Pregnancy and maternal behavior induce changes in glia, glutamate and its metabolism within the cingulate cortex

An upregulation of the astrocytic proteins GFAP and bFGF within area 2 of the cingulate cortex (Cg2) occurs within 3 hours of parturition in rats. These changes are the result of an interaction between hormonal state and maternal experience and are associated with increased dendritic spine density in this area. Here, we examined whether this upregulation of astrocytic proteins generalized to other glial markers and, in particular those associated with glutamate metabolism. We chose glial markers commonly used to reflect different aspects of glial function: vimentin, like GFAP, is a marker of intermediate filaments; glutamine synthetase (GS), and S-100beta, are used as markers for mature astrocytes and GS has also been used as a specific marker for glutamatergic enzymatic activity. In addition, we examined levels of proteins associated with glutamine synthetase, glutamate, glutamine and two excitatory amino acid transporters found in astrocytes, glt-1 and glast. S100beta immunoreactivity did not vary with reproductive state in either Cg2 or MPOA suggesting no change in the number of mature astrocytes across these conditions. Vimentin-ir did not differ across groups in Cg2, but expression of this protein decreased from Day 1 postpartum onwards in the MPOA. By contrast, GS-ir was increased within 24 h postpartum in Cg2 but not MPOA and similarly to GFAP and bFGF this upregulation of GS resulted from an interaction between hormonal state and maternal experience. Within Cg2, upregulation of GS was not accompanied by changes in the astrocytic glutamatergic transporters, glt-1 and glast, however, an increase in both glutamate and glutamine proteins were observed within the Cg2 of postpartum animals. Together, these changes suggest postpartum upregulation of glutamatergic activity and metabolism within Cg2 that is stimulated by pregnancy hormones and maternal experience.     

Changes in dendritic spine density on layer 2/3 pyramidal cells within the cingulate cortex of late pregnant and postpartum rats

A rapid upregulation of astrocytic protein expression within area 2 of the cingulate cortex (Cg2) of the maternal rat occurs within 3 h postpartum and persists throughout lactation. Previous studies have shown that similar changes in astrocytic proteins can signal changes in local synapses and dendritic spines. Thus, here we used the Golgi-Cox impregnation technique to compare spine density in layer 2 and 3 pyramidal cells of Cg2, the CA1 region of the hippocampus and the parietal cortex (ParCx) among metestrus, late pregnant (LP), 3-hour postpartum (3H PP) and 16-day postpartum rats (D16 PP). Rats in the 3H PP group had higher numbers of dendritic spines/10 μm on the apical dendrites of pyramidal neurons in both Cg2 and CA1 than the other groups, which did not differ. A similar pattern was observed in basilar dendrites but this failed to reach significance. In Cg2, Sholl analysis revealed that rats in the D16 PP group had a significantly greater extent of dendritic arborization in the basilar region than any other group. These data suggest that the changes in astrocytic proteins that occur in Cg2 in the postpartum period are associated with neuronal plasticity in pyramidal layers 2 and 3.     

哺乳、哺乳-禁乳及哺乳-禁乳后再哺乳对大鼠下丘脑胖素A免疫反应神经元的影响

为观察下丘脑胖素 A在哺乳期摄食增加和能量代谢中的作用,本研究采用脑连续切片之免疫组织化学和图像定量分析技术,对分娩后第 12 天非哺乳、持续哺乳、持续哺乳后禁哺乳过夜和持续哺乳 - 禁哺乳后再急性哺乳大鼠下丘脑胖素A免疫反应神经元的免疫反应性进行了观察和半定量分析。结果表明,分娩后持续哺乳 11 d, 大鼠的日摄食量较同期分娩的非哺乳大鼠明显增加(180%),一夜禁哺乳则明显降低哺乳大鼠的日摄食量(45%); 哺乳12 d, 大鼠下丘脑胖素 A免疫反应神经元的数目和平均染色强度较非哺乳大鼠明显增加(P<0.001,P<0.05); 禁哺乳过夜(15 h)明显降低哺乳大鼠胖素A免疫反应神经元的数目和平均染色强度(P<0.001,P<0.05),与非哺乳大鼠比较无明显差异;禁哺乳过夜后再急性哺乳2 h 明显增加禁哺乳大鼠胖素 A 免疫反应神经元的数目和平均染色强度(P<0.001,P<0.05),急性哺乳 5 h 后,虽亦明显增加禁哺乳大鼠胖素 A免疫反应性(P<0.05),但与急性哺乳 2 h 比较作用减弱。上述结果表明,持续哺乳和禁乳后再哺乳均导致下丘脑胖素A明显增加,提示哺乳期胖素A可能表达上调并可能与哺乳期摄食增加有关, 且吸乳动作与下丘脑胖素A样神经元之间可能存在某种神经或体液性联系途径。     

Postpartum aggression in mice: The influence of suckling stimulation

The influence of suckling stimulation upon postpartum aggression was studied by removing the nipples (thelectomy) of female mice at various times during pregnancy and lactation. Prepartum thelectomy, regardless of whether it was performed prior to mating or shortly before parturition, in combination with the fostering of young, prevented the exhibition of aggression. The aggressive behavior of females thelectomized following either 2 or 5 days of suckling experience was similar to that of normal lactating females. However, only 25% of animals thelectomized following 24 hr of suckling experience exhibited aggressive behavior. The results demonstrate that suckling stimulation is important for the initiation of postpartum aggression but is not essential in animals that have had at least 48 hr of suckling experience.     

Effects of the intensity of the suckling stimulus and ovarian steroids on pituitary gonadotropin-releasing hormone receptors during lactation

These studies examined whether the decrease in pituitary responsiveness to gonadotropin-releasing hormone (GnRH) observed during lactation in the rat results from a change in pituitary GnRH receptors. GnRH binding capacity was determined by saturation analysis using D-Ala6 as both ligand and tracer. During the estrous cycle, the number of GnRH binding sites increased from 199 +/- 38 fmol/mg protein on estrus to 527 +/- 31 fmol/mg protein on the morning of proestrus, whereas there was no change in receptor affinity (Ka, 6-10 X 10(9) M-1), During lactation, females nursing 8 pups on Days 5 or 10 postpartum had 50% fewer GnRH receptors (109-120 fmol/mg protein) than observed during estrus or diestrus 1 (199-242 fmol/mg protein) although receptor affinity was similar among all the groups. No deficits in pituitary GnRH receptors were observed in females nursing 2 pups on Day 10 postpartum. Removal of the 8-pup suckling stimulus for 24 or 48 h resulted in a dramatic increase in GnRH receptor capacity by 24 h from 120 +/- 16 to 355 +/- 39 fmol/mg protein. The rise in GnRH receptors after pup removal was accompanied by an increase in serum luteinizing hormone (LH) and estradiol concentrations. To assess the role of ovarian steroids in determining GnRH receptor capacity during lactation, females were ovariectomized (OVX) on Day 2 postpartum. Suckling of a large litter (8 pups) completely blocked the postcastration rise in serum LH and in pituitary GnRH receptors on Day 10 postpartum (OVX+ 8, 77 +/- 12 fmol/mg protein; OVX+ 0, 442 +/- 38 fmol/mg protein).(ABSTRACT TRUNCATED AT 250 WORDS)     

Suckling behaviour in the pipistrelle bat (Pipistrellus pipistrellus)

A sub-colony of 20 pipistrelle bats was studied during late pregnancy and lactation. Seventeen females gave birth to single young, with 16 young surviving to weaning. Free association and flight were permitted, and mother-infant pairs were identified and marked on the day of birth (day 0). Suckling associations were identified daily (n = 195) and females were never found suckling young other than their own. A series of choice experiments using young aged 0–10 days and 15–25 days indicated that females would permit only their own offspring to suckle during both early and late lactation. During early lactation, mothers actively selected their young and encouraged suckling but did not do so during late lactation. Mothers were most often indifferent to approaches by alien young but occasionally aggressively rejected them. Mothers never aggressively rejected their own young but frequently were indifferent to them. Young attempted to suckle from any mother without discrimination.     

Elevated Fos-like immunoreactivity in the brains of postpartum female prairie voles, Microtus ochrogaster

This study investigated the expression of the protein product of the immediate early gene c-fos in the brains of female prairie voles (Microtus ochrogaster) in association with pregnancy and postparturient activities including maternal behavior, lactation and postpartum estrus. Fos expression was assessed in female voles that were late in pregnancy, nonpregnant or at one of three different times postpartum (0-8, 12-24, and 24-48 h, respectively). A significant increase in the number of cells displaying Fos immunoreactivity (Fos-ir) was observed during the 0-8 h and 12-24 h postpartum time periods in the accessory olfactory bulbs, medial preoptic area, hypothalamus (specifically, the supraoptic nucleus, ventro-medial hypothalamus, and paraventricular nucleus), lateral septum, bed nucleus of the stria terminalis, and primary somatosensory area of the brain. The number of Fos-ir cells decreased after 24 h postpartum. There were no significant changes in Fos-ir cell numbers in the primary olfactory bulbs, hippocampus, or caudate putamen. The neural activation of the medial preoptic area, accessory olfactory bulbs, hypothalamus, and bed nucleus is consistent with reports in rats of Fos induction associated with the onset of maternal behavior. In voles postpartum estrous behavior begins and ends 0-12 h after parturition. Maternal behavior, including lactation, is initiated at the same time but persists for several weeks. The highest Fos-ir cell numbers reported here coincide with the timing of postpartum estrous behavior in this species.     

Effects of reduced suckling on days to estrus, conception during lactation and embryo survival in sows

Trials to investigate the effects of limited suckling on sow reproduction and piglet growth were conducted using 41 first parity and 32 second parity Yorkshire sows. Separation of sows from their litters (22 hrs/day, days 21-35 postpartum) induced estrus in 60% of primiparous and 72% of second parity sows during lactation. Compared to control group animals, primiparous sows had higher weaning weights and second parity sows higher rates of embryo survival on day 30 of gestation. Piglets subjected to suckling restriction had weights, at 2 weeks after weaning, equal to those weaned after 5 weeks of unrestricted suckling. We conclude that suckling restriction can provide the dual benefits of an extended piglet nursing period and a decreased breeding to breeding interval in sows.     

Loss of calcineurin from the medial preoptic area of primiparous rats

Western blot analyses reveal that calcineurin A (CNA), which is present in the hippocampus, basolateral amygdala, parietal cortex, and MPOA of virgin males and females, is undetectable only in the MPOA of primiparous females regardless of whether they had postpartum pup contact or not. In contrast, CNB was expressed at unchanging levels in the PC and MPOA. Similarly, G(alphao) and PKA(RI) were expressed at high levels in all of the brain regions of virgin males, virgin females, and primiparous females, supporting the concept that this loss of CNA is a specific event. Understanding how and why the expression of CNA, the sole neuronal Ca2+/CaM-dependent protein phosphatase, is down-regulated specifically in the MPOA of primiparous females may yield some insight into the signal transduction events that mediate the onset of mammalian maternal behavior.     

Effects of suckling mode on endocrine control of reproductive activity resumption in Texel ewes lambing in July or November

To study the effects of the overlapping of seasonal and lactation anestrus and the influence of the suckling mode on the resumption of reproductive activity in Texel ewes, two experiments were carried out after the July and November lambings. The frequency and the duration of suckling decreased with the age of the lambs, and the suckling intensity in the lambs that were allowed to suck unrestrictedly was three times higher than in lambs restricted to sucking three times a day. The overlapping of seasonal anestrus and lactation delayed the resumption of ovarian and estrus activity, but no difference was observed between dams suckling single lambs and twins. The resumption of ovarian and estrus activity was much earlier after the November lambing than after the July lambing. Weaning after the November lambing shortened the interval between parturition to first estrus but not to the first luteal function. The reduction of suckling intensity by suckling management had no effect on the resumption of ovarian and estrus activity. In early postpartum, suckling inhibited the luteinizing hormone (LH) pulsatile secretion and consequently the first LH discharge. However, the earlier restoration of gonadotropin stimulation in dry ewes was not immediately followed by ovarian activity. The suckling inhibition may be due to a temporary disturbance in hormonal balance, the rise in prolactin (PRL) and cortisol secretions. Plasma estradiol 17beta (E2) concentrations were higher in dry than in suckling ewes in early postpartum. Follicle stimulating hormone (FSH) secretion was not involved in the process of delaying the resumption of reproductive activity after lambing.     

Interaction of late pregnancy and lactation in rats.

The effect of pregnancy on lactation was studied during the third week of lactational pregnancy in postpartum pregnant rats with a delay in implantation of only 1 day (1d-LP rats). In an experimental design in which the suckling litter was prevented from consuming solid food, lactational performance was estimated by weighing the ten-pup suckling litters on days 16-21 of lactation or by measuring maternal weight loss after a nursing spell on day 21. In 1d-LP rats, food consumption as well as lactational performance was lower than it was in nonpregnant lactating rats (L rats) and pregnant-lactating rats with a normal long delay of implantation of at least 6 days (LP rats). The time spent by the pups sucking at the nipples was not different among the three groups, but the number of milk ejections was diminished in 1d-LP dams. Restriction of daily food supply during days 16 to 21 of lactation diminished lactational performance more strongly in 1d-LP rats than it did in L rats; 1d-LP rats conserved protein stores and mobilized fewer minerals than did L rats. The weight and composition of the litter in vitro were not affected by the food restriction. In pregnant-lactating rats (LP and 1d-LP rats), the number of early resorptions was increased in comparison with pregnant rats, showing that lactation can affect the earlier stages of pregnancy. It was concluded that late pregnancy does not affect nursing behaviour, but suppresses lactation by restricting maternal food intake and mobilization of maternal stores. Measurements in serum indicate a causative role for oestradiol, but not for leptin.     

Increased cholesterol 7alpha-hydroxylase expression and size of the bile acid pool in the lactating rat

Maximal bile acid secretory rates and expression of bile acid transporters in liver and ileum are increased in lactation, possibly to facilitate increased enterohepatic recirculation of bile acids. We determined changes in the size and composition of the bile acid pool and key enzymes of the bile acid synthetic pathway [cholesterol 7alpha-hydroxylase (Cyp7a1), sterol 27-hydroxylase (Cyp27a1), and sterol 12alpha-hydroxylase (Cyp8b1)] in lactating rats relative to female virgin controls. The bile acid pool increased 1.9 to 2.5-fold [postpartum (PP) days 10, 14, and 19-23], compared with controls. A 1.5-fold increase in cholic acids and a 14 to 20% decrease in muricholic acids in lactation significantly increased the hydrophobicity index. In contrast, the hepatic concentration of bile acids and small heterodimer partner mRNA were unchanged in lactation. A 2.8-fold increase in Cyp7a1 mRNA expression at 16 h (10 h of light) demonstrated a shift in the diurnal rhythm at day 10 PP; Cyp7a1 protein expression and cholesterol 7alpha-hydroxylase activity were significantly increased at this time and remained elevated at day 14 PP but decreased to control levels by day 21 PP. There was an overall decrease in Cyp27a1 mRNA expression and a 20% decrease in Cyp27a1 protein expression, but there was no change in Cyp8b1 mRNA or protein expression at day 10 PP. The increase in Cyp7a1 expression PP provides a mechanism for the increase in the bile acid pool.     

Studies of pericopulatory pregnancy blockage and the gestation period in deer mice (Peromyscus maniculatus)

It has previously been shown that female deer mice in cycling estrus that copulate with or are exposed to more than one male at the time of mating have a lower probability of delivering a litter than when mating with and being exposed to just one male. In the present research the effect of post-mating exposure to the bedding of a strange versus a familiar male was studied in females in postpartum estrus. In four experiments females in postpartum estrus were protected from pericopulatory pregnancy blockage, presumably by suckling-induced elevated prolactin levels. Varying female experience, number of ejaculations, and the presence of suckling pups had no effect on this protection. Thus, both the pericopulatory block and the Bruce effect (a more delayed type of pregnancy blockage) occur in cycling, but not suckling, females. This suggests that the two may have a common mechanism and leads to a new perspective in the search for adaptive significance. In addition, the subsequent gestation periods were prolonged in females nursing young; the length was affected by the number of pups weaned from the previous litter and the number of pups being carried.     

Role of pup age, estradiol-17 beta and pituitary responsiveness in the differences in the suckling-induced prolactin response during early and late lactation

The suckling-induced prolactin (Prl) response was studied in 10- and 20-day postpartum female rats. The response in 20-day postpartum mothers has a slower onset, has markedly reduced peak values and returns to baseline somewhat sooner than the response in 10-day postpartum mothers. The blunted response of late lactation was seen in mothers suckled for 30 min and was maintained over a longer interval in mothers continuously suckled for 120 min. This refractory phenomenon was not due to decreased suckling intensity provided by 20-day relative to 10-day-old pups. Pituitary gland Prl release in response to thyrotropin releasing hormone (TRH) and haloperidol challenges also did not distinguish 10- from 20-day postpartum mothers. Significantly higher estradiol-17 beta levels were found in 20-day compared with 10-day postpartum mothers, a finding which cannot account for the blunted response. Pup separation from 10-day postpartum mothers for 4, 24, 48 or 72 h did not produce a blunted response like that seen in late lactation. It is suggested that the hypothalamic mechanism mediating suckling-induced Prl release becomes refractory to the suckling stimulus during the preweaning period.     

NDRG2 as a marker protein for brain astrocytes

The protein NDRG2 (N-myc downregulated gene 2) is expressed in astrocytes. We show here that NDRG2 is located in the cytosol of protoplasmic and fibrous astrocytes throughout the mammalian brain, including Bergmann glia as observed in mouse, rat, tree shrew, marmoset and human. NDRG2 immunoreactivity is detectable in the astrocytic cell bodies and excrescencies including fine distal processes. Glutamatergic and GABAergic nerve terminals are associated with NDRG2 immunopositive astrocytic processes. Müller glia in the retina displays no NDRG2 immunoreactivity. NDRG2 positive astrocytes are more abundant and more evenly distributed in the brain than GFAP (glial fibrillary acidic protein) immunoreactive cells. Some regions with very little GFAP such as the caudate nucleus show pronounced NDRG2 immunoreactivity. In white matter areas, NDRG2 is less strong than GFAP labeling. Most NDRG2 positive somata are immunoreactive for S100ß but not all S100ß cells express NDRG2. NDRG2 positive astrocytes do not express nestin and NG2 (chondroitin sulfate proteoglycan 4). The localization of NDRG2 overlaps only partially with that of aquaporin 4, the membrane-bound water channel that is concentrated in the astrocytic endfeet. Reactive astrocytes at a cortical lesion display very little NDRG2, which indicates that expression of the protein is reduced in reactive astrocytes. In conclusion, our data show that NDRG2 is a specific marker for a large population of mature, non-reactive brain astrocytes. Visualization of NDRG2 immunoreactive structures may serve as a reliable tool for quantitative studies on numbers of astrocytes in distinct brain regions and for high-resolution microscopy studies on distal astrocytic processes.     

Effect of suckling on the hypothalamic-pituitary axis in postpartum beef cows, independent of ovarian secretions

Thirty-two postpartum (PP) cows were used to investigate the effect of suckling on secretion of luteinizing hormone (LH). Calves remained with their dams (suckled; S), or they were removed within 24 h of birth (nonsuckled; NS). To evaluate the relationship between suckling and negative feedback regulation of LH, cows were ovariectomized on Day 5 PP, then injected intravenously with estradiol-17 beta (E) or vehicle (V) on Day 10 PP. To investigate the influence of suckling on the gonadotropin-releasing hormone (GnRH)-induced release of LH, cows were injected with 80 micrograms of GnRH on a single day varying from 18 to 85 days PP. Suckling inhibited the postcastration rise in LH, as LH concentrations increased at a faster rate in NS compared with S cows [0.031 +/- 0.02 ng/(ml X day) LH: P less than 0.05]; this was not influenced by basal amounts of E since amounts did not differ between S and NS cows at ovariectomy (5.37 +/- 0.36 vs. 5.34 +/- 0.48 pg/ml E; P greater than 0.05). Serum concentrations of LH were negatively related to total follicular E only in S cows (r = -0.71; P less than 0.01). Estradiol-17 beta caused a decrease not only in the level but also the variability in LH concentrations in both S and NS cows: LH in S cows was less variable after E than in NS cows (P less than 0.001), but the magnitude of LH suppression was not influenced by suckling (P greater than 0.25). The regression of LH response on days PP was essentially the same over time for both S (P greater than 0.25) and NS (P greater than 0.25) cows, indicating that LH response to a GnRH injection was not influenced by suckling or days PP. Suckled cows had a tendency to release more LH relative to their baseline in response to GnRH as time PP increased (P less than 0.10), but NS cows did not. These results indicate that even though ovarian secretions inhibit LH release from the pituitary, other inhibitory influences may have a major effect in S cows. Concentrations of LH were lower in S cows than NS cows on Day 10 PP, following removal of the ovaries on Day 5, suggesting that suckling had a direct effect on the hypothalamic-pituitary axis.     

Basic fibroblast growth factor, neurofilament, and glial fibrillary acidic protein immunoreactivities in the myenteric plexus of the rat esophagus and colon

The enteric nervous system consists of a number of interconnected networks of neuronal cell bodies and fibers as well as satellite cells, the enteric glia. Basic fibroblast growth factor (bFGF) is a mitogen for a variety of mesodermal and neuroectodermal-derived cells and its presence has been described in many tissues. The present work employs immunohistochemistry to analyze neurons and glial cells in the esophageal and colic enteric plexus of the Wistar rat for neurofilament (NF) and glial fibrillary acidic proteins (GFAP) immunoreactivity as well as bFGF immunoreactivity in these cells. Rats were processed for immunohistochemistry; the distal esophagus and colon were opened and their myenteric plexuses were processed as whole-mount preparations. The membranes were immunostained for visualization of NF, GFAP, and bFGF. NF immunoreactivity was seen in neuronal cell bodies of esophageal and colic enteric ganglia. GFAP-immunoreactive enteric glial cells and processes were present in the esophageal and colic enteric plexuses surrounding neuronal cell bodies and axons. A dense net of GFAP-immunoreactive processes was seen in the ganglia and connecting strands of the myenteric plexus. bFGF immunoreactivity was observed in the cytoplasm of the majority of the neurons in the enteric ganglia of esophagus and colon. The two-color immunoperoxidase and immunofluorescence methods revealed bFGF immunoreactivity also in the nucleus of GFAP-positive enteric glial cells. The results suggest that immunohistochemical localization of NF and GFAP may be an important tool in the study of the plasticity in the enteric nervous system. The presence of bFGF in neurons and glia of the myenteric plexus of the esophagus and the colon indicates that this neurotrophic factor may exert autocrine and paracrine actions in the enteric nervous system.     

Emergence and selection of the dominant follicle and gonadotropin dynamics in postpartum lactating versus non-postpartum cycling mares

Among female livestock, the mare has the shortest interval from parturition to first ovulation. Due to the scarcity of research on postpartum mares, little progress has been made on the characterization of the resumption of ovarian cyclicity after parturition. This study compared follicular and gonadotropin dynamics during follicle emergence and deviation in postpartum lactating (PP Lactating) versus non-postpartum cycling (N-PP Cycling) mares. On the day of parturition, every PP Lactating mare was paired with a N-PP Cycling mare. Comparisons were made by considering the partum-ovulation interval and the postpartum interovulatory interval for the PP Lactating mares, and two interovulatory intervals for the N-PP Cycling mares. The results presented herein demonstrate that during the postpartum period, lactating mares have some similarities in follicular and hormonal profiles around emergence and deviation when compared with non-postpartum cycling mares. However, some peculiar and important characteristics were noticed during the postpartum period in lactating mares: (1) The emergence of the DF occurs around the day of parturition; (2) follicle deviation in the ovulatory wave occurs earlier during the foal heat than in other intervals; (3) lower FSH and LH systemic concentrations were not detrimental enough to prevent the rapid resumption of ovarian activity just after parturition; and (4) the association between parturition and season can have an additional and confounding effect during postpartum ovarian activity in mares. The novel findings of this study provide better knowledge of the resumption of ovarian activity after parturition and may help provide insight into the reproductive management of this species.     

Reduced anxiety in postpartum rats requires recent physical interactions with pups, but is independent of suckling and peripheral sources of hormones

Changes in emotional behavior occur across the reproductive cycle in female rodents, with reduced anxiety found during the postpartum period, but relatively little is known about factors contributing to this decreased anxiety. Using increased duration of time spent in the open arms of an elevated plus-maze as an indicator of reduced anxiety, it was found in a series of experiments that (1) anxiety is significantly reduced in Long-Evans females during the first week of lactation, but not thereafter, (2) relatively recent contact with pups before testing (within 4 h) is necessary for their reduced anxiety, (3) dams that receive only distal sensory cues from pups for the 4 h prior to testing do not show reduced anxiety, (4) the absence of nipples, and therefore a lack of suckling by pups, has no effect on dams' anxiety, (5) cesarean delivery of pups 2 days prior to expected parturition did not alter later anxiety in dams, (6) hypophysectomy during mid-pregnancy or ovariectomy within 24 h after parturition also did not prevent reduced anxiety in dams, and (7) differences in anxiety between lactating and virgin females are greatest 4-8 min after being placed in the plus-maze. Therefore, exposure to their own peripheral hormones through mid-pregnancy is sufficient to prime female rats to show reduced anxiety, but only if they later have recent physical interaction with pups. Furthermore, because suckling and the peripheral hormones released during suckling appear to be unnecessary, decreased anxiety in maternal rats may instead be regulated by the transient intracerebral release of neuropeptides or neurotransmitters while dams receive other types of tactile inputs from their infants.     

Suckling effects in sows: importance for mammary development and productivity

An understanding of the mechanisms regulating milk yield in sows is crucial for producers to make the best management decisions during lactation. Suckling of mammary glands by piglets is one factor that is essential for development of these glands during lactation and for the maintenance of lactation in sows. The process of mammary development is not static as the majority of it takes place in the last third of gestation, continues during lactation, is followed by involution at weaning and starts over again in the next gestation. During involution, the mammary glands undergo a rapid and drastic regression in parenchymal tissue, and this can also occur during lactation if a gland is not suckled regularly. Indeed, the pattern of regression is similar for glands that involute at weaning or during lactation. Suckling during 12 to 14 h postpartum is insufficient to maintain lactation and the process of involution that occurs in early lactation is reversible within 1 day of farrowing but is irreversible if a gland is not used for 3 days. However, milk yield from a gland which is ‘rescued’ within the first 24 h remains lower throughout lactation. Suckling does not only affect milk yield in the ongoing lactation, but it also seems to affect that of the next lactation. Indeed, non-suckling of a mammary gland in first-parity sows decreased development and milk yield of that gland in second parity. Nursing behaviour of piglets in early lactation was also affected, where changes were indicative of piglets in second parity being hungrier when suckling glands that were not previously used. It is not known, however, if the same effects would be seen between the second and third lactation. Furthermore, the minimum suckling period required to ensure maximal milk yield from a gland in the next lactation is not known. This review provides an update on our current knowledge of the importance of suckling for mammary development and milk yield in swine.