A kilonova as the electromagnetic counterpart to a gravitational-wave source.

Gravitational waves were discovered with the detection of binary black-hole mergers and they should also be detectable from lower-mass neutron-star mergers. These are predicted to eject material rich in heavy radioactive isotopes that can power an electromagnetic signal. This signal is luminous at optical and infrared wavelengths and is called a kilonova. The gravitational-wave source GW170817 arose from a binary neutron-star merger in the nearby Universe with a relatively well confined sky position and distance estimate. Here we report observations and physical modelling of a rapidly fading electromagnetic transient in the galaxy NGC 4993, which is spatially coincident with GW170817 and with a weak, short γ-ray burst. The transient has physical parameters that broadly match the theoretical predictions of blue kilonovae from neutron-star mergers. The emitted electromagnetic radiation can be explained with an ejected mass of 0.04 ± 0.01 solar masses, with an opacity of less than 0.5 square centimetres per gram, at a velocity of 0.2 ± 0.1 times light speed. The power source is constrained to have a power-law slope of -1.2 ± 0.3, consistent with radioactive powering from r-process nuclides. (The r-process is a series of neutron capture reactions that synthesise many of the elements heavier than iron.) We identify line features in the spectra that are consistent with light r-process elements (atomic masses of 90-140). As it fades, the transient rapidly becomes red, and a higher-opacity, lanthanide-rich ejecta component may contribute to the emission. This indicates that neutron-star mergers produce gravitational waves and radioactively powered kilonovae, and are a nucleosynthetic source of the r-process elements.

An ultraviolet-optical flare from the tidal disruption of a helium-rich stellar core.

The flare of radiation from the tidal disruption and accretion of a star can be used as a marker for supermassive black holes that otherwise lie dormant and undetected in the centres of distant galaxies. Previous candidate flares have had declining light curves in good agreement with expectations, but with poor constraints on the time of disruption and the type of star disrupted, because the rising emission was not observed. Recently, two 'relativistic' candidate tidal disruption events were discovered, each of whose extreme X-ray luminosity and synchrotron radio emission were interpreted as the onset of emission from a relativistic jet. Here we report a luminous ultraviolet-optical flare from the nuclear region of an inactive galaxy at a redshift of 0.1696. The observed continuum is cooler than expected for a simple accreting debris disk, but the well-sampled rise and decay of the light curve follow the predicted mass accretion rate and can be modelled to determine the time of disruption to an accuracy of two days. The black hole has a mass of about two million solar masses, modulo a factor dependent on the mass and radius of the star disrupted. On the basis of the spectroscopic signature of ionized helium from the unbound debris, we determine that the disrupted star was a helium-rich stellar core.

Sexual behaviors of aging female rats: influence of age and hormonal state of male partners.

Proceptive and receptive behaviors were examined in aging and young ovariectomized female rats treated with estradiol benzoate and progesterone. Young females approached males confined in wire mesh cages more frequently and exhibited more proceptive behaviors toward these males than did middle-aged females. Both middle-aged and young females approached young males more often than they approached old males, but young females showed greater preferences for young males and for males with high levels of testosterone than did middle-aged females. Young females also exhibited more proceptive behaviors than did middle-aged and old females during tests of sexual behavior. The frequencies of lordosis responses, however, did not differ among these three groups of females.

Sexual solicitation by middle-aged and old rhesus females.

The sexual behavior of old and middle-aged ovariectomized estradiol treated females was studied in pair tests with old and middle-aged male rhesus macaques (Macaca mulatta). The ages of the subjects (females, N = 21 and males, N = 16) ranged from 9 to 26 years. We found no significant differences between old and middle-aged females in the overall rates of soliciting sexual behavior, and they solicited sexual behavior as readily from old as from middle-aged males. Old males were less active sexually than middle-aged males, but the two groups did not behave differently toward old and middle-aged females. Old and middle-aged females were equally proceptive toward old and middle-aged males and equally attractive to them. Because the ratio of female presents to male contacts did not differ between the old and middle-aged females, the two groups may be considered equally receptive as well. We found no evidence that the capacity to respond sexually to extradiol treatment is diminished in old rhesus females.

Testosterone is more effective than dihydrotestosterone plus estradiol in activating sexual behavior in old male rats.

Sexual behavior declines in old male rats, and testosterone therapy does not restore the behavior to levels found in young males. If as a result of aging, old males have less capacity to aromatize or reduce testosterone, dihydrotestosterone plus estradiol treatment should be more effective than testosterone treatment in restoring sexual behavior in old castrated males. In a test of this hypothesis, the sexual behavior of old (24 months) castrated Fischer 344 males given injections of testosterone propionate (TP) or dihydrotestosterone propionate (DHTP) plus estradiol benzoate (EB) and that of old sham-operated males given injections of vehicle were observed. The DHTP/EB proved to be less effective overall than the TP in increasing sexual behavior in old castrated males. In a second experiment, young (3 months) and old (30 months) males were tested to verify that the reduced effectiveness of DHTP/EB treatment was age-related. Testosterone propionate and DHTP/EB were equally effective in restoring most measures of sexual behavior in young castrated males. In old castrated males, DHTP/EB treatment was no more or less effective than TP treatment in increasing these same measures. Neither hormone increased the behavior of old males to the level found in young males. Since DHTP/EB treatment is less effective than TP treatment in stimulating sexual behavior in old males, a reduced capacity to aromatize or reduce testosterone is not a likely explanation for decreased responsiveness to testosterone in old male rats.

Sexual behavior in response to testosterone in old long-term-castrated rhesus males.

The sexual behavior of 20-year-old long-term-castrated and intact male rhesus macaques (Macaca mulatta) was observed. In the first experiment we compared the sexual behavior of males tested in 1980 with their performance in 1970 (before castration and sham castration) and in 1971 (1 year after the operation). In 1971 and 1980, the castrated males were tested while under testosterone propionate treatment. The castrated and intact males did not differ in any measures of sexual behavior over the 10-year period, but both groups showed a decline in the percentage of tests with intromissions and ejaculations; a decline in the rates of contacting, mounting, and intromitting; and an increase in the latencies to contact, intromit, and ejaculate. In a second experiment, the sexual performances of old castrated and intact males were compared to those of young intact males (8 to 12 years). The intromission rate and the percentage of tests with intromissions were significantly greater in young males than in old intact males, but did not differ from values for old castrated males. The old castrated and old intact males did not differ from each other in these measures. Young males had a higher percentage of tests with ejaculation than either group of old males. We have concluded that old long-term-castrated rhesus males retain the potential to display sexual behavior at levels comparable to those observed in old intact males.

Age-related deficits in brain androgen binding and metabolism, testosterone, and sexual behavior of male rats.

Brain androgen binding and metabolism, serum testosterone (T), and sexual behavior were measured in old and young male Fischer 344 rats. After completion of sexual behavior tests, blood was collected for T assay and brains were removed for simultaneous measurements of cytosolic (ARc) and nuclear (ARn) androgen receptors and aromatase activity (AA) in the preoptic area (POA), hypothalamus (HYP) and amygdala (AMG). In Experiment 1, old and young intact males were examined. None of the old males ejaculated in any of the tests of sexual behavior whereas all of the young males ejaculated. The old males had lower levels of serum T, lower levels of ARn in the POA and HYP and lower levels of AA in the POA. The ARc levels of the old and young males did not differ. Experiment 2 was designed to determine if the deficits in brain androgen binding and metabolism were due to low levels of T. Old and young T-treated gonadectomized (GX-T) males and young intact (I) males were examined. T levels were comparable in the young and old GX-T males and were higher in each of these groups than in the young I males. In sexual behavior tests, all of the young but only 25% of the old GX-T males ejaculated. Although ARn levels in the old GX-T males were lower than in the young GX-T males, they were comparable to the young I male levels. No age-related differences in T induction of AA were observed.(ABSTRACT TRUNCATED AT 250 WORDS)

Correlation of diurnal changes in hormones with sexual behavior and age in male rhesus macaques.

We designed this study to determine whether the sexual behavior of male monkeys changes on a diurnal schedule that coincides with changes in the levels of hormones such as luteinizing hormone (LH), testosterone, estradiol, and cortisol, and whether these differences vary with age. Old (n = 8) and young (n = 8) males were bled and given sexual behavior tests eight times at 0900 and eight times at 2100. The old and young males did not differ in mean serum levels of testosterone, estradiol, cortisol, or LH. Testosterone and LH levels were lower at 0900 than at 2100 (p less than 0.01), and estradiol and cortisol levels were higher at 0900 than at 2100 (p less than 0.01). The young males had higher percentages of tests with erections, mounts, intromissions, and ejaculations than the old males (p less than 0.01). The rates of contacting, mounting, and intromitting were higher at 0900 than at 2100 (p less than 0.05). We failed to confirm previously obtained correlations between hormone levels and sexual performance, This failure led us to conclude that any significant correlation between sexual behavior and hormone levels must be regarded as tentative unless repeated in successive independent studies.

Differences among ovariectomized female rhesus macaques in the display of sexual behavior without and with estradiol treatment.

Ovariectomized, estrogen-primed, adult female rhesus macaques were preselected for the display of high (HLP) and low (LLP) levels of sexual performance. Eight sexually vigorous adult males were selected as partners. For 13 consecutive days each month, females received 0, 2.5, 5.0, and 10.0 micrograms of estradiol benzoate (EB) and were tested on Days 12 and 13 of injection. Blood samples were taken before and on the last day of each of the four injection series, and serum levels of estradiol, progesterone, and testosterone were determined with a radioimmunoassay. The HLP females (n = 5) displayed higher levels of receptivity and proceptivity and received more mounts, intromissions, and ejaculations when tested without EB treatment than LLP females (n = 5) did when they were tested with 10.0 micrograms of EB. Female receptivity, proceptivity, and attractiveness cannot be explained by estrogen action alone. The effectiveness of estrogen is limited by the somatic characteristics of the individual on which it acts.

Apomorphine, deprenyl, and yohimbine fail to increase sexual behavior in rhesus males.

Various doses of apomorphine, deprenyl, and yohimbine were administered to old (20-26 years) rhesus males that had been sexually active when younger and to younger (6-17 years) males that were characteristically sexually sluggish. These neuropharmacological agents have been reported to increase sexual behavior in male rats. In Experiment 1, 10 old intact rhesus males were tested after injection of vehicle and apomorphine, and 6 old testosterone-treated castrated males were tested after treatment with deprenyl and yohimbine and the vehicles for each drug. In experiment 2, the 5 younger males were tested after treatment with each of the drugs and with the vehicles for each drug. There were a few minor changes in behavior associated with certain doses of each of the drugs and as many depressive as facilitative effects on sexual behavior. This suggests that there are basic differences between rats and rhesus macaques in the systems mediating sexual behavior.

Sensitivity of male, female, and androgenized female rats to testosterone during extinction of a conditioned taste aversion.

Experiment 1 tested the hypothesis that male and female rats differ in the amount of testosterone (T) required to prolong extinction of a conditioned taste aversion. Gonadectomized male and female rats were implanted with empty or 30-, 60-, or 120-mm T-filled capsules. The males had slower extinction rates than females when both were implanted with 30-mm and 60-mm capsules but not when implanted with 120-mm capsules. The dimorphic sensitivity was not due to differences in plasma T levels: the levels for males and females were not different. Experiment 2 tested the hypothesis that the presence of T during the perinatal period results in a greater sensitivity to T in adulthood. Females exposed to T during the perinatal period showed prolonged extinction when given a 30-mm T-filled capsule as an adult, whereas unexposed females did not. These results support the hypothesis that the amount of T required to activate the prolonged extinction in an adult depends on perinatal exposure to T.

Cooling the area postrema induces conditioned taste aversions in male rats and blocks acquisition of LiCl-induced aversions.

The experiments presented in this article were designed to examine whether area postrema (AP) lesions attenuate LiCl-induced conditioned taste aversions (CTAs) by disruption of information about the illness-producing properties of LiCl or by a lesion-induced malaise. Reversible lesioning of the AP caused by cooling induced a CTA in male rats. The cooling-induced CTA could be blocked if males were exposed to cooling for several days before acquisition day. Acquisition of a LiCl-induced CTA was blocked in males if they were exposed to cooling before acquisition day and during LiCl administration on acquisition day was attenuated but not blocked in males if they were exposed to cooling only before acquisition day, and was unchanged in males if they were exposed to cooling only during LiCl administration. Taken together these results indicate that the AP is important for acquisition of LiCl-induced CTAs but that inactivation of this area is so aversive it will induce CTAs that can obscure the attenuation of LiCl-induced aversions.

Age-related deficits in brain estrogen receptors and sexual behavior of male rats.

Neural estrogen receptors (ER), serum testosterone (T), estradiol (E2) and luteinizing hormone (LH), and masculine sexual behavior were measured in young (5 months) and old (24 1/2 months) Fischer 344 male rats. We found that old intact males, which displayed significantly lower levels of sexual behavior, T, and LH than young intact males, also had lower levels of nuclear ER (ERn) in the amygdala (AMG). The age difference in ER binding did not appear to be a consequence of altered blood E2 levels because circulating E2 did not differ between the two age groups. Gonadectomy eliminated ejaculatory behavior and significantly reduced ERn in young males. When we administered exogenous T to gonadectomized males in doses that approximated levels found in young intact males, we found that sexual performance of old males was stimulated to precastration levels but not to levels found in young males. Moreover, such treatment failed to increase ERn in the AMG of old males to the levels measured in the AMG of young males. These results suggest that there is an association between the inability of T to increase ERn concentration in the AMG and the deficits in sexual performance that are characteristic of old males. Thus, the capacity of neural tissue to bind estrogen, presumably derived from circulating T, may be a limiting factor in the determination of androgen responsiveness in aging males.

Decreased testosterone levels and accelerated extinction of a conditioned taste aversion in fluid-deprived male rats.

The hypothesis that fluid deprivation accelerates extinction of a conditioned taste aversion in male Sprague-Dawley-derived rats by reducing serum testosterone levels was tested. Serum testosterone levels were found to be lower in fluid-deprived males than in nondeprived males (Experiments 1 and 2). Exogenous testosterone treatment that results in high physiological levels of serum testosterone slowed the extinction of fluid-deprived gonadectomized males to rates comparable with those of nondeprived sham males (Experiment 3). It was noted, however, that testosterone treatment was less effective in slowing extinction in fluid-deprived gonadectomized males than in nondeprived gonadectomized males even though the serum testosterone levels were the same (Experiments 3 and 4). These results provide strong support for the original hypothesis, but they suggest that fluid deprivation also reduces sensitivity to testosterone.

The role of the dura in conditioned taste avoidance induced by cooling the area postrema of male rats.

Experiments were designed to assess the contribution of the dura mater to the formation of conditioned taste avoidance induced by cooling the area postrema. The results of the first experiment verified that the temperature of the dura showed a temperature gradient at various distances from the tip of the cold probe. In the second and third experiments, a circle of dura was cut away so that different amounts of the area postrema could be cooled without cooling the overlying dura. Cooling the dura plus the area postrema did not produce a stronger avoidance than just cooling the area postrema. In the fourth experiment, the cerebellar cortex was cooled with and without cooling the dura. Cooling the cerebellar cortex produced conditioned taste avoidance, and cooling the dura plus the cerebellar cortex did not produce a stronger avoidance. Taken together, these results suggest that cooling the dura mater does not contribute to the conditioned taste avoidance induced by cooling the area postrema. The results of the fifth experiment showed that cooling the area postrema produced a stronger conditioned taste avoidance than cooling the cerebellar cortex. It is suggested that the avoidance induced by cooling both of these structures is the result of physiological changes occurring when neurons in these structures are inactivated and when the subdural meninges are cooled. Furthermore, these changes are more severe when the area postrema is cooled.

The effects of cooling the area postrema of male rats on conditioned taste aversions induced by LiC1 and apomorphine.

Although permanent lesion studies have demonstrated that the area postrema (AP), a chemoreceptor trigger zone, is part of the neural mechanism for conditioned taste aversions (CTAs), its exact role remains questionable. It has been suggested that the attenuated acquisition of a CTA after permanent lesions of the AP is the result of an inability to recognize the conditioned taste as novel. The present series of experiments was designed to test the hypothesis that lesions of the AP interfered with LiCl processing and not recognition of taste novelty. This was accomplished by using the reversible lesioning procedure, cooling, only during administration of the illness-inducing agent. In Expt. 1, measurement of thermal lines around the tip of the cold probe in the AP indicated that our cooling procedures allowed the majority of the AP to be cooled to temperatures that suppress neuronal activity and transsynaptic transmission, but not axonal transmission. In Expts. 2 and 3, rats were injected with either LiCl or apomorphine after consumption of a 10% sucrose solution. Cooling of the AP was initiated 5 min before administration of one of the illness-inducing agents and was continued for 55 min after injection. The rats were tested later for acquisition while the neural function of the AP was preserved. Our experimental results demonstrated that cooling the AP could attenuate the CTA induced by LiCl, but had no effect on the CTA induced by apomorphine. Since the AP was functional when the rats encountered the novel sucrose solution both before and after conditioning, but not functional when LiCl was given, these results do not support the recognition of taste novelty hypothesis.

Failure of ACTH to prolong extinction of a conditioned taste aversion in the absence of the testes.

Both corticotropin (ACTH) and testosterone prolong the extinction of a conditioned taste aversion in water-deprived intact male rats. An investigation was made to determine whether ACTH affects extinction in the absence of the testes and also to determine the effect of ACTH on serum testosterone levels. Water-deprived intact males showed prolonged extinction after ACTH injections; water-deprived gonadectomized males and intact females did not. All three of these groups showed elevated testosterone levels after ACTH administration, but testosterone levels were higher in the intact males than in the gonadectomized males or intact females. These results clearly show that in the absence of the testes ACTH is unable to prolong extinction. It is proposed that the increased level of testosterone following ACTH injection in water-deprived intact males is responsible for the prolonged extinction of a conditioned taste aversion. Although testosterone levels may increase in females and castrated males following ACTH injection, the increase is not sufficient to prolong extinction in these water-deprived animals.

Progesterone, estradiol, testosterone and dihydrotestosterone: effects on rate of extinction of a conditioned taste aversion in rats.

An investigation was made to determine why the prolonged rate of extinction of a conditioned taste aversion induced by testosterone is diminished when the ovarian system is intact. In the first experiment, 36 gonadectomized female rats received injections of progesterone, testosterone propionate (TP), progesterone plus TP, or sesame oil. Progesterone did not reduce the slow extinction rate induced by TP. In a second experiment, 36 gonadectomized female rats received injections of estradiol dipropionate (EP), TP, EP plus TP, or sesame oil. Estradiol dipropionate reduced the effectiveness of TP in prolonging the extinction rate. These same results (the ineffectiveness of progesterone and the effectiveness of EP in blocking TP-induced slow extinction) were also observed in male rats in a third experiment. Dihydrotestosterone, as well as testosterone, has been shown to prolong extinction: hence, in a fourth experiment 30 gonadectomized female rats received injections of EP, TP, dihydrotestosterone propionate (DHTP), EP plus DHTP, or sesame oil. Estradiol dipropionate reduced the DHTP-induced slow extinction. All the above data are consistent with the hypothesis that it is estradiol from the ovaries that diminishes the effect of testosterone on the rate of extinction of a conditioned taste aversion in intact females.

Sexual performance of old and young male rhesus macaques following treatment with GnRH.

GnRH has been reported to facilitate sexual performance in a number of species. To determine whether the same was true for rhesus monkeys we measured sexual behavior and serum levels of testosterone (T) and luteinizing hormone (LH) following control tests and after treatment with two doses of GnRH. In the first experiment, old intact (N = 11) and old T-implanted, castrated (N = 4) rhesus macaques were examined. Mean intromission rate of old intact males was significantly lower following treatment with 100 micrograms of GnRH than following control injections. Other measures of sexual behavior did not differ across treatments. There were no significant treatment effects among the old castrated males. The failure to facilitate sexual performance may have been due to the age of the males and not to the species under study. Thus, in a second experiment the effects of GnRH were examined in young rhesus males. Again, there was no facilitation of sexual performance. This cannot be accounted for by a failure of GnRH to produce a physiological response. For both old and young intact males serum levels of LH and T increased significantly after treatment with both doses of GnRH. LH but not T levels increased significantly in the T-implanted males.

Blockage of the effects of testosterone on extinction of a conditioned taste aversion by estradiol: time of action.

Testosterone (T) prolongs the extinction of a conditioned taste aversion only if it is present during extinction. Experiments were conducted to determine whether estradiol (E) blocks the effects of T by acting during acquisition or extinction. In the first experiment, gonadectomized male and female rats injected with estradiol dipropionate (EP) and testosterone propionate (TP) during extinction had significantly faster extinction rates than those only injected with TP. Treating gonadectomized rats with TP prior to as well as during extinction did not prevent EP from blocking the effects of T. In Experiment 2, E was equally effective in preventing T from prolonging extinction when it was implanted in gonadectomized males during acquisition, extinction, or both acquisition and extinction. Thus, E does not have to be present concurrently with T during extinction to be effective. This suggests that E does not act on a T-related mechanism but rather acts independently of T.