Cortisol levels and aggression in neutered and intact free-roaming female cats living in urban social groups.

Free-roaming domestic cats in urban areas often live in defined social groups, and the breeding females in these groups tend to form structures of a matrilineal nature. In recent years, resulting from the growing populations of free-roaming cats in many cities worldwide, these cats are being managed using the Trap-Neuter-Return (TNR) method. The aims of this study were to explore the hypotheses that (a) neutering reduces aggression in the females living in such social groups; and (b) if such reduction does occur, that it might be accompanied by a reduction in cortisol levels. The study was conducted on eight cat feeding groups in residential neighbourhoods in Tel Aviv, Israel. The municipal veterinary department offers TNR services upon request. Cats are collected from the feeding group and returned to their original group after neutering. We found that neutered females showed reduced aggressiveness as well as reduced cortisol levels compared to the intact females. In addition, those intact females that displayed more aggression had higher cortisol levels compared to the less aggressive intact females. Based on the results of this study it is possible to suggest for the first time a possible relationship between cortisol levels and aggression in free-roaming female domestic cats. This study is an initial step in assessing the long-term effects of Trap-Neuter-Return (TNR) on the welfare of the individual cat. If cortisol levels in female cats are reduced after neutering, partly as a result of reduced social and reproductive pressures (as expressed by lower aggression of the neutered females), it is possible that TNR has an added beneficial role in cat welfare in addition to that of control of population size.

Magnetic compass orientation in the blind mole rat Spalax ehrenbergi.

The blind mole rat Spalax ehrenbergi is a solitary, subterranean rodent that digs and inhabits a system of branching tunnels, with no above-ground exits, which it never leaves unless forced to. To survive, the mole rat must be able to orient efficiently in its tunnel system. The sensory channels available for spatial orientation in the subterranean environment are restricted in comparison with those existing above ground. This study examined the possibility that the mole rat is able to perceive and use the earth's magnetic field to orient in space. Experiments were performed using a device constructed from a pair of electromagnetic 'Helmholtz coils', which create a magnetic field whose direction and strength can be altered. In the first experiment, we tested a group of mole rats (N=33) in an eight-armed maze under the earth's natural magnetic field to determine whether they have directional preferences for the location of their sleeping nest, food chamber and toilet site. A second group of mole rats (N=30) was tested for their directional preference after the earth's magnetic field had been experimentally shifted by 180 degrees. We found that the first group exhibited a significant preference (P<0.001) to build both their sleeping nest and their food store in the southern sector of the maze, whereas the second group shifted the location of their nests (P<0.01) and food store (P<0.05), to the northern sector of the maze, corresponding to the shift in the magnetic field. In the second experiment, we tested whether the magnetic compass orientation found in the first experiment depends on a light stimulus by testing a group of mole rats in the eight-armed maze under total darkness. No significant difference in directional preference between light and dark test conditions was observed. It can be concluded, therefore, that, in contrast to some amphibians and birds, magnetic compass orientation in the mole rat is independent of light stimulation. In the third experiment, we examined whether mole rats (N=24) use the earth's magnetic field as a compass cue to orient in a labyrinth. In the first stage (trials 1-13), the animals were trained to reach a goal box at the end of a complex labyrinth until all individuals had learned the task. In the second stage (trial 14), half the trained mole rats underwent another labyrinth trial under the earth's natural magnetic field, while the other half were tested under a magnetic field shifted by 180 degrees. We found a significant decrease (P<0.001) in performance of the mole rats tested under the shifted magnetic field compared with the group tested under the natural magnetic field. The findings from these experiments prove that the mole rat is able to perceive and use the earth's magnetic field to orient in space.

Reversed palatal perforation by upper incisors in ageing blind mole-rats (Spalax ehrenbergi).

Blind mole-rats (Spalax ehrenbergi) are fossorial solitary rodents that present striking morphological, physiological and behavioural adaptations to the subterranean environment in which they live. Previous studies have shown that mole-rats are specialised in tooth-digging. The rapid eruption-rate of their incisors has evolved to compensate for their excessive wear by excavation. Males use their incisors more than females for digging and fighting, and their rate of incisor eruption is significantly more rapid than in females. Since mole-rats use their incisors for digging throughout the year, we suggest that continuous mechanical pressure on their oral tissues concentrated at the apical sites of the upper incisors leads to cell and tissue fatigue. We provide evidence for 5 stages of palatal perforation by the upper incisors at their apical sites, with maximum perforation characterising aged males. Interspecies comparisons with 7 other fossorial and semi-fossorial rodent species, and with beavers, which expose their incisors to enormous mechanical pressure, revealed that this palatal perforation is unique to the male mole-rat. We suggest that while the fast eruption rate of incisors in the mole-rat compensates for the rapid wear resulting from digging, evolutionary adaptation to continuous tooth-digging is still ongoing, since the physical pressure of digging at the apical sites of the upper incisors leads to tissue destruction, breakage of the palatal bone and possibly to death, as a result of maxillary inflammation.

The structure of mast cell secretory granules in the blind mole rat (Spalax ehrenbergi).

Eosinophils, basophils, and mast cells produce and secrete active substances whose role is to attack invading parasites and protect the host. In this study we use morphometric methods to study mast cells in the blind mole rat (Spalax ehrenbergi). The subterranean and solitary way of life of this species has led to the evolutionary development of special anatomical, morphological, behavioral, and physiological adaptations. Because of its particular lifestyle, the mole rat is less exposed to parasites than other rodents. This could provide a unique model for research into the pathobiology of mast cells. The paracrystalline structure of the mast cell granule content is composed of parallel plates. Diffraction analysis of electron micrographs of thin sections of araldite-embedded tissues indicated that each crystal line plate is a periodic array of parallelograms. The crystal unit cell volume is approximately 930 nm(3), suggesting that each unit cell is composed of one heparin molecule and one to three additional adsorbed proteins. Morphometric data show that characteristics of the secretory granules of mast cells of the blind mole rat resemble those of other rodents. The mast cell unit granule volume in the present study was calculated to be 0.055 microm(3), similar to that of rat peritoneal mast cells.

Effect of photoperiod variation on testes and accessory sex organs in the male blind mole rat Spalax ehrenbergi.

The blind mole rat is a seasonally breeding fossorial rodent that is perceptionally blind. This study examines the effect of photoperiod on the morphology and histology of the male mole rat reproductive system, three groups of male mole rats were maintained in the laboratory under short day (SD) conditions (9L: 15D); long day (LD) conditions (15L:9D); and constant darkness (CD), and compared to animals trapped in the field (FL). It was found that the field animals revealed higher testes and prostate gland weights, higher prostate tubuli volume (v*) and lower testes tubuli volume (v*) compared to the other three groups. Distribution of the tubuli in the testes (Vv) was low in the FL group compared to the SD and LD groups but still higher than in the CD group. Distribution of lumen in the testes (Vv) was higher in the CD group in comparison to the other three groups. Distribution of interstitial tissue in the testes (Vv) was higher in the FL group than in the other three groups. Electrolytes and elements secreted from the prostate gland did not differ among the four groups. In the FL group distribution of the tubuli (Vv) in the prostate gland was low and lumen ratio (Vv) was high compared to the other three groups. Distribution of connective tissue in the prostate gland did not differ among all four groups. Testosterone levels and total sperm count was highest in the FL group. Sperm production was noted in all groups; however spermatid and spermatozoa cell production was higher in the FL group. This study shows that photoperiod could be important in initiating timing in the breeding season but that certain other conditions which are absent in the laboratory appear to be responsible for successful breeding in the field.

Urinary testosterone levels in the male blind mole rat (Spalax ehrenbergi) affect female preference.

This study investigated the sexual attraction of female blind mole rats to four groups of male mole rats: (a) intact males raised in captivity; (b) intact males trapped in the field; (c) captive males injected with testosterone; (d) captive castrated males. In the first part we measured blood testosterone, androstenedione, and dihydrotestosterone (DHT) levels, by radioimmunoassay; and urine testosterone levels, measured by GC-MS. The second part examined the relationship between urine testosterone levels in males and their attractiveness to females. Higher blood and urine testosterone levels were found in the field animals and in those injected with testosterone compared to captive intact or castrated animals: urine testosterone levels in the two other groups were not detectable. Blood androstenedione levels were also higher in the field animals and in those injected with testosterone compared to captive intact or castrated mole rats. Blood dihydrotestosterone levels were not detectable in all four experimental groups. Female mole rats chose to spend a longer period of time next to males with high blood and urine testosterone levels and high blood androstenedione levels than next to those with lower levels of these hormones. Because courtship and sexual behavior are influenced both by high levels of blood and urine testosterone and high levels of blood androstenedione, we suggest that the low levels of courtship and other sexual behavior in captive mole rats may be related to the lack of female attraction to these males, which display low levels of all three parameters.

Seismic communication signals in the blind mole-rat (Spalax ehrenbergi): electrophysiological and behavioral evidence for their processing by the auditory system.

Based on morphological and behavioral findings we suggest that the seismic vibratory signals that blind mole-rats (Spalax ehrenbergi) use for intraspecific communication are picked up from the substrate by bone conduction and processed by the auditory system. An alternative hypothesis, raised by others, suggest that these signals are processed by the somatosensory system. We show here that brain stem and middle latency responses evoked by vibrations are similar to those evoked by high-intensity airborne clicks but are larger in their amplitudes, especially when the lower jaw is in close contact with the vibrating substrate. Bilateral deafening of the mole-rat or high-intensity masking noise almost completely eliminated these responses. Deafening also gradually reduced head-drumming behavior until its complete elimination about 4-6 weeks after surgery. Successive vibrations, at a rate of 0.5 vibrations/s, elicited prominent responses. At rates higher than 2 vibrations/s the amplitude of the brain stem response did not change, yet the middle latency response disappeared almost completely. It is concluded that the seismic signals that mole rate use for long distance communication are indeed processed primarily by the auditory system.

Social stress in neighboring and encountering blind mole-rats (Spalax ehrenbergi).

Blind mole-rats (Spalax ehrenbergi) are solitary aggressive subterranean rodents. They inhabit individual territories, comprised of branched tunnels. Each such tunnel system is completely separate from that of any neighboring mole-rat. Although intraspecific encounters between neighbors are infrequent, when they do occur, they may result in the injury or death of one or both animals. Avoidance of encounters may be due to the awareness of a neighbor's whereabouts through scent-marking and/or seismic (vibratory) communication. The present study was intended to examine whether encounters between individual mole-rats result in physiological stress. Two experimental conditions were designed to simulate natural situations: a brief encounter between two neighboring mole-rats, taking place either once or several times and long-term residency of neighbors whose only contact was either vibratory or vibratory plus odor communication. Blood samples were taken before, during, and after encounters in the first experiment and at set intervals in the second. The blood variables measured were blood glucose levels (BGL) and neutrophil/lymphocyte ratio (N/L). Blood glucose levels and neutrophil/lymphocyte ratio ratios increased in both members of encountering pairs. Long-term residency with a neighbor resulted in the establishment of a dominant-subordinate relationship through vibratory communication only, with increased neutrophil/lymphocyte ratio ratio in the subordinate males. However, long-term residency of males exposed to both vibrations and odors of neighboring males resulted in the death of both individuals. It seems that brief direct encounters and long-term neighboring conditions without physical contact are sufficient to cause severe stress to mole-rats. It is possible that in the wild, in some situations in which neighboring mole-rats cannot avoid constant exposure to each other's vibratory and odor signals, the consequent extensive stress may result in death.

Morphology and cytology of the nasal cavity and vomeronasal organ in juvenile and adult blind mole rats (Spalax ehrenbergi).

The blind mole rat (Spalax ehrenbergi) is a fossorial solitary rodent which exhibits extensive intraspecific aggression and uses scent markings to deter contraspecific invaders. Mole rats of different ages were captured near Tel Aviv, Israel, and sacrificed by an overdose of Xylazine hydrochloride. Olfactory epithelium sites from the nasal cavity (NC) and the vomeronasal organ (VNO) were dissected and fixed for light and electron microscopy. The mole rat's olfactory epithelium of the NC consists of several cell types, of which two types are supporting cells that comprise both microvilli and cilia but differ in staining and the presence of rough endoplasmic reticulum. The third type has no cilia. Secretory goblet cells were frequent among supporting cells of adults alone. Two types of receptor cells protrude into the NC with olfactory knobs at their apical region; one type has up to 177.6 +/- 9.4 cilia per knob plus microvilli, while the other type has only microvilli. The third type of sensory cell has no knob and contains microvilli only. The basal epithelium layer consists of short-bodied cells with round nuclei. The VNO of the mole rat is situated beneath the nasal septum, consisting of supporting, sensory, and basal cell types, with many cilia at the apical portion. At its anterior part, the VNO is connected to the NC by narrow canals. The abundance of cilia and microvilli in the mole rat olfactory cells provides the first anatomical evidence for their olfactory acuity. Such acuity is important in mole rats, compensating for their loss of vision and enabling them to detect and avoid rivals prior to potential aggressive encounters as well as to select food plants during foraging.

Mole-rat harderian gland secretions inhibit aggression

The harderian gland secretions of mole-rats, Spalax ehrenbergiusually released by self-grooming, include odorous substances which are sex dependent. Male secretions were the most attractive to both sexes, while female secretions were attractive to males but not to other females. The rate of attacks by females towards intact males was higher than towards males whose harderian gland had been removed. However, grooming by intact male mole-rats decreased the rate of attacks by their opponents, while grooming by males without harderian glands did not; thus the male harderian secretions appear to have appeasement qualities. Grooming by females with and without harderian glands failed to reduce aggression. Unlike intact males, those without harderian glands had almost no volatiles on their fur, and thus are probably not considered to be a threat to conspecifics. Gas chromatography spectra showed that substances of harderian origin were added to the fur during grooming. Some of these substances remained on the fur long after the animal ceased grooming, and appear to give the animal its specific odour, but some volatile substances peaked briefly after grooming, and were probably responsible for the decline of aggression that occurred after grooming. Although grooming has long been considered to be a displacement activity, we suggest that in the mole-rat its performance is too risky to be merely this, and it has acquired the meaning of appeasement through the release of chemical cues.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour

Circadian patterns of locomotor activity and body temperature in blind mole-rats, Spalax ehrenbergi.

A wide variety of organisms exhibit circadian rhythms, regulated by internal clocks that are entrained primarily by the alternating cycle of light and darkness. There have been few studies of circadian rhythms in fossorial species that inhabit a microenvironment where day-night variations in most environmental parameters are minimized and where exposure to light occurs only infrequently. In this study, daily patterns of locomotor activity and body temperature (Tb) were examined in adult blind mole-rats (Spalax ehrenbergi). These fossorial rodents lack external eyes but possess rudimentary ocular structures that are embedded in the Harderian glands and covered by skin and fur. Most individual mole-rats exhibited circadian rhythms of locomotor activity, but some animals were arrhythmic. Individuals that did exhibit robust rhythms of locomotor activity also showed rhythms of Tb. In most cases, Tb was highest during the phase of intense locomotor activity. Locomotor activity rhythms could be entrained to light:dark cycles, and several mole-rats exhibited entrainment to non-24-h light cycles (T-cycles) with period lengths ranging from T = 23 h to T = 25 h. Some individuals also showed entrainment to daily cycles of ambient temperature. There was considerable interindividual variation in the daily patterns of locomotor activity among mole-rats in virtually all the conditions of environmental lighting and temperature employed in this study. Thus, whereas it appears likely that photic cues have a significant role in the entrainment of circadian rhythms in mole-rats, the amount of variability in rhythm patterns among individuals appears to be much greater than for most species that have been studied.

Putative melatonin receptors in the blind mole rat harderian gland.

The blind mole rat (Spalax ehrenbergi) displays daily and seasonal rhythms. Melatonin, secreted nocturnally by the pineal gland, is also produced in the harderian gland and affects its morphology in rodents. We report here on the presence of putative melatonin receptors in the blind mole rat harderian gland, located in the microsome-enriched fraction of the cells. Equilibrium 125I-melatonin binding studies indicated high- and low-affinity melatonin binding sites in the female (apparent Kd 10 pM and 2.4 nM, respectively) and low-affinity sites in the male (apparent Kd 2.6 nM) mole rat. The binding sites were not significantly affected by season. Castration increased the density of high-affinity binding sites in males and low-affinity binding in females. 125I-melatonin binding to the gonadectomized mole rat preparation was inhibited by serotonin > 2-iodomelatonin > or = memelatonin > 5-methoxytryptamine. The guanine nucleotide analogs, guanosine 5'-O-[3-thio-triphosphate] and guanosine 5'-O-[2-thio-diphosphate], inhibited specific 125I-melatonin binding, whereas 5'-guanylyl imido-diphosphate was less potent. These results indicate for the first time the presence of GTP-sensitive melatonin binding sites in the blind mole rat harderian gland, and suggest that their expression is under control of sex steroids.

Effect of the neuroactive steroid alpha-THDOC on staircase test behavior in mice.

This study examined the effect of the neuroactive steroid 3 alpha, 5 alpha-tetrahydrodeoxycorticosterone (alpha-THDOC) as compared to the benzodiazepines diazepam and midazolam and the barbiturate phenobarbital on the number of rearing events and the number of steps ascended in the mouse staircase test. The benzodiazepines, phenobarbital and alpha-THDOC all reduced rearing activity at doses that did not affect climbing. The rearing-suppression effect of the benzodiazepines and alpha-THDOC, but not of phenobarbital, was blocked by the benzodiazepine antagonist flumazenil. It appears that, although such neuroactive steroids, like barbiturates, bind to distinct sites within the chloride ion channel of the gamma-aminobutyric acid type A (GABAA) receptor complex, alpha-THDOC behavioral activity is modulated by the benzodiazepine recognition site.

Quantitative microscopic changes in the mole rat's accessory sex organs during an annual cycle.

BACKGROUND: The mole rat, Spalax ehrenbergi, is a solitary, aggressive subterranean rodent. The present study summarizes a year-round investigation of morphological changes in the mole rat's accessory sex organs. METHODS: Mole rats maintained in the laboratory were killed after 3 months of acclimation; additional animals trapped in the wild were killed immediately. The accessory sex organs were processed for routine histological examination. Tissues were fixed in Bouin's solution, embedded in paraffin blocks, and stained by hematoxylin-eosin. A systemic sampling approach was used to photomicrograph the tissues for histomorphometric assessment. RESULTS: The volume fraction (Vv, mean +/- SEM) of prostate connective tissue from animals kept in captivity increased significantly in January (0.49 +/- 0.05 mm3/mm3) and April (0.43 +/- 0.04 mm3/mm3) but only 0.26 +/- 0.03 mm3/mm3 in November. In the field group, the Vv of prostate connective tissue was significantly higher in January (0.58 +/- 0.08 mm3/mm3) and April (0.62 +/- 0.08 mm3/mm3) and lower in November (0.44 +/- 0.03 mm3/mm3) and February (0.43 +/- 0.03 mm3/mm3), with a concomitant decrease in prostate tubuli and lumen. The prostate tubuli star volume (v*) in laboratory animals increased in November (0.009 +/- 0.002 mm3) and May (0.09 +/- 0.02 mm3). The same pattern was shown in the field group, with a significant increase in December (0.012 +/- 0.002 mm3) and March (0.007 +/- 0.001 mm3). The Cowper tubuli Vv in the captive animals increased during February (0.24 +/- 0.02 mm3/mm3), with a concomitant reduction in the connective tissue (0.05 +/- 0.02 mm3/mm3). The Cowper tubuli v* in same animals increased in December, April, and July (1.37 +/- 0.18 x 10(-4) mm3, 0.94 +/- 0.10 x 10(-4) mm3, 1.52 +/- 0.20 x 10(-4) mm3, respectively). In field group, a slight decrease in star volume took place from November to May (1.25 +/- 0.16 mm3 to 0.39 +/- 0.05 mm3, respectively). Testosterone levels appeared to be higher in the field group than in the laboratory group. In December the values were 1.62 +/- 0.15 ng/ml in the field group and 0.55 +/- 0.12 ng/ml in the laboratory group, and in May the laboratory group values were 1.66 +/- 0.12 ng/ml. CONCLUSIONS: In captivity and in the field, male mole rats probably undergo an annual cycle of accessory gland tissue structural changes that are correlated with testosterone secretion.

Grooming secretions and seasonal adaptations in the blind mole rat (Spalax ehrenbergi).

The postulated thermoregulatory function of grooming in the "blind" mole rat (Spalax ehrenbergi), a solitary, highly aggressive subterranean rodent, was tested by subjecting individuals to extreme hot and cold environments and measuring their grooming frequency and thermoregulatory capacity. It was found that exposure to heat stress during both the winter and the summer failed to elicit grooming in isolated mole rats, even though their body temperature was significantly elevated. Thus, unlike Mongolian gerbils (Meriones unguiculatus), mole rats do not seem to have a mechanism for lowering body temperature by grooming. However, at low temperatures, grooming behavior significantly increased during both seasons, although a decline in body temperature was recorded only during the summer. Because grooming in mole rats expresses Harderian lipids onto the fur, it is suggested that in the winter, under cold stress, this behavior might prevent a drop in body temperature through spreading insulating Harderian lipids.

Quantitative histomorphology of the blind mole rat harderian gland.

Anatomical, histological and morphometric studies have been performed on the harderian gland and its surroundings in the blind mole rat (Spalax ehrenbergi). The gland is tubuloalveolar with no true duct system. All ducts within the gland are formed by a single epithelial cell type and drain into a wide secretory duct. This opens into the conjunctival sac which serves as a reservoir for harderian secretions. Drainage from the conjunctival sac follows 2 possible routes: one through the nasolacrimal duct to the external nasal cavity, the other through a unique excretory duct that emerges from the anteromedial part of the conjunctival sac and runs through the dermis to the skin, opening at the base of a hair follicle. The function of this newly described duct is discussed. Morphometric studies revealed that the lumen volume fraction in the female, slightly smaller than that of the male during the summer, becomes significantly greater during the winter breeding season. The dimorphism and seasonal variations found in the gland acini suggests that the gland may be implicated in pheromone production.

Interbirth interval and duration of pregnancy in hares.

The possibility of inducing superfetation in hares by artificial insemination (AI) was investigated. AI performed on various days during the second half of gestation did not result in new pregnancies, but all ongoing pregnancies were terminated 1-4 days after AI. We suggest that copulation during the last week of pregnancy, common among captive hares, may have a similar effect to that of AI in terminating pregnancies and inducing early deliveries. Intervals between successive deliveries that are shorter than the normal duration of gestation do not necessarily indicate superfetation and this phenomenon may be rare among hares in captivity as well as in the wild.

Retinal projections in the blind mole rat: a WGA-HRP tracing study of a natural degeneration.

The mole rat Spalax ehrenbergi is a fossorial rodent. Although its peripheral visual system--eye and optic nerve--is highly degenerated, it shows some sensitivity to light. However, in the usual sense, it is essentially blind. An auditory take-over of the visual lateral geniculate nucleus and at least part of the visual cortices was recently demonstrated. In order to visualize the retinal projections during ontogeny, we used an anterograde tracing technique, with monocular injection of wheat germ agglutinin-labeled horseradish peroxidase (WGA-HRP). In the newborn mole rat the retina projects to most of its normal targets as compared with seeing rodents, with bilateral projections to the suprachiasmatic nuclei, the dorsal and ventral lateral geniculate nuclei, the lateroposterior nuclei, the optic tract nuclei and the superior colliculi. During the course of ontogeny, the retinohypothalamic connection is stabilized but the main optic tract undergoes progressive degeneration. In adults, only a few retinal fibers enter the contralateral ventral lateral geniculate nucleus, the lateroposterior nucleus, the optic tract nucleus and the superior colliculus. No retinal fibers could be detected in the dorsal lateral geniculate nucleus. Thus, the retinofugal projections in the adult mole rat could explain its reduced sensitivity to light, whereas the complete degeneration of the retino-dorsal lateral geniculate nucleus projection could underlie the invasion of auditory input into this normally visual center.

Hypothalamic neuronal activity associated with onset of pseudopregnancy in the rat.

The electrical activity of several hypothalamic sites, before and following cervical stimulation, was examined in an attempt to evaluate the central nervous system processes involved in the induction of pseudopregnancy in the rat. Cervical stimulation, resulting in pseudopregnancy, induced a sequence of neuronal changes at: 3.8, 4.4, 11.6, 20.1, 30.8, 44.5, 51.2, 51.7, 62.4, 85.1 and 111.5 min after stimulation in the preoptic area, lateral hypothalamus, ventromedial hypothalamus, preoptic area, suprachiasmatic nucleus, lateral hypothalamus, preoptic area, ventromedial hypothalamus, lateral hypothalamus and anterior hypothalamus, respectively. These results suggest that the preoptic region contains both the facilitatory neuronal mechanism and also an inhibitory system. The latter could tonically inhibit the expression of the nocturnal prolactin surge. We conclude that the sequence of neuronal events observed in various hypothalamic areas is in part responsible for the process which results in pseudopregnancy.