Change of nipple preference between successive offspring in Japanese macaques

Selective use of nipples among free-ranging infant Japanese macaques (Macaca fuscata) was studied. Each infant held only one of its mother's two nipples in the mouth during an average of 90.9% ± 1.5% (SEM) of total nipple contact time. The next offspring born to the same mother tended to use the nipple that its previous sibling had not used. Maternal behavior did not cause use of one nipple or the other.     

Fighting by Kittens and Piglets during Suckling: What Does it Mean?

Newborn kittens and piglets show some notable similarities in suckling‐associated behaviours. Both develop a pattern of teat use in which each member of the litter predominantly occupies one or two particular nipples or teats, both show agonistic behaviour towards littermates during suckling, and both are born with ‘weapons’–kittens with sharp claws and piglets with sharp needle teeth. Traditionally, the differential use of nipples or teats has been thought to be due to differential quality of the mammary glands. However, neither our findings in kittens nor an examination of the literature on piglets support this view. As mammary glands usually are of equal quality and as there are normally more glands than young, there seems little reason for littermates to fight. Furthermore, there is also little evidence that individual performance in suckling contests or use of ‘weapons’ is related to weight gain or to dominance relations. Given the present lack of satisfying adaptive explanations for the seemingly aggressive behaviour and use of weapons by kittens and piglets, we suggest these to be epiphenomena of developmental processes anticipating the use of these behaviours and structures in later adaptive contexts.     

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.     

Suckling Behavior of the Collared Peccary (Tayassu tajacu)

Based on Trivers' (1974) theory of parent/offspring conflict, we predicted that weaning would be accelerated under poor dietary conditions in an aseasonally breeding, monomorphic ungulate, the collared peccary (Tayassu tajacu). Suckling behavior was observed in 18 captive litters under a dietary treatment of high or low protein formulated to mimic forage under ecological conditions of normal rainfall vs. drought. Litters on the low protein diet weaned later (mean of 27 weeks) than litters on the high protein diet (mean of 21 weeks). Other indices of cumulative suckling effort (weaning weight, cumulative minutes suckling, cumulative bouts) did not differ between dietary treatments. No differences in suckling effort were found relative to offspring sex or litter size. Milk did not completely compensate for the low protein diet affecting growth of young in the second and third phases of lactation. Some nursers did not conserve bodily resources as would be expected if the benefits of short interbirth intervals outweighed the costs of lactation during unpredictable drought conditions.     

Differences in fertility and suckling patterns between primiparous and multiparous rhesus mothers (Macaca mulatta)

Primiparous females gave birth around the same time as multiparous nonlactating females, and earlier than did multiparous lactating females. No differences in birth sex ratio were found between primiparous and multiparous females. During the breeding season following birth, primiparous mothers returned to oestrus later than did multiparous mothers, and while few primiparous mothers conceived successfully during that season, most multiparous mothers did. Primiparous females suckled their infants more frequently than did multiparous females at all ages; infants of primiparous females also made more nipple contacts per bout, and had shorter sucking bouts. When mothers came into oestrus, suckling frequency drastically increased for primiparous females, but not for multiparous females, magnifying the differences between the two groups. After the first oestrus, suckling frequency declined for all mothers, but multiparous mothers had consistently lower suckling frequencies than did primiparous mothers. The high suckling frequency, and numerous nipple contacts per bout, found among primiparous mothers are likely to be related to the low reproductive chances that these females faced during the breeding season. Multiparous mothers seemed to compensate for their low suckling frequency by lengthening the suckling bouts, and this suckling pattern did not hinder their reproduction. It is argued that primiparous mothers might have to suckle their infants more frequently because they can only produce milk at slow rates, being in this way forced into a reproductively inhibiting suckling pattern. However, the delay in subsequent reproduction could be ultimately advantageous for primiparous mothers if it enhanced infant survival, and allowed the mothers to regain physical condition before reproducing again.     

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.     

Tenacious nipple attachment in rodents: the sibling competition hypothesis

Tenacious attachment to the mother's nipples by suckling pups has been observed in many rodent species, but apeears to be absent in others. The natural history of this behaviour is reviewed, along with the possible adaptive value. Four hypotheses are extracted from the literature: the antipredator, anti-falling, anti-drowning, and infant-nipple contact hypotheses. These hypotheses are found to be ad hoc and supported by only anecdotal evidence. An alternative hypothesis is presented, that tenacious nipple attachment is an adaptation to sibling competition for maternal resources (nipples and milk). Morphological specialization, developmental time course, and patterns of pup mortality support the new model. The behaviour has been noted only in the Muridae. Quantitative interspecific comparisons within the Muridae demonstrate that tenacious nipple attachment is associated with reproductive tactics that promote sibling competition: breeding by means of litter overlap, low mammary number, and more offspring per mammary.     

Calcitonin gene-related peptide (CGRP) in the nipple of the rat mammary gland

The distribution of nerve fibres immunoreactive to calcitonin gene-related peptide (CGRP) was investigated by immunohistochemistry in nipples and mammary glands from lactating and non-lactating rats and compared to the immunoreactivity of other neuropeptides including substance P (SP), neuropeptide Y (NPY), vasoactive intestinal peptide (VIP) and somatostatin (SOM). The study revealed an extensive innervation of the mammary nipples, in which CGRP-immunoreactive (IR) nerve fibres were abundantly present in the epidermis, dermal connective tissue and intralobular connective tissue of the mammary gland parenchyma. Several of the dermal CGRP-IR fibres seemed to follow blood vessels, or formed ringlet-like structures. The latter were mostly observed in the dermal connective tissue of the nipple from the lactating rat and may have a mechanoreceptive function, e.g. for the suckling stimuli. The location of SP-IR appeared to be comparable to CGRP-IR, but in fewer fibres. Dense NPY-IR networks of nerve fibres were closely associated with the fascicles of smooth musculature in the core of the nipple base. In contrast, VIP-IR fibres were only sparsely present, and SOM-IR was not detected in the mammary nipples. The immunoreactive content of CGRP and SP was determined by radioimmunoassays. The total amount of immunoreactive CGRP was significantly higher in the nipples from the pregnant and the lactating rats when compared to SP. The maximum concentration of CGRP (65.9±4.0 pmol/g) measured in the nipples of the pregnant (day 10) rats exceeded almost ninefold the maximum concentration of SP (7.7±2.0 pmol/g). Thus, the immunoreactive content of CGRP in the nipples confirmed the immunohistochemical observations, and the present results strongly suggest that CGRP is a major neuropeptide in the rat nipple.     

A note on the effects of co-mingling piglet litters on pre-weaning growth, injuries and responses to behavioural tests

The purpose of this study was to determine how co-mingling litters affected piglets’ pre-weaning growth, ear injuries, suckling behaviour and responses to behavioural tests used to measure coping abilities. Thirty sows and their respective litters were housed in standard farrowing crates until day 13 after birth. On day13, the partition between two neighbouring pens was removed for 20 litters allowing piglets to interact (forming 10 co-mingled litters). The remaining 10 control litters were kept in standard farrowing crates throughout the experiment. Three focal piglets from each litter were used for data collection. Focal piglets were weighed and ear injuries recorded on days 2, 4, 9, 12, 15 and 18 after birth. There were no differences in piglets’ weight gain before or after co-mingling. Ear injuries were more abundant in co-mingled litters on day 15 (P < 0.05) but these differences disappeared by day 18. Suckling behaviour was recorded on days 5, 8, 10, 14, 16 and 18 after birth. There were no differences in teat fidelity, suckling frequency and mother fidelity between treatments. Three behavioural tests, social challenge, isolation, and backtest, were performed before and after co-mingling. There were no treatment effects on piglets’ response to the isolation test and backtest. Co-mingled piglets showed longer latency for the first aggressive interaction (P < 0.05), spent more time in proximity to one another (P < 0.05) and performed less single bites (P < 0.05) than control piglets during the social challenge. In addition, the duration and frequency of aggressive interactions (P < 0.05) were lower in co-mingled piglets than control piglets. Co-mingling did not affect the frequency of single head thrusts or oral–nasal contact, but did tend to increase the frequency of escape attempts (P < 0.10). Our results suggest that co-mingling litters during lactation affects piglets’ social behaviour, by primarily decreasing aggressive interactions during social challenges.     

Light on the moth-eye corneal nipple array of butterflies

The outer surface of the facet lenses in the compound eyes of moths consists of an array of excessive cuticular protuberances, termed corneal nipples. We have investigated the moth-eye corneal nipple array of the facet lenses of 19 diurnal butterfly species by scanning electron microscopy, transmission electron microscopy and atomic force microscope, as well as by optical modelling. The nipples appeared to be arranged in domains with almost crystalline, hexagonal packing. The nipple distances were found to vary only slightly, ranging from about 180 to 240 nm, but the nipple heights varied between 0 (papilionids) and 230 nm (a nymphalid), in good agreement with previous work. The nipples create an interface with a gradient refractive index between that of air and the facet lens material, because their distance is distinctly smaller than the wavelength of light. The gradient in the refractive index was deduced from effective medium theory. By dividing the height of the nipple layer into 100 thin slices, an optical multilayer model could be applied to calculate the reflectance of the facet lenses as a function of height, polarization and angle of incidence. The reflectance progressively diminished with increased nipple height. Nipples with a paraboloid shape and height 250 nm, touching each other at the base, virtually completely reduced the reflectance for normally incident light. The calculated dependence of the reflectance on polarization and angle of incidence agreed well with experimental data, underscoring the validity of the modelling. The corneal nipples presumably mainly function to reduce the eye glare of moths that are inactive during the day, so to make them less visible for predators. Moths are probably ancestral to the diurnal butterflies, suggesting that the reduced size of the nipples of most butterfly species indicates a vanishing trait. This effect is extreme in papilionids, which have virtually absent nipples, in line with their highly developed status. A similar evolutionary development can be noticed for the tapetum of the ommatidia of lepidopteran eyes. It is most elaborate in moth-eyes, but strongly reduced in most diurnal butterflies and absent in papilionids.     

Suckling behaviour and fertility in rhesus macaques (Macaca multatta)

Lactation has been proposed as the main proximate mechanism regulating female fertility among non–seasonally breeding mammals. Less is known about the effects of lactation upon fertility among seasonal species, where photoperiodic control of reproduction is widely believed to have an overriding influence. In neither case is it clear which of the different components of lactation influence fertility and in which ways. In this paper, the links between suckling patterns and the likelihood of conception during the following breeding season are examined. Data were collected at the Madingley rhesus colony, where reproduction is strictly seasonal. It is shown that fertility was depressed in lactating females when compared to non–lactating ones. Within the lactating group, mothers who suckled their infants frequently, and with a great number of independent nipple contacts in each bout, failed to conceive during the following breeding season. It is suggested that early differences in suckling patterns might reflect differences in maternal condition; and that differences during the breeding season may affect the quality of the ovulations. A considerable decrease in both the frequency of suckling and the number of nipple contacts per bout seemed to be required for conception to occur. On the other hand, neither the duration of the suckling bouts, nor the duration of the intervals between bouts, were found to be associated with the likelihood of conception.     

The S nipple-areola reconstruction

Nipple reconstruction is performed as a last stage in breast reconstruction following mastectomy. Various methods of nipple reconstruction have been described, most of them utilizing either free composite grafts or local flaps. The main problem encountered using either method is the gradual absorption and flattening of the nipple. The technique we used in reconstructing 22 nipples, in preference over the various methods accepted in breast reconstruction, achieves a long-standing, protruding nipple constructed from two large local flaps raised from an S-shape design. The technique is simple and permits freedom in choosing the height of the nipple, even in the presence of a mastectomy scar. The size of the nipple thereby constructed is in excess of what was expected. Shrinkage occurs during the first 2 months, and the resulting size is more than adequate. The areola is reconstructed by a full-thickness skin graft harvested from a nonhairy area of the upper inner thigh. The local flaps lack the necessary color, which is achieved by tattooing.     

Litter Size Determines Circadian Periodicity of Nursing in Rabbits

Nursing in rabbits occurs inside the nest with circadian periodicity. To determine the contribution of suckling stimulation in regulating such periodicity, we varied the size of the litters provided (1, 2, 4, or 6–8 pups). Nursing does, kept under a 14:10 (L:D) photoperiod, were continuously videotaped from parturition into lactation day 15. Although parturitions occurred throughout the day, a significant negative linear correlation (p?<?0.0001; r?=??0.68) was evident between time of delivery and time of nursing on lactation day 1, regardless of newborn number: longer intervals between these two events were seen in does delivering in the early morning than in those that gave birth late in the day. In rabbits suckling 6–8 pups, a Rayleigh analysis revealed that the population vector best describing their nursing pattern (across lactation days 1–15) had a phase angle?=?58° (corresponding to solar time 0352?h and rho?=?0.78; p?<?0.001). In contrast, the nursing pattern of does nursing litters smaller than 6 pups did not show circadian periodicity; rather, mothers showed multiple entrances into the nest box throughout the day. Cluster analysis revealed that the main equilibrium point of intervals between suckling bouts shifted from 24?h (6–8 pups) to 6?h (4 and 2 pups) and to as low as 4?h with 1 pup. In the groups nursing 2, 4, or 6–8 pups, most nursing episodes were followed by food and water intake. Those mothers also showed self-grooming of the ventrum and nipples after nursing. The incidence of these behaviors was lower in does nursing 1 pup. In conclusion, nursing in rabbits spontaneously occurs with circadian periodicity, but it is largely modulated by a threshold of suckling stimulation. (Author correspondence: gabygmm@gmail.com)     

A new tool for measuring the suckling stimulus during breastfeeding in humans: the orokinetogram and the Fourier series

The Fourier series was used to analyse the oral movements recorded by the orokinetogram during breastfeeding in human babies. This is a new method that allows recording of oral movements without introducing any extrinsic element between the nipple and the mouth of the baby. The advantage of displaying suckling activity after fast Fourier transform (FFT) is that this algorithm allows storage, quantification and frequency analysis of the oral movements throughout a suckling bout, which enables the total oral activity to be measured. Two types of oral movements are found: slow high amplitude (SHA) and fast low amplitude (FLA). FLA movements may be derived from peristaltic movements of the tongue that result in tickling stimuli to the mechanoreceptors of the nipple and milk expression. The frequency bandwidth of oral movements is wider (0-8 Hz) than has been described previously (0-3 Hz) and this is due to the presence of the FLA oral movements. An indirect measurement of the energy of oral movements during suckling is obtained by the pattern of energy distribution used in each individual frequency band by oral movements. This pattern changes in relation to the periods of continuous and intermittent suckling activity. SHA and FLA oral movements are more intense during continuous suckling. Statistical analysis showed a correlation between the energy of SHA and FLA waves throughout the suckling bout, and also that the highest level of energy during suckling activity is displayed during the first 2 min. The novel tools described in this paper allow investigation of the role of suckling stimulus in reflex hormone release and other mother-infant interactions.     

Studying nanostructured nipple arrays of moth eye facets helps to design better thin film solar cells

Nipples on the surface of moth eye facets exhibit almost perfect broadband anti-reflection properties. We have studied the facet surface micro-protuberances, known as corneal nipples, of the chestnut leafminer moth Cameraria ohridella by atomic force microscopy, and simulated the optics of the nipple arrays by three-dimensional electromagnetic simulation. The influence of the dimensions and shapes of the nipples on the optics was studied. In particular, the shape of the nipples has a major influence on the anti-reflection properties. Furthermore, we transferred the structure of the almost perfect broadband anti-reflection coatings to amorphous silicon thin film solar cells. The coating that imitates the moth-eye array allows for an increase of the short circuit current and conversion efficiency of more than 40%.     

Isotope recycling in lactating dogs (Canis familiaris)

Isotope-based techniques for the measurement of water turnover, energy expenditure, and milk intake often assume that there is no recycling of isotopes once they have left the labeled animal. In experiments involving lactating females or their suckling offspring, there are several possible routes of isotope recycling. These include the consumption of labeled milk by offspring, the ingestion of labeled excreta, and the rebreathing of exhaled labeled CO(2) or water vapor by both mother and offspring. Isotope recycling might be especially important during lactation because the offspring are in close contact with each other and their mother for prolonged periods. We show here in 24- to 30-day-old domestic dog Canis familiaris puppies that there was no detectable transfer of (18)O or (2)H from labeled to unlabeled pups in two litters (16 pups, 8 labeled, 8 unlabeled) that were weaned early and independent of their mother. However, there was a significant transfer of both isotopes from labeled to unlabeled pups and from labeled pups to their mothers in nine equivalent nursing litters of the same age (27 labeled, 26 unlabeled pups). The increases in enrichment of isotopes in unlabeled offspring were greater than the increases in enrichment of the mothers. This indicates that maternal ingestion of offspring excreta and subsequent transfer of isotope in milk is not the sole pathway of recycling. Additional routes must also be important, such as exchange of isotope between pups on saliva-coated nipples and perhaps direct ingestion of excreta by unweaned young. Recycling is unlikely to be an important factor when determining maternal metabolic rate during peak lactation in domestic dogs. However, experiments that are designed to assess the energy demands of pups and isotope-based estimates of water turnover in offspring may need to take into account any effects of isotope recycling. In a theoretical example, removing the effects of recycling increased the measured energy expenditure in pups by up to 7% and increased the calculated elimination rates of both isotopes by up to 11.1% in (18)oxygen and 10.9% in (2)hydrogen.     

Major role of suckling stimulation for inhibition of estrous behaviors in lactating rabbits: acute and chronic effects

Lactation in rabbits induces anestrus: sexual receptivity and scent-marking (chinning) are reduced despite the brevity of suckling (one daily nursing bout, lasting around 3 min). The mechanisms underlying this effect are unknown but, as chinning, lordosis, and ambulation in an open field are immediately inhibited by the peripheral stimulation received during mating we hypothesized that, across lactation, suckling stimulation would provoke a similar effect. To test this possibility we provided litters of 1, 3, 5, or 10 pups across lactation days 1-15 and quantified chinning and ambulation frequencies, the lordosis quotient, and milk output. Baseline chinning frequency, determined before the daily nursing bout, was low across lactation days 1-15 in does nursing 3, 5 or 10 pups but it increased steadily across days 1-10 in rabbits suckling one pup. Yet, a single young was sufficient to abolish chinning for about 1h, after which this behavior rose again. Suckling litters of all sizes reduced (but did not abolish) ambulation frequency, both chronically (baseline levels declined across days 1-5) and acutely. Sexual receptivity was significantly reduced on lactation day 15 only in does that had nursed 10 pups. Large litter size promoted a larger milk output and a normal duration of nursing episodes. Results support a major role of suckling stimulation for the suppression of estrous behaviors and ambulation through as yet unidentified mechanisms.     

Role of nipple stimulation in the suppression of estrous cyclicity during extended lactation in the rat

The role of nipple stimulation in the suppression of the estrous cycle during extended lactation was studied in rats subjected to either total, partial, or sham excision of the nipples. Each female cohabited with 4 pups, 4-14 days old, over a period of 70 days postpartum, during which vaginal smears were recorded daily. Initially, regardless of the presence of nipples, all rats exhibited a postpartum diestrus that lasted for 12-20 days. Intact females (bearing 6 pairs of nipples) continued to exhibit successive prolonged diestrous phases over 70 days of lactation. A comparable result was obtained with females bearing only the anterior pair of nipples, which, in a separate experiment, was found to be the most frequently suckled pair. However, females devoid of nipples resumed regular (4-day) estrous cycles between Days 12 and 27 postpartum, in spite of their continuous contact with pups. Thus, when lactation is prolonged beyond the normal time of weaning (Day 21 postpartum), stimulation of the nipples by sucking becomes indispensable for the continued arrest of the estrous cycle. The possible mechanisms underlying this phenomenon are discussed.     

强壮水螅的特征及其与寡水螅的种间差别问题

一种广泛分布于黑龙江省的大型具柄水螅被暂定为强壮水螅Hydra robusta(IT(?) 1947)。这一中国品系与IT(?)(1947)在日本报告的新种主要特征相同,包括具有精巢乳头。只是精巢乳头不稳定。在实验室条件下第一次有性生殖时每个雄体都发生精巢乳头,但第二次有性生殖时,同一群的后代可全部失去精巢乳头。这是介于强壮水螅与寡水螅Hydra oligactis Pallas(1766)之间的中间性状。从而导致作者做出结论,认为过去其他学者所主张的,以精巢乳头的有无做为鉴别特征来区分强壮水螅和寡水螅是不可靠的。因此,本文又检验了此种水螅的体细胞染色体,证明2n=30,其中第二对染色体上有明显近中位置的次缢痕,最小染色体的长度为最大染色体长度的二分之一以上。这些可做为此种水蝗的鉴别特征。对Niiyama(1944)和Datta(1970)在寡水螅上所做的染色体研究也做了比较及讨论。     

Randomised controlled trial of breast shells and Hoffman's exercises for inverted and non-protractile nipples.

OBJECTIVE--To determine the value of recommending breast shells or Hoffman''s exercises, or both, to pregnant women with inverted or non-protractile nipples who intend to breast feed. DESIGN--Randomised controlled trial with a two treatment by two level factorial design. SETTING--Antenatal clinics in a district general hospital and the community. SUBJECTS--96 nulliparous women recruited between 25 and 35 completed weeks in a singleton pregnancy with at least one inverted or non-protractile nipple. MAIN OUTCOME MEASURES--Anatomical change of nipples, judged blindly before first breast feeding, and success of breast feeding reported by postal questionnaire six weeks postnatally. RESULTS--Sustained improvement in nipple anatomy was more common in the untreated groups but the differences were not significant (52% (25/48) shells v 60% (29/48) no shells; difference -8% (95% confidence interval -28% to 11%) and 54% (26/48) exercises v 58% (28/48) no exercises; -4% (-24% to 16%)). 24 (50%) women not recommended shells and 14 (29%) recommended shells (21%; 40% to 2%) were breast feeding six weeks after delivery (p = 0.05), reflecting more women recommended shells both deciding to bottle feed before delivery and discontinuing breast feeding. The same number of women in exercise and no exercise groups were successfully breast feeding (0%; -20% to 20%). 13% of women approached about the trial (and planning to breast feed) did not attempt breast feeding. CONCLUSIONS--Recommending nipple preparation with breast shells may reduce the chances of successful breast feeding. While there is no clear evidence that the treatments offered are effective antenatal nipple examination should be abandoned.