Neurogenesis and sexual behavior.

Different conditions induce proliferation, migration and integration of new neurons in the adult brain. This process of neurogenesis is a clear example of long lasting plastic changes in the brain of different species. Sexual behavior is a motivated behavior that is crucial for the survival of the species, but an individual can spend all his life without displaying sexual behavior. In the present review, we briefly describe some of the effects of pheromones on neurogenesis. We review in detail studies describing the effects of sexual behavior in both males and females on proliferation, migration and integration of new cells and neurons. It will become evident that most of the studies have been done in rodents, assessing the effects of this behavior on neurogenesis within the dentate gyrus of the hippocampus and in the subventricular zone - rostral migratory stream - olfactory bulb system.

Subtropical goats ovulate in response to the male effect after a prolonged treatment of artificial long days to stimulate their milk yield.

The objectives of this study were to determine (i) if in subtropical goats that gave birth during mid-December, the exposition to an artificial long-day photoperiod consisting in only 14 hr of light per day can increase the milk yield and (ii) to test whether these females can respond to the male effect at the end of the prolonged photoperiodic treatment. In experiment 1, 17 lactating goats were maintained under natural short days (control group), while another 22 goats were maintained under artificial long days (treated group) consisting in 14 hr light and 10 hr darkness starting at day 10 of lactation. The continuous exposition to an artificial long-day photoperiod produced an increase in the milk yield level during the first 110 days of lactation (time × treatment interaction; p = .01), while none of the milk components were modified due to the photoperiodic treatment (p > .05). In experiment 2, all control and treated anovulatory goats were submitted to the male effect using photostimulated males. All females showed oestrous behaviour within the first 10 days that were in contact with males (100% in both groups; p > .05). Thus, the latency to onset of oestrus did not differ between females from control (58.2 ± 3.0 hr) and treated (62 ± 4.6 hr) groups. Male exposition provoked ovulation independently if females were previously under long days or natural photoperiod (96 vs 100%, respectively; p = .79). It was concluded that exposure to 14 hr of light per day in subtropical goats that gave birth in late autumn stimulates milk yield without preventing the ovulation in response to the male effect at the end of the prolonged photoperiodic treatment.

Four hours of daily contact with sexually active males is sufficient to induce fertile ovulation in anestrous goats.

The study was conducted on two consecutive years to determine whether ovulatory activity can be induced in anovulatory goats by exposing them to sexually active bucks for 4, 8, 12 or 16 h per day during 15 consecutive days. In experiment 1, females remained continuously in the experimental pens where they were in contact with males. One group remained isolated from males (controls) and four other groups were exposed to sexually active males for 4, 8, 12 or 16 h per day. In experiment 2, females were taken away to "resting" pens free of male odours between the periods of contact with bucks. They were allocated to 5 groups as in experiment 1. Ovulations were determined by progesterone plasma levels and transrectal ultrasonography. Pregnancy was determined by abdominal ultrasonography. In both experiments, more than 90% of females exposed to the bucks had at least one ovulation during the whole experiment whereas only 11 or 0% (experiments 1 and 2, respectively) did so in the control group (P<0.001). Furthermore, the proportion of females ovulating did not differ among groups depending on duration of contact with bucks (P>0.05). In both experiments, pregnancy rates were not affected by the daily duration of contact with males (P>0.05). To conclude, 4h of daily contact with sexually active males is sufficient to stimulate ovulatory activity in anovulatory goats and this effect is not due to the presence of olfactory cues from the males remaining in the pens.

Undernutrition reduces the body weight and testicular size of bucks exposed to long days but not their ability to stimulate reproduction of seasonally anestrous goats.

In semiarid conditions, feed is often scarce and variable with underfeeding being common; these factors can potentially induce fertility reductions in both sexes. Sexually active bucks are able to very efficiently fertilize out-of-season goats, but we do not know whether underfeeding would reduce the ability of bucks to fertilize goats during these periods. Two experiments were conducted to determine (i) testicular size and change of odor intensity of undernourished bucks exposed to long days and (ii) the ability of these bucks to stimulate reproductive activity in seasonally anestrous goats. In experiment 1, bucks (n = 7) were fed 1.5 times the normal maintenance requirements from September to May and formed the well-fed group. Another group of bucks (n = 7) were fed 0.5 times the maintenance requirements and formed the undernourished group. All bucks were subjected to artificially long days from 1 November to 15 January; this period was followed by a natural photoperiod until 30 May. Body weight, scrotal circumference and male odor intensity changes were determined every 2 weeks. In experiment 2, two groups of female goats (n = 26 each) were exposed to well-fed (n = 2) or undernourished bucks (n = 2) on 31 March. Ovulations and pregnancy rates were determined by transrectal ultrasonography. In experiment 1, a treatment by time interaction was detected for BW, scrotal circumference and odor intensity changes (P < 0.001). The BWs of well-fed bucks were greater than those of the undernourished bucks from October to May (P < 0.01), as were the scrotal circumferences from December to March (P < 0.05) and odor intensities from February to May (P < 0.05). In experiment 2, the proportions of females that ovulated at least once (100% v. 96%) or those that were diagnosed as pregnant (85% v. 77%; P > 0.05) did not differ significantly between the goats exposed to well-fed or undernourished bucks. The interval between the introduction of bucks and the onset of estrous behavior was shorter in goats exposed to well-fed bucks compared to the interval for those goats exposed to undernourished bucks (2.5 ± 0.2 v. 9.5 ± 0.6 days; P < 0.05). We conclude that undernourishment reduces the testicular size and odor intensity responses in bucks exposed to long days, but that undernourished bucks are still able to stimulate reproductive activity in seasonally anestrous goats, as is also the case for well-fed bucks.

Presence of a sexually active goat buck enhances ovulation occurrence in seasonally anestrous does after ovulation and luteolysis induction in hormonally-treated goats in seasonal anestrus.

In seasonally anestrous goat does, ovulations can be induced by combining a treatment regimen including progestagen, eCG and prostaglandins. Nonetheless, ovulations occur only once and then does return to a seasonally anestrous state. This study was performed to determine whether the presence of a sexually active buck can stimulate a second ovulation after induced luteolysis using prostaglandins following the first ovulation. Three groups of seasonally anestrous does were treated to induce ovulations using an intra-vaginally inserted sponge containing a progestin combined with eCG and prostaglandin administrations. Goats that had ovulations were treated with a prostaglandin 11 days after progestin sponge removal. After the prostaglandin injection, does continued to be isolated from bucks (n = 8), were penned with a control buck (n = 9), or were penned with a sexually active buck (n = 10). The proportion of goats having ovulations after imposing the ovulation-induction protocol was greater than 80% and did not differ among treatment groups (P >  0.05). The proportion of does having ovulations after injecting prostaglandins was greater when does were penned with a sexually active buck (8/10) than does penned with a control buck (0/9) or that were isolated from bucks (0/8; P < 0.05). It is concluded that in seasonally anestrous goat does induced to have ovulations using a hormonal treatment regimen, the presence of a sexually active buck can induce a second ovulation when there is an induced luteolysis.

Fifteen minutes of daily contact with sexually active male induces ovulation but delays its timing in seasonally anestrous goats.

The present experiment was conducted to determine (1) whether the sexually active bucks are able to stimulate the ovulatory activity of seasonal anestrous goats when the daily contact is reduced to 15 min/day during 15 days and (2) the exact ovulatory activity in anestrous goats exposed to bucks for 2 hours or less per day during 15 days. One group of goats (n = 15) was isolated from bucks. The other six groups (n = 15 each) were exposed to sexually active bucks (n = 1 each) for 15 minutes, 30 minutes, 1 hour, 2 hours, 4 hours, or 24 hours during 15 days. Goats with plasma concentrations of progesterone greater than 0.5 ng/mL were considered to have ovulated. More than 93% of females exposed to bucks ovulated throughout the experiment regardless of the duration of contact with males, whereas none of them ovulated in the isolated group (P < 0.0001). The proportions of females that ovulated at least once did not differ among groups as well as the proportions of goats that displayed normal or short ovulatory cycles. The interval between the introduction of males and the first ovulation did not differ among groups of goats in contact with bucks for 15 minutes, 30 minutes, 1 hour, or 2 hours. However, in these groups, this ovulation occurred about 2 days later than in females in contact with males during 4 or 24 hours (6.3 vs. 4.4 days; P < 0.05). We conclude that 15 minutes of daily contact with sexually active buck is sufficient to stimulate the ovulatory activity in seasonally anestrous goats, but the first ovulation is delayed in goats exposed to males for 2 h/day or less during 15 days in comparison with those in contact with males for 4 or 24 h/day.

Sexually active bucks counterbalance the seasonal negative feedback of estradiol on LH in ovariectomized goats.

We showed previously that the permanent presence of bucks rendered sexually active by photoperiodic treatments, thereafter called photostimulated bucks, prevents the occurrence of seasonal anovulation; also, the introduction of these sexually active bucks induces ovulations during seasonal anestrus. Here, we studied the response of ovariectomized goats bearing 12-mm subcutaneous implants filled or not with estradiol to sexually active males to determine (1) whether the permanent presence of such bucks prevents the decrease of LH despite the presence of a negative feedback by estradiol mimicking that of seasonal anestrus (experiment 1) and (2) whether the introduction of photostimulated bucks increases the plasma LH concentrations in spite of this negative feedback (experiment 2). In experiment 1, one group of goats remained in contact with sexually active bucks, whereas the other group remained in contact with control bucks. Plasma LH concentrations were high and did not differ with time or between groups of females from November to February (P > 0.05), when both types of bucks were sexually active. Afterward, in goats in contact with control and sexually inactive bucks, LH concentrations decreased from March (P ≤ 0.01) and remained low until May, whereas LH levels remained high from March to May in goats in contact with the photostimulated bucks (P > 0.05). In experiment 2, 2 groups of females bearing empty subcutaneous implants, and 2 groups of goats bearing subcutaneous implants filled with estradiol, were exposed to control or photostimulated bucks. Plasma LH concentrations did not increase in goats bearing empty implants, when exposed to control or photostimulated bucks (from 2.01 ± 0.26 to 1.98 ± 0.31 ng/mL, and from 2.45 ± 0.29 to 2.42 ± 0.21 ng/mL respectively; P > 0.05). In contrast, plasma LH concentrations increased from 0.97 ± 0.41 to 2.80 ± 0.62 ng/mL in goats exposed to the photostimulated bucks and bearing estradiol implants (P < 0.05). Thus, the permanent presence of sexually active bucks prevented the decrease of plasma LH concentration in OVX + E2 goats during the seasonal anestrus, and the introduction of the photostimulated bucks increased the plasma LH concentrations in OVX + E2 goats during the seasonal anestrus. Therefore, we conclude that in both cases, the photostimulated bucks are able to reduce or counterbalance the seasonal negative feedback of estradiol on LH secretion.

Efficiency of the male effect with photostimulated bucks does not depend on their familiarity with goats.

In ewes, the ovulatory response of females exposed to familiar rams is lower than the response of those exposed to novel ones. In goats, males rendered sexually active by exposure to long days are more efficient to induce ovulation in seasonal anestrous females than untreated males. Two experiments were conducted to determine 1) whether male goats remain familiar to females after 45days of separation; and 2) whether photostimulated males are able to stimulate the sexual activity of females, independently of their familiarity with them. In Experiment 1, three groups of goats (n=10 goats per group) were put in contact with males (n=2 per group) during 10days in November (familiarization period). These males were called familiar males. After 15, 30 and 45days of separation from the males, females of each group were exposed to familiar or novel males during 10min. In each test, goats in contact with novel males displayed more distress bleats, escapes, head butts, and sniffing than those in contact with familiar males (P<0.05). In Experiment 2, we used sexually inactive (n=4 control males), and sexually active males (n=4 photostimulated males). In February, two groups of goats (n=50 each) were put in contact with control or photostimulated males (n=2 each) during 10days ("familiar" control or photostimulated male, respectively). After 45days of separation from the males, both groups of females were further divided into two groups (n=25 goats per group). In April, two groups were re-exposed to "familiar" control or "familiar" photostimulated males (n=2 per group), whereas the other two groups were exposed to "novel" control or "novel" photostimulated males (n=2 per group). The photostimulated males displayed a higher level of sexual behavior than the controls. The proportion of goats that ovulated and displayed estrus was higher when exposed to the photostimulated males than when exposed to control ones (≥80% vs. 0%; P<0.05). These proportions did not differ between groups exposed to familiar or novel photostimulated males (P>0.05). We concluded that after 45days of separation, males are still familiar to females. The photostimulated males are able to induce the sexual activity of seasonally anestrous goats independently of their familiarity with them.

Switching photo-stimulated males between groups of goats does not improve the reproductive response during the male effect.

We aimed to determine whether the daily exchange of photo-stimulated males among subgroups of females improved the reproductive response of anestrous goats exposed to males. Bucks were rendered sexually active during the rest season by exposure to 2.5 months of long days from November 1st. In April, males (n=3) were put in contact with three subgroups of anestrous goats (one male per 12 females) where they remained throughout the study, constituting the fixed-group. Other males (n=3) were put in contact with three subgroups of females (one male per 11-12 females) and were rotated daily among them, constituting the rotated-group. The sexual behavior of all males was registered from 08:00 to 09:00 on days 0, 1, 2, and 8 after exchanging the males from the subgroups of females. Ovulation and pregnancy rates were determined by transrectal ultrasonography. The occurrences of ano-genital sniffing, nudging (days 1, 2, and 8), and mounting attempts (days 2 and 8) were greater in the rotated than in the fixed-group (P<0.01). The proportions of females that ovulated did not differ among goats from the fixed (92%) and rotated-group (94%; P>0.05). The proportion of pregnant females and the fertility at kidding did not differ between those from the rotated (79% and 59%) and fixed-group (83% and 61%; P>0.05). We conclude that the daily exchange of photo-stimulated males among subgroups induced an increase of their sexual behavior, but does not improve the pregnancy rates in seasonal anestrous goats.

Two or 24 h of daily contact with sexually active males results in different profiles of LH secretion that both lead to ovulation in anestrous goats.

Two experiments were conducted to (a) determine whether sexually active males are able to stimulate the sexual activity of anestrous female goats when duration of contact is reduced to an intermittent contact shorter than 4 daily hours and (b) compare the pattern of secretion of LH when anestrous goats are exposed either permanently or intermittently to males. In the first experiment, 4 groups of anovulatory goats were exposed to sexually active males for 24, 4, 2, or 1 h/d during 15 consecutive days, whereas control females remained isolated. More than 89% of females in the groups exposed to the sexually active bucks ovulated, whereas only 5% did so in the control group (P < 0.001). However, the proportion of females ovulating before day 4 was greater in the 2-, 4-, or 24-h contact groups than in the control, whereas it did not differ between the control group and the 1-h contact group (P = 0.02, <0.001, <0.001 and 0.23, respectively). In the second experiment, 3 groups of anovulatory goats were exposed permanently (24 h/d) or intermittently (2 h/d) to bucks during 5 d or remained isolated. We found that pulsatility of luteinizing hormone (LH) increased in the intermittent and permanent contact groups after males were introduced to females (P = 0.05); this pulsatility of LH remained elevated in the permanent-contact group, whereas it decreased in the intermittent-contact group, once the male was removed (P = 0.32 and 0.05, respectively). We conclude that 1 or 2 daily hours of contact with sexually active males is sufficient to stimulate ovulatory activity in anovulatory goats; however, ovulation is obtained through a different pattern of secretion of LH.

Reducing exposure to long days from 75 to 30 days of extra-light treatment does not decrease the capacity of male goats to stimulate ovulatory activity in seasonally anovulatory females.

The response of male goats exposed to different durations of long days (LD) during an extra-light treatment in autumn-winter, and their ability to induce ovulations in seasonally anovulatory goats were investigated in 2 experiments. In experiment 1, control males were exposed to natural photoperiod (n = 5), whereas 4 additional groups (n = 5/group) were exposed to 16 h of light per d during 75, 45, 30, or 15 d of LD. In the 4 groups, photoperiodic treatments ended on January 15th. Plasma concentrations of testosterone were determined in blood samples obtained once a week from October 15th to May 30th. The rise of testosterone levels occurred earlier in males from the 75-LD and 45-LD groups than in those from the 30-LD, 15-LD, and control groups (P < 0.05). In addition, the time during which levels of testosterone remained >5 ng/mL was longer in males from the 75-LD and 45-LD than in those from the 30-LD and 15-LD groups (P < 0.05). In experiment 2, a group of anovulatory goats (n = 13) was isolated from males, while 3 additional groups were put in contact during 15 d with males previously exposed to 75, 45, or 30 days of LD (n = 25, 27, and 26 females/group, respectively and n = 3 males per group). The proportion of goats that ovulated was higher in the 3 groups in contact with the photo-stimulated males (range: 88%-92%) than in the group isolated from them (0%; P < 0.05). The proportion of pregnant females did not differ between the 3 groups of does in contact with photo-stimulated males (range: 78%-92%; P > 0.05). We conclude that, in our experimental conditions, a photoperiodic treatment as short as 30 d of LD during autumn-winter, stimulated testosterone secretion of bucks during their period of sexual rest and rendered them able to induce ovulations in seasonal anestrous goats and to obtain pregnancies in these females.

Importance of intense male sexual behavior for inducing the preovulatory LH surge and ovulation in seasonally anovulatory female goats.

The present study was carried out to determine whether the presence of photostimulated sedated male goats could stimulate the LH preovulatory surge and ovulation in seasonal anestrous goats. Sexually experienced male goats were treated with artificial long days (16 hours light per day) from 1 November to 15 January to stimulate their sexual activity in March and April, corresponding to the natural sexual rest. A female group of goats (n=20) was exposed to non-sedated males who displayed an intense sexual behavior and provided strong odor (non-sedated group). Another female group of goats (n=20) was exposed to the photo-stimulated male goats, but these males were sedated with Xylazine 2% to prevent the expression of sexual behavior (sedated group). The sedated males also provided a strong odor. Females of both groups had full physical and visual contact with non-sedated or sedated males. In both groups, the males remained with females during 4 days. The LH preovulatory surge of 10 female goats per group was measured by determination of LH plasma concentrations in samples taken every 3 hours. In addition, in all goats, (n=20 by group), ovulation was determined by measuring plasma concentrations of progesterone. The proportion of female goats showing a preovulatory LH surge was higher in goats exposed to non-sedated (10/10) than in those exposed to sedated bucks (0/10; P<0.0001). Similarly, most of does in contact with non-sedated males ovulated (19/20), but none of those in contact with sedated males did so (0/20; P<0.0001). We conclude that the expression of an intense sexual behavior by male goats is necessary to induce LH preovulatory surge and ovulation in seasonally anovulatory goats.

Sexually active bucks are able to stimulate three successive groups of females per day with a 4-hour period of contact.

Bucks rendered sexually active by a photoperiod treatment of long days can induce fertile ovulation in a group of goats with only 4 h of contact daily with a male:female ratio of 1:10. Here we tested whether such bucks could induce fertile ovulations when stimulating successively three different groups of anovulatory goats when interacting 4 h per day during 15 consecutive days. Control males (n=3) were introduced in the control group (n=25) of does at 8:00 h and were removed at 12:00 h. Experimental males (n=3) were in contact with the experimental groups of does: from 8:00 h to 12:00 h with a first group (n=27), from 12:00 h to 16:00 h with a second group (n=26) and with a third one (n=27) from 16:00 h to 20:00 h. Bucks were then placed until next day in another pen. Both in the control and the experimental groups, more than 85% of females ovulated, and the proportions did not differ between the control and experimental groups (P≥0.67) or between the three experimental groups (P≥0.67). Moreover, the ovulation rate did not differ significantly between the control and the experimental females nor between the three experimental groups. Bucks were able to fertilize more than 72% of does independently of the number of females they were exposed to (P≥0.17). Finally, more than 58% of females kidded and fertility did not differ between the control and experimental groups (P=1) nor among experimental groups (P≥0.77). We conclude that sexually active bucks are able to induce fertile ovulation in three successive groups of anovulatory goats even when the period of contact between sexes is reduced to 4 h per day.