H. Hernández

Sensory and physiological determinants of maternal behavior in the goat (Capra hircus)

Maternal behavior in the goat appears at the time of parturition, partly under the activating influence of vaginocervical stimulation. Mothers actively lick their neonate and rapidly establish a selective bond with their kid through olfactory recognition. They also develop visual and acoustic recognition of the kid within 4 h following birth. Acoustic recognition is present at 48 h. The establishment of maternal recognition can be impaired by underfeeding during the second half of pregnancy. There is no indication that the mechanisms controlling the onset of maternal behavior and bonding are different from those reported in sheep, despite the fact that lambs start to follow their mother within a few hours after birth and kids hide for about a week. During lactation, the cues provided by the kid are necessary for the maintenance of maternal responsiveness, but suckling itself does not appear of primary importance. The presence of the kid also modulates the hormonal response to udder stimulation and influences recovery of postpartum sexual activity when kidding (i.e. birthing) takes place in autumn. Finally, the rapid establishment of mutual attachment between mother goats (does) and their kids offers the possibility to investigate an aspect of mother-young affiliation that is not present in many laboratory species.

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.

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.

Differential effects of undernutrition during pregnancy on the behaviour of does and their kids at parturition and on the establishment of mutual recognition

We investigated whether undernutrition during the second half of pregnancy impaired the behaviour of does and their kids at parturition and early mutual recognition. Twenty-two control and 22 underfed mixed-breed, multiparous dairy goats were used, together with their respective kids (control, n = 31: nine singles, 16 twins and six triplets; underfed, n = 32: 11 singles, 18 twins and three triplets). Undernutrition involved limiting protein and energy intake at 70% of the nutritional requirements for maintenance and foetal growth from day 70 of pregnancy until birth. The behaviour of mothers and their two first-born kids was observed for 90 min from the birth of each kid. Maternal olfactory recognition of the kid was assessed at 4 h post partum by testing selective nursing behaviour. Non-olfactory recognition was assessed at 8 h in a two-choice test excluding olfactory cues. In kids, preference for the mother was assessed in a two-choice test at either 12 or 24 h post partum. Bodyweight of does and kids were lower in the underfed group up to 2 weeks post partum. At parturition, licking, maternal bleating frequency and latency to nursing did not differ between nutritional groups. Control kids were faster than underfed kids to stand, search for and reach the udder, but underfed kids bleated more and tended to spend more time at the udder. Both control and underfed does accepted their own kid and rejected the alien in the selectivity test at 4 h. In contrast, at 8 h post partum, only control goats showed a significant preference for their own kid in the non-olfactory recognition test. Both control and underfed kids showed a preference for their own mother at 12 and 24 h and undernutrition during pregnancy had little influence on the performance of kids. However, 12 h-old underfed kids tended to be less active than control kids and visited their own mothers less than control kids. There were no significant correlations between the behaviour of the mother or of the kid at parturition and their performance in the discrimination tests. Overall, undernutrition in the second half of pregnancy appears to be more detrimental for the behaviour of the mother than for the kid. Furthermore, it has more impact on the establishment of maternal non-olfactory recognition than on maternal care at parturition or the establishment of maternal selectivity.

The ovulatory response of anoestrous goats exposed to the male effect in the subtropics is unrelated to their follicular diameter at male exposure.

This study was designed to determine whether the follicular diameter at the introduction of the bucks influences the ovarian response in does exposed to males during the anoestrous season in the subtropics. Bucks (n = 4) were subjected to 2.5 months of long days from November 1st to stimulate their sexual activity. On 29th March, one of the four treated males was joined with the females (n = 13), being exchanged with other males every 12 h, during 10 days. Oestrous behaviour was recorded twice daily. Ultrasound examinations of the ovaries were performed once daily from Day -7 to -1 and twice daily from Day 0 to 6. Follicles that ovulate were categorized according to the diameter at the moment when females were joined with males, as Small (<3.9 mm), Medium (4.0-5.9 mm) or Large (>6.0 mm). All females ovulated (13/13) and 12 came into oestrus during the first 5 days after exposure to males. The growth rate of the follicles increased after the introduction of the bucks from 1.1 ± 0.1 mm per day to 1.5 ± 0.1 mm per day (p < 0.05). The percentage of follicles from each category that ovulated did not differ (p > 0.05; Small 47.8%; Medium 34.8% and Large 17.4%). From follicles that ovulated, the growth rate of those that were Small at the moment of the introduction of the bucks was greater (2.1 ± 0.1 mm per day; p < 0.05) than that observed in those that were Medium (1.3 ± 0.1 mm per day) and Large follicles (1.1 ± 0.1 mm per day). In 12 does, the largest follicle present in the ovaries was growing when bucks were introduced. From these follicles, five finally ovulated and seven finally regressed. In conclusion, the follicular diameter at the introduction of the bucks is not related to the oestrous behaviour and ovulatory responding patterns in female goats exposed to sexually active bucks in the subtropics.

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.

Artificial long-day photoperiod in the subtropics increases milk production in goats giving birth in late autumn

Two experiments were conducted to determine whether exposure to a photoperiod of artificial long days in autumn increased milk yield in subtropical goats milked once (Exp. I) or twice daily (Exp. II). In Exp. I, starting at d 10 of lactation, 1 group of does was kept under naturally decreasing photoperiod (DD1X; n = 8), whereas the other group was submitted to an artificial photoperiod of long days (LD1X; n = 8; 16 h light:8 h darkness). The kids were weaned 28 d after parturition, and dams were manually milked once daily. Milk yield and milk components (fat, protein, and lactose) were assessed up to 140 d of lactation. From d 0 to 28 of lactation (suckling phase), mean daily milk yield did not differ between DD1X and LD1X goats (2.3 ± 0.2 kg vs. 2.4 ± 0.2 kg; P = 0.717). However, between d 29 and 84 (early milking phase), mean daily milk yield was greater in LD1X does than in DD1X does (2.6 ± 0.1 kg vs. 2.1 ± 0.1 kg; P = 0.001). Finally, between d 85 and 140 (late milking phase), mean daily milk yield was greater in LD1X goats than in DD1X goats (P ≤ 0.05) only during the first 2 wk. In Exp. II, one group of goats was exposed to a photoperiod of naturally decreasing days (DD2X; n = 8) and another group was submitted to an artificial photoperiod of long days (LD2X; n = 7). In both groups, kids were weaned on d 28 of lactation and the dams were manually milked twice daily. During the nursing phase, mean daily milk yield did not differ between the DD2X and LD2X groups (2.5 ± 0.3 kg vs. 2.6 ± 0.2 kg; P = 0.767). In the early milking phase, mean daily milk yield was greater in LD2X than in DD2X goats (3.3 ± 0.2 kg vs. 2.8 ± 0.2 kg; P = 0.022), whereas during the late milking phase, milk yield did not differ between the 2 groups (P = 0.946). In both experiments, milk composition was not significantly influenced by exposure to long-day photoperiod. We conclude that, in subtropical female goats that start lactation in late autumn, exposure to an artificial long-day photoperiod stimulates milk yield, even if goats are milked once daily. In addition, combining exposure to long days with twice-daily milking will increase further milk yield in such goats without affecting milk components.

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.

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.

Feeding corn during the last 12 days of gestation improved colostrum production and neonatal activity in goats grazing subtropical semi-arid rangeland.

The objectives were to investigate if partial substitution of a grazing diet with a supplemental high-starch feed during the last 12 d of gestation improved colostrum yield and neonatal activity in goat does grazing semi-arid rangeland. For the first objective, 25 pregnant does were randomly assigned to 1 of 2 treatments: 1) grazing only (control; n = 11), and 2) grazing plus 0.6 kg/d of supplemental corn (as fed), a high starch feed, during the last 12 ± 1.0 d before parturition (SC; n = 14). Colostrum was collected at parturition and again at 1, 3, 6, and 10 h postpartum for yield and composition. In all goats, blood concentrations of glucose and plasma progesterone were determined. Total colostrum yield/10 h was greater (P = 0.002) in the SC does (1,102 ± 144 g) than in control does (405 ± 50 g). From 6 to 10 h postpartum, contents of colostrum protein and solids not fat were less (P ≤ 0.012) whereas lactose contents were greater (P = 0.035) in the SC does than in control does. Concentration of glucose in the blood at parturition was greater (P = 0.037) in the SC does (160 ± 13 mg/dL) than in control does (115 ± 12 mg/dL). A significant decrease (P = 0.001) in plasma progesterone concentrations occurred 1 d before parturition in control does, whereas the decrease (P = 0.008) occurred 3 d before parturition in SC does. For the second objective, 20 does (10 per group) and their single kids were randomly assigned to the same treatments as above for evaluation of dietary treatment on neonatal activity. Kid activity was assessed using videos recorded during the first 90 min after birth. Frequency of low-pitched bleats was greater (P < 0.001) in kids of SC does than in kids of controls. Frequencies and durations of teat seeking from 30 to 90 min after birth and of suckling activity were greater (P ≤ 0.015) in kids of SC does than in kids of controls. Duration of parturition was longer (P = 0.001) in control does (58 ± 10.2 min) than in SC does (21 ± 2.8 min). Finally, percentage of fetal malpresentation was greater (P = 0.036) in control does (33.3%) than in SC does (9.0%). We conclude that partial substitution of the grazing diet with a high-starch feed, corn, during the last 12 d of gestation improved colostrum yield and neonatal activity in goats grazing subtropical semi-arid rangeland.