Exposure of rams in sexual rest to sexually activated males in spring increases plasma LH and testosterone concentrations.

Eight stimulating rams, and twelve stimulated rams, were used to determine whether a similar endocrine response to the introduction of sexually active males in spring in a flock of ewes is observed in a flock of rams. The stimulating rams (n = 4) were induced into a sexually active state by exposure to 2 months of long days (16 h light/d) (15 December-15 February). At the end of the long-day period, rams were returned to the natural photoperiod. Control-stimulating rams (n = 4) were kept under the natural photoperiod. On April 20, stimulated rams were divided into 2 groups, and joined with activated (ACT; n = 6) or control stimulating rams (C; n = 6). On the day of ram introduction, stimulated rams were blood sampled for 8 h at 20-min intervals, from 4 h before to 4 h after ram introduction, and next day from 24 to 28 h after ram introduction, and analyzed for plasma LH concentrations, and 10, 20 and 30 days after ram introduction to measure plasma testosterone levels. Mean (±SEM) plasma LH concentrations (ng/ml) of stimulated rams were similar during the 4 h before stimulating-ram introduction (ACT: 0.59 ± 0.03; C: 0.53 ± 0.04; P > 0.05). The introduction of the photoperiod-treated stimulating rams increased LH concentrations of stimulated rams during the 4 h after their introduction (1.14 ± 0.37) compared with the C group (0.51 ± 0.03; P < 0.05), especially during the first hour (ACT: 0.93 ± 0.16; C: 0.49 ± 0.03; P < 0.05), and during the blood sampling period 24-28 h after ram introduction (0.75 ± 0.07 vs. 0.58 ± 0.04; P < 0.05). Before the introduction of stimulating rams, the LH pulse frequencies and amplitudes did not differ between groups; however, LH pulsatility was higher at 4 h (0.58 ± 0.11 pulses/h; P < 0.05), and had trend to be higher 24 h (0.50 ± 0.06) (P = 0.10) after the introduction of the photoperiod-treated stimulating rams compared with the control-stimulating rams (0.29 ± 0.08 and 0.29 ± 0.10, respectively). As for LH pulses, there was an effect of group (P < 0.05) on LH amplitude, which presented a trend to be higher in ACT rams 4 h after ram introduction (1.68 ± 0.30; P < 0.10) and higher 24 h (1.07 ± 0.08; P < 0.05) after ram introduction, compared with LH amplitudes of C rams (0.71 ± 0.06 and 0.82 ± 0.07, respectively). Plasma testosterone concentrations of rams exposed to photoperiod-treated activated rams were higher than those of rams exposed to control-stimulating rams, at 4 h, 20 and 30 days after ram introduction (P < 0.05). In conclusion, sexually active rams in spring are able to stimulate LH and testosterone secretion of other rams in sexual rest, a phenomenon we called "ram-to-ram effect".

Milk let-down in goats can be influenced by the type of stimuli applied or to which goats are exposed during suckling and hand milking.

The present experiments on goats were conducted with the aim of verifying (a) whether the identity of the goat kids can affect the characteristics of milk let-down during suckling (Study 1) and (b) whether the presence of the goat kids may improve the milk let-down during hand milking (Study 2). In Study 1, 16 lactating goats with twins and 12 with single kids were used. Dams with single goat kids ejected more milk when suckling their kids at Days 5 and 20 of lactation, than when suckling aliens. Dams with twins ejected more milk when suckling their own kids than aliens only at Day 5 of lactation. Taking into account all the dams, they ejected more milk with more fat, protein, and lactose during suckling their own litter than when suckling aliens. In Study 2, when dams (n = 13) were hand milked in the presence of their own litter or in its absence, they ejected more milk with more fat, protein, and lactose than when milked in the presence of alien kids. We conclude that milk let-down in goats can be influenced by the type of stimuli applied or to which they are exposed to during suckling and milking.

Photoperiod-Treatment in Mediterranean Bucks Can Improve the Reproductive Performance of the Male Effect Depending on the Extent of Their Seasonality.

This study examines the effectiveness of the photoperiod treatment (extra light for 88 days) to render the bucks sexually active during the seasonal anoestrous in order to induce a male effect, depending on the reproductive seasonality of the breed of the bucks used. In April, 57 anoestrous Blanca Andaluza does were distributed into four groups with three males each: 13 were exposed to control Murciano-Granadina bucks (lower seasonality); 15 were exposed to photostimulated Murciano-Granadina bucks; 14 were exposed to control Blanca Andaluza bucks (higher seasonality), and 15 were exposed to photostimulated Blanca Andaluza bucks. After male introduction, the sexual behaviour of the bucks was assessed, and harness marks recorded doe oestrous behaviour. Ovulation was confirmed from plasma progesterone, and the ovulation rate was assessed by transrectal ultrasonography. Fecundity, fertility, prolificacy and productivity were also determined. All of the does in all of the groups showed ovulation. Interaction between both sources of variation was observed: the percentage of females showing oestrous (p < 0.01) and productivity (p < 0.05) was the lowest in the Blanca Andaluza control group (50% and 0.36 ± 0.17 goat kids born/female, respectively). In conclusion, photoperiod-treated bucks efficiently induce a male effect, but photostimulation could be more necessary for breeds with deep seasonality.

Sexually active bucks are a critical social cue that activates the gonadotrope axis and early puberty onset in does.

In rodents, early exposure to adult male is well known to induce an early puberty in females (Vandenbergh effect). This phenomenon has been less studied in other mammals. In goats, despite our extensive knowledge about the "male-effect" phenomenon in adults (i.e. ovulation induced by the introduction of the male during the anestrous), there are few data on the consequences of an early exposure of females to males. Here, we evaluated the puberty onset of young alpine goats when raised since weaning with intact bucks (INT), with castrated bucks (CAS) or isolated from bucks (ISOL). The INT group had the first ovulation 1.5 month before the two other groups. Despite the earlier puberty the INT group of females had normal and regular ovarian cycles. Morphological study of the genital tract showed that at 6 months, uterus of INT goats was 40% heavier than CAS and ISOL goats. Moreover, INT females had a myometrium significantly thicker and INT was the only group having corpora lutea. In our study, INT females were pubescent in the month following the entry of bucks into the breeding season, suggesting that only sexually active bucks provide the signal responsible for puberty acceleration. By removing direct contact with the bucks, we showed that somatosensory interactions were dispensable for an early puberty induction. Finally, no difference in the GnRH network (fiber density and number of synaptic appositions) can be detected between pubescent and non-pubescent females, suggesting that the male stimulations triggering puberty onset act probably on upstream neuronal networks, potentially on kisspeptin neurons.

Sexually active bucks are efficient to stimulate female ovulatory activity during the anestrous season also under temperate latitudes.

Goats are seasonal breeders and photoperiod is the main cue controlling the onset and offset of the breeding season. Nevertheless introducing a sexually active buck in a group of females during anestrous can stimulate their reproductive function and induce ovulation. This "male-effect" is very efficient under subtropical latitudes, when using sexually active males previously stimulated by a photoperiodic treatment. However, there is less evidence of its feasibility under temperate latitudes where the more important variation in day length could be responsible for a stronger inhibition of female sexual activity. The aim of this study was therefore to determine whether intense sexual activity can be induced in alpine bucks during the non-breeding season by a long-day treatment under temperate latitude and if these males could be used to produce an efficient male-effect. Bucks (n=21) were divided in two groups, one submitted to a photoperiodic treatment from November 1st to January 15th and then switched to natural photoperiod, while the other group remained entirely under the natural photoperiod. The ones submitted to this light treatment exhibit higher testicular volume and testosterone level 6 weeks after the end of the treatment. At the end of March, bucks were used to stimulate anestrous does (n=41) continuously for 15 days. We showed that (a) light treatment was efficient to induce an increase of sexual activity in bucks and (b) that the introduction of stimulated bucks among females induced a significantly higher proportion of ovulation in anestrous does than control bucks (86% vs 5%). Our results indicate that under temperate latitudes induction of ovulation in females during the anestrous season is feasible using bucks treated with long-days during winter.

Effects of exposure to artificial long days on milk yield, maternal insulin-like growth factor 1 levels and kid growth rate in subtropical goats.

This study was designed to determine whether any relationship exists between exposure to artificial long days, milk yield, maternal plasma insulin-like growth factor 1 (IGF-1) levels, and kid growth rate in goats. One group of lactating goats was maintained under naturally decreasing day length (control group; n = 19), while in another one, they were kept under artificial long days (LD group; n = 19). Milk yield was higher in goats from the LD group than that in the control group (P < 0.05). Maternal IGF-1 levels at day 57 of lactation were higher (P < 0.05) in goats from the LD group than the levels in the control group and were positively correlated with the total milk yields per goat at days 43 and 57 of lactation (r = 0.77 and r = 0.84, respectively; P < 0.01). Daily weight gain at week 4 was higher (P < 0.01) in kids from the LD group than that in kids from the control group and was correlated with total and average IGF-1 maternal levels (r = 0.60 and r = 0.60, P < 0.05). It was concluded that submitting lactating goats to artificial long days increases milk yield, plasma IGF-1 maternal levels and the growth rate of the kids.

One day of contact with photostimulated bucks is sufficient to induce ovulation in seasonally anestrous goats.

In seasonally anovulatory goats, daily contact with photostimulated bucks for 15 days stimulates ovulations in most females. In this study, we determined whether ovulation could be induced in goats exposed to photostimulated bucks for less than 15 days. Bucks were rendered sexually active during the nonbreeding season by exposure to 2.5 months of long days from November 1. The control group of females was exposed to one photostimulated buck for 15 days (n = 12). Other three experimental groups were exposed to males (n = 1 per group) for 1, 5, or 10 days (n = 14 or 15 females per group). Ovulations were determined by measurement of daily plasma progesterone concentrations during 17 days. All females from the control and experimental groups ovulated at least once during the experiment (P > 0.05). Furthermore, the proportions of goats that displayed a short luteal phase followed by a new ovulation, or a normal luteal phase after being in contact with males, did not differ between groups depending on the duration of time of contact with the photostimulated males (P > 0.05). In contrast, the proportions of females that displayed a short luteal phase followed by anovulation were greater in goats in contact with males for 1 day than in those in contact with males for 10 and 15 days (P < 0.05), whereas they did not differ from females exposed to males for 5 days (P > 0.05). We conclude that 1 day of contact with sexually active males is long enough to stimulate the ovulatory activity in seasonally anovulatory goats. However, a significantly higher proportion of females exposed to males for 1 day did not ovulate again after showing a short luteal phase.

Male goat vocalizations stimulate the estrous behavior and LH secretion in anestrous goats that have been previously exposed to bucks.

We investigated whether live vocalizations emitted by bucks interacting with anestrous females stimulate secretion of LH, estrous behavior and ovulation in anestrous goats. In experiment 1, bucks rendered sexually active by exposure to long days followed by natural photoperiod were exposed in a light-proof-building to five anestrous females. Buck vocalizations were reproduced through a microphone-amplifier-loudspeaker system to an open pen where one group of goats (n=6) was exposed for 10 days to these live vocalizations. Another group of females (n=6) was isolated from males and vocalizations. The proportion of goats displaying estrous behavior was significantly higher in females exposed to buck vocalizations than in females isolated from males. The proportion of goats that ovulated did not differ between the 2 groups (exposed to males versus isolated). In experiment 2, female goats that either had previous contact with males (n=7), or no previous contact with males (n=7) were exposed to live buck vocalizations, reproduced as described in experiment 1, for 5 days. The number and amplitude of LH pulses did not differ between groups before exposition to buck vocalizations. Five days of exposure to male vocalizations significantly increased LH pulsatility only in females that had previous contact with males, while LH pulse amplitude was not modified. We concluded that live buck vocalizations can stimulate estrous behavior and LH secretion in goats if they have had previous contact with bucks.

Lack of sexual experience does not reduce the responses of LH, estrus or fertility in anestrous goats exposed to sexually active males.

We investigated whether LH secretion, estrous behavior and fertility would differ between sexually inexperienced and experienced anestrous goats exposed to the males. Male goats were rendered sexually active during the reproductive rest season by exposure to 2.5 months of artificial long days. Two groups of anovulatory sexually inexperienced and sexually experienced does were exposed to males during 15 days (n = 20 per group). LH pulsatility was determined every 15 min from 4h before to 8h after introducing males (Day 0). Estrous behavior was recorded twice daily. Pregnancy rates were determined on Day 50. Fertility was determined at parturition. Male sexual behavior was registered on days 1 and 2 during 1h. Before introducing the males, the number of LH pulses did not differ between groups. After introduction of the males, all females increased their LH pulsatility, but the number of pulses did not differ between sexually inexperienced and experienced goats. The proportion of females displaying estrous behavior with a high pregnancy rate and fertility did not differ between inexperienced and experienced goats. The sexual behavior of the males did not differ significantly between those interacting with sexually inexperienced or experienced goats. We conclude that goats can show substantial endocrine and reproductive responses to males, even in the absence of previous sexual experience, when sexually active bucks are used.

Ovulatory activity of female goats adapted to the subtropics is responsive to photoperiod.

The objective of this study was to determine whether reproductive seasonality of local well-fed female goats from subtropical Mexico (26 degrees N) can be controlled by photoperiod. The control group (n=12) remained in an open pen under natural photoperiod variations. The two experimental groups (n=8 each) were placed in light-proof rooms and exposed for 2 yr to alternations of 3 months of long days and 3 months of short days. One group was first exposed to long days, Group 1, and the other one to short days, Group 2. Blood samples were obtained twice a week to determine ovulation status by progesterone plasma concentrations. Goats from the control group displayed a seasonality of ovulations. The mean (+/-SEM) dates of the onset and end of the ovulations were September 10+/-5 d and February 16+/-4 d, respectively. In contrast, in both experimental groups, ovulations were modified (P<0.05) by the photoperiodic treatments such that ovulations started and ended during short and long days, respectively. In Groups 1 and 2, ovulations started 67+/-2 and 66+/-2 d (P>0.05), respectively, after the transfer from long to short days. In contrast, the timing of the cessation of ovulations after the transfer from short to long days differed (P<0.05) between groups (19+/-3 and 31+/-3 d for Groups 1 and 2, respectively). Local female goats from subtropical Mexico are, therefore, sensitive to photoperiodic changes that they are exposed to at this latitude and this environmental cue may control the timing of the breeding season in natural conditions.

Male sexual behavior contributes to the maintenance of high LH pulsatility in anestrous female goats.

The objective of this study was to determine the importance of male sexual behavior in stimulating LH secretion in anovulatory female goats. Two groups of females (n=10 per group) were each exposed to a buck in sexual rest and submitted to natural daylength. In one group, the buck was awake, whereas in the other group, it was sedated to prevent its sexual behavior. Two other groups of goats (n=10 per group) were exposed to sexually active bucks that had been exposed to 2.5 months of long days. In one group, the buck was awake, and in the other group, it was sedated. LH secretion was determined every 15 min from 4 h before introducing the bucks to 8 h after, then every 15 min again from 20 to 24 h after introducing the bucks. The bucks submitted to natural daylength did not stimulate LH secretion (P>0.05), whether they were sedated or not. In contrast, both the awake and the sedated light-treated bucks induced an increase (P<0.05) of LH pulsatility in the first 4 h following their introduction. However, pulsatility remained elevated until 24 h in the females exposed to the light-treated awake buck, whereas in the group with the light-treated sedated buck, pulsatility diminished (P<0.05) after the first 4 h of stimulation by the buck. In conclusion, the sexual behavior of males contributes to the maintenance of a high LH pulsatility up to 24 h after introduction into a group of anovulatory goats.

Evidence that the photoperiod controls the annual changes in testosterone secretion, testicular and body weight in subtropical male goats.

The aim of this study was to determine whether the reproductive seasonality of local male goats from subtropical Mexico (26 degrees N) is controlled by photoperiod. The control group (n = 7) remained in an open shed under natural daylight. The two experimental groups (n = 6 each) were placed in light-proof buildings and exposed for 2 years (yr) to alternations of 3 months (mo) of long days and 3 mo of short days. One group was first exposed to long days and the other one to short days. Body and testicular weights were determined every 2 wk. Blood samples were obtained weekly to determine testosterone plasma concentrations. In the control group, the body weight exhibited variations (P < 0.0001) and it increased during the non-breeding season. In both treated groups, long days stimulated weight gain and short days inhibited it (P < 0.0001). In the control group, testicular weight displayed variations (P < 0.0001), and high values were registered in June. In the treated groups, a testicular weight reduction occurred 6-9 mo after the onset of the study. Afterwards, the changes in testicular size varied according to daylength (P < 0.01). The pattern of plasma testosterone concentration in the control group varied over the study (P < 0.0001) and the levels were higher from May-June to November. In both treated groups, the changes in testosterone secretion occurred according to photoperiod changes (P < 0.0001). Short days enhanced testosterone secretion one photoperiodic cycle after the onset of the study and long days inhibited it. Local male goats from subtropical Mexico are sensitive to photoperiodic changes and this environmental cue may control the timing of the breeding season in natural conditions.

Management of photoperiod to control caprine reproduction in the subtropics.

Reproductive seasonality is observed in some breeds originating from or adapted to subtropical latitudes. In 'photoperiodic flexible breeds', such as Australian cashmere goats, the annual breeding season can be manipulated through nutrition, whereas in 'photoperiodic rigid breeds', such as Creole goats from subtropical Mexico, sexual activity can be controlled by altering the photoperiod. In males from the latter breed, artificial long days, whether or not accompanied by the administration of melatonin, stimulate sexual activity during the non-breeding season. These treated males are able to induce the sexual activity of anoestrous females through the male effect under intensive or extensive conditions. Photoperiodic treatments and the male effect can be easily integrated into different breeding management systems in subtropical latitudes.

Maternal olfaction differentially modulates oxytocin and prolactin release during suckling in goats.

In postparturient goats, olfactory recognition of the young allows the establishment of a selective bond between the mother and her kids. Once this bond is formed, the mother rejects alien young that attempt to suckle. We tested whether the development of the maternal selective bond in goats modulates prolactin (PRL) and oxytocin (OT) release in response to suckling. On day 37 of lactation, serial blood samples were taken during nursing of the mother's own or alien kid(s) in 10 intact/selective goats and in 10 goats rendered anosmic/nonselective through prepartum peripheral ZnSO(4) irrigation. Spontaneous nursing behavior was also studied weekly from day 7 to 30 of lactation, at which time milk production was measured. Maternal selectivity had no effect on PRL release, in contrast to OT release, which was significantly affected by this factor. Intact mothers released OT only when nursing their own kids, but not with aliens, while anosmic/nonselective dams showed an increase in OT levels regardless of the identity of the kids. In addition to these effects on maternal selectivity, the amplitude of the response of both hormones was lower in anosmic mothers than in intact mothers. Finally, nursing behavior and milk production were not significantly affected by anosmia. We conclude that maternal selective behavior in goats, which relies on the individual olfactory signature of the kid, modulates the OT, but not the PRL, response to suckling. In addition, perception of the smell of the young appears to have a general facilitatory effect, independent of the kid's identity, on the release of both hormones.