of human attributes in the framework of a historical-narrative explanation of hominid origins.