Phase response curve to melatonin in a putatively diurnal rodent, Octodon degus

The degu (Octodon degus) is a diurnal rodent, although phase inversions to nocturnal behavior have been reported under specific laboratory conditions. The reliability of this animal as a diurnal model of sleep therefore requires further characterization of intrinsic circadian pacemaker properties. A phase response curve to light has been reported in the degu, and is consistent with other diurnal animals. This study reports a phase response curve to melatonin in the degu, which is distinct in orientation from the light curve.     

Odor-specific effects on reentrainment following phase advances in the diurnal rodent, Octodon degus

Reentrainment following phase shifts of the light-dark (LD) cycle is accelerated in Octodon degus in the presence of olfactory social cues (i.e., odors) produced by conspecifics. However, not all odors from conspecifics were effective in facilitating reentrainment after a phase advance. In the current experiments, we examined whether nonanimal odors, odors from another species, or conspecific odors, including those manipulated by steroid hormones, can cause the same increased reentrainment of wheel-running activity as odors from an intact, adult female degu. A variety of odors, each selected to probe a particular aspect of the reentrainment acceleration phenomenon, were presented to a group of phase-shifting female degus. The shifting females (test animals) responded to odors of intact, female degu donors with decreased reentrainment time, but odors of ovariectomized (OVX), OVX with a single hormone replacement capsule (estradiol or progesterone) or phase-shifting females had no effect. Multiple males were effective odor donors, whereas a single male was ineffective in earlier studies. Rats and cloves were not effective in accelerating reentrainment. Furthermore, odors from rats delayed reentrainment. We conclude that the odors that effectively accelerate degu reentrainment after a phase advance of the LD cycle are species specific. We also report that repeated phase shifts, followed by complete recovery of phase relationships, do not alter the rate of recovery from a phase shift over time. These data suggest that in O. degus, a social species, odors may reinforce and strengthen the salience of the photic zeitgeber and/or facilitate synchronization of rhythms between animals.     

Chronotype changes during puberty depend on gonadal hormones in the slow-developing rodent, Octodon degus

During puberty, human adolescents develop a later chronotype, exhibiting a delay in the timing of rest and activity as well as other daily physiological rhythms. The purpose of this study was to determine whether similar changes in chronotype occur during puberty in a laboratory rodent species, and, if so, to determine whether they are due to pubertal hormones acting on the circadian timekeeping system. To test this hypothesis, we carefully tracked daily activity rhythms across puberty in the slow-developing rodent Octodon degus. We confirmed that male degus showed a large reorganization of activity rhythms that correlated with secondary sex development during puberty, including a loss of bimodality and a 3-5 h phase-advance. Similar to humans, this circadian reorganization showed distinct sex differences, with females showing little change during puberty in two separate experiments. Prepubertal gonadectomy (GDX) eliminated the changes, whereas SHAM gonadectomy had little impact. Therefore, gonadal hormones are likely to play a role in pubertal changes in chronotype in this rodent species. Using evidence from a variety of species, including our recent studies in the rat, we conclude that chronotype changes during puberty are a well-demonstrated phenomenon in mammals.     

Testicular hormones modulate circadian rhythms of the diurnal rodent, Octodon degus

Sex differences have been identified in a variety of circadian rhythms, including free-running rhythms, light-induced phase shifts, sleep patterns, hormonal fluctuations, and rates of reentrainment. In the precocial, diurnal rodent Octodon degus, sex differences have been found in length of free-running rhythm (tau), phase response curves, rates of reentrainment, and in the use of social cues to facilitate reentrainment. Although gonadal hormones primarily organize circadian rhythms during early development, adult gonadal hormones have activational properties on various aspects of circadian rhythms in a number of species examined. Gonadectomy of adult female O. degus did not influence tau, phase angle of entrainment, or activity patterns in previous experiments. The present experiment examined the role of gonadal hormones in adult male degus' circadian wheel-running rhythms. We predicted that male gonadal hormones would have an activational effect on some aspects of circadian rhythms, particularly those in which we see sex differences. Phase angles of entrainment, tau, length of the active period (alpha), maximum and mean activity levels, and activity amplitude were examined for intact and castrated males housed in LD 12:12. Responses to light pulses while housed in constant darkness (DD) were also compared. Castration had no significant effect on tau or light-induced phase shifts. However, castration significantly increased phase angle of entrainment and decreased activity levels. The data indicate that adult gonadal steroids are not responsible for the sex differences in endogenous circadian mechanisms of O. degus (tau, PRC), although they influence activity level and phase angle of entrainment. This is most likely due to masking properties of testosterone, similar to the activity-increasing effects of estrogen during estrus in O. degus females.     

Phase Response Curve to Melatonin in a Putatively Diurnal Rodent,Octodon degus

The degu (Octodon degus) is a diurnal rodent, although phase inversions to nocturnal behavior have been reported under specific laboratory conditions. The reliability of this animal as a diurnal model of sleep therefore requires further characterization of intrinsic circadian pacemaker properties. A phase response curve to light has been reported in the degu, and is consistent with other diurnal animals. This study reports a phase response curve to melatonin in the degu, which is distinct in orientation from the light curve.     

Circadian organization of the diurnal Caviomorph rodent, Octodon degus

The Octodon degus, or degu, is an excellent animal model for studying the theoretical and neural underpinnings of diurnality. The power of this model comes from their unique evolutionary lineage, long lives, and relative ease of care in the laboratory for a non-domesticated species. We have summarized the field and laboratory data indicating the critical variables that influence the degus' phase preference and the possible mechanisms for the phase flexibility observed in the field and laboratory. We also review studies examining the physiology and anatomy of light and non-photic inputs to the degu circadian system and studies of the circadian pacemaker itself, with particular emphasis placed on characteristics that appear to be convergent adaptations to a diurnal niche. Finally, we begin to seek the origin for the diurnally-phased activity output of the degu, although we conclude that significant work remains to be done.     

No infanticide in the hystricognath rodent, Octodon degus: does ecology play a role?

The behavior of male and female Octodon degus (Hystricognathi; Octodontidae) was studied in captivity to examine the occurrence of non-parental infanticide, which involves the killing of immature infants by adult conspecifics other than the genetic parents. Sexually inexperienced male and female, and lactating female degus were tested for their behavior toward genetically unrelated, and socially unfamiliar, degu pups in a neutral arena. No male or female degu showed any sign of aggression toward the pups. Lactating females tended to exhibit the shortest latency to first behavioral interaction with the pup and the highest rate of social interactions with the pup, and they spent a relatively high proportion of their time in proximity with the pup. In contrast, males tended to show the longest latency to first pup contact and a reduced rate of interactions with the pup, and they spent a relatively small fraction of their time with the pup. The behavior of non-breeding females seemed intermediate between that of males and lactating females. Given that social and ecological conditions posed to promote non-parental infanticide in other rodents seemed not particularly different from what is known of degu biology, ecology, and social behavior, lack of degu infanticide may reflect phylogenetic inertia instead of an absence of conditions favoring infanticide. Received: 8 January 2000 / Received in revised form: 16 June 2000 / Accepted: 20 June 2000     

Seasonal variation in the range areas of the diurnal rodent Octodon degus

Both breeding activity and abundance and quality of available food are expected to influence daily movements of animals. Animals are predicted to range over large areas to meet high energy demands associated with reproduction (females) or to increase mating success (males). However, animals should expand their range areas whenever food conditions deteriorate. To examine the extent to which breeding activity versus food availability influence space use, we compared the size and location of range areas (home ranges) of the degu (Octodon degus), a diurnal rodent from semiarid environments of north-central Chile, during the austral winter and summer seasons. Degus produce young during the austral spring (September-October) when high-quality food is readily available. In contrast, degus do not breed during the austral summer (January-March) when food is scarce and of low quality. We predicted that degus would range over smaller areas in winter if the availability of food has a greater influence on space than breeding activity. Individuals were radiotracked in winter and the following summer over a 3-year period. Surveys of herbaceous cover were conducted during winter and summer to determine seasonal changes in the abundance and quality of primary food. In summer degus expanded and moved the location of their range areas to locations with available food. Given that preferred food was less abundant in summer than winter, we suggest that degu range areas are strongly influenced by food conditions.     

Gonadal hormone effects on entrained and free-running circadian activity rhythms in the developing diurnal rodent Octodon degus

The slowly maturing, long-lived rodent Octodon degus (degu) provides a unique opportunity to examine the development of the circadian system during adolescence. These studies characterize entrained and free-running activity rhythms in gonadally intact and prepubertally gonadectomized male and female degus across the first year of life to clarify the impact of sex and gonadal hormones on the circadian system during adolescence. Gonadally intact degus exhibited a delay in the phase angle of activity onset (Psi(on)) during puberty, which reversed as animals became reproductively competent. Gonadectomy before puberty prevented this phase delay. However, the effect of gonadal hormones during puberty on Psi(on) does not result from changes in the period of the underlying circadian pacemaker. A sex difference in Psi(on) and free-running period (tau) emerged several months after puberty; these developmental changes are not likely to be related, since the sex difference in Psi(on) emerged before the sex difference in tau. Changes in the levels of circulating hormones cannot explain the emergence of these sex differences, since there is a rather lengthy delay between the age at which degus reach sexual maturity and the age at which Psi(on) and tau become sexually dimorphic. However, postnatal exposure to gonadal hormones is required for sexual differentiation of Psi(on) and tau, since these sex differences were absent in prepubertally gonadectomized degus. These data suggest that gonadal hormones modulate the circadian system during adolescent development and provide a new model for postpubertal sexual differentiation of a central nervous system structure.     

Testosterone suppresses circadian responsiveness to social cues in the diurnal rodent Octodon degus

The diurnal, social rodent Octodon degus displays a robust sex difference in the ability to use social cues to facilitate reentrainment following a phase advance of the light cycle. Adult females housed with a female social cue donor reentrained 25% to 40% faster than did females reentraining alone. However, reentrainment rates of males were unaffected by exposure to female social cues during reentrainment. The authors hypothesized that males were less sensitive to the reentrainment-enhancing effects of social cues and that their higher threshold to the stimuli could be overcome if the social cues were either increased in strength or salience. Housing a male with two females significantly shortened the time to reentrain following a 9-h phase advance (p = 0.002). Housing with a sister had no effect on reentrainment. Therefore, male degus are able to respond to social cues but require the stimulus to be stronger than that for females. The effect of testosterone was tested by comparing reentrainment rates of castrated males before and after testosterone replacement both with and without a female social cue donor. Castrated males responded to a single female social cue donor, reentraining 35% faster than when housed alone (p = 0.006), whereas the time to reentrainment of intact males and males with testosterone capsule implants did not differ. Intact females were also implanted with testosterone and phase shifted with and without donors. Testosterone treatment eliminated the increase in reentrainment rates in the presence of social cues. The authors conclude that the rate of recovery from odor-enhanced phase shifts is modulated by activational effects of testosterone in male degus. Testosterone is also effective in suppressing social cue responsiveness in females, suggesting that testosterone's effects on responsiveness are not sexually dimorphic. This hormonal effect likely occurs by altering sensory system functions or CNS response to sensory information.     

Social cues and hormone levels in male Octodon degus (Rodentia): a field test of the Challenge Hypothesis

Social interactions are important factors determining and regulating individual behaviors. Testosterone has been related to agonistic interactions, while glucocorticoids have been related to social stress, especially during interactions of dominance. We compared testosterone and cortisol concentrations in male degus (Octodon degus, Rodentia) under laboratory conditions without male social interactions, with data from wild males in nature. Under natural conditions, males should present higher levels of testosterone during the breeding season due to social interactions (Challenge Hypothesis). Alternatively, intense social instability could act as a stressing environment, raising glucocorticoids, which inhibit testosterone concentrations. Our results show a significant increase in agonistic interactions between males during the breeding season, and disappearance of non-agonistic male interactions during this period. Hormone levels in breeding season show nonsignificant differences between laboratory groups, but testosterone concentrations in field males were significantly higher than in laboratory males. Testosterone levels were similar among pre-breeding and breeding periods, but in field animals the concentration was approximately 30% higher than in laboratory degus. In field animals, we found two different mating strategies: resident males, with territorial behavior, and transient males, displayed an opportunistic approach to females. Finally, cortisol presents a similar pattern in both laboratory and field animals; pre-breeding values of cortisol are higher than during the breeding season. This suggests that social interactions in O. degus activate a rise in testosterone, supporting the Challenge Hypothesis, and could be considered as partial support of the Social Stress Hypothesis.     

Burrow limitations and group living in the communally rearing rodent, Octodon degus

Group living is thought to evolve whenever individuals attain a net fitness advantage due to reduced predation risk or enhanced foraging efficiency, but also when individuals are forced to remain in groups, which often occurs during high-density conditions due to limitations of critical resources for independent breeding. The influence of ecological limitations on sociality has been studied little in species in which reproduction is more evenly shared among group members. Previous studies in the caviomorph rodent Octodon degus (a New World hystricognath) revealed no evidence that group living confers an advantage and suggest that burrow limitations influence formation of social groups. Our objective was to examine the relevance of ecological limitations on sociality in these rodents. Our 4-year study revealed no association between degu density and use of burrow systems. The frequency with which burrow systems were used by degus was not related to the quality of these structures; only in 1 of the 4 years did the frequency of burrow use decrease with decreasing abundance of food. Neither the number of females per group nor total group size (related measures of degu sociality) changed with yearly density of degus. Although the number of males within social groups was lower in 2008, this variation was not related clearly to varying density. The percentage of females in social groups that bred was close to 99% and did not change across years of varying density. Our results suggest that sociality in degus is not the consequence of burrow limitations during breeding. Whether habitat limitations contribute to variation in vertebrate social systems is discussed.     

Sociality, glucocorticoids and direct fitness in the communally rearing rodent, Octodon degus

While ecological causes of sociality (or group living) have been identified, proximate mechanisms remain less clear. Recently, close connections between sociality, glucocorticoid hormones (cort) and fitness have been hypothesized. In particular, cort levels would reflect a balance between fitness benefits and costs of group living, and therefore baseline cort levels would vary with sociality in a way opposite to the covariation between sociality and fitness. However, since reproductive effort may become a major determinant of stress responses (i.e., the cort–adaptation hypothesis), cort levels might also be expected to vary with sociality in a way similar to the covariation between sociality and fitness. We tested these expectations during three years in a natural population of the communally rearing degu, Octodon degus. During each year we quantified group membership, measured fecal cortisol metabolites (a proxy of baseline cort levels under natural conditions), and estimated direct fitness. We recorded that direct fitness decreases with group size in these animals. Secondly, neither group size nor the number of females (two proxies of sociality) influenced mean (or coefficient of variation, CV) baseline cortisol levels of adult females. In contrast, cortisol increased with per capita number of offspring produced and offspring surviving to breeding age during two out of three years examined. Together, our results imply that variation in glucocorticoid hormones is more linked to reproductive challenge than to the costs of group living. Most generally, our study provided independent support to the cort–adaptation hypothesis, according to which reproductive effort is a major determinant, yet temporally variable, influence on cort–fitness covariation.     

Distribution of aquaporins in the colon of Octodon degus,a South American desert rodent

Octodon degus is a desert rodent of northern Chile, adapted to survive with a limited supply of water. This rodent has a high degree of fecal dehydration, related to colon water absorption. With the hypothesis that aquaporins (AQPs) might be present in the colon epithelium of O. degus and involved in fluid absorption, we studied colon water absorption in vivo and the distribution of AQPs and Na(+) transporters by immunocytochemistry. AQP-1 was found in apical and basolateral membranes of surface-absorptive and crypt epithelial cells. AQP-8 was found in the cytoplasm of enterocytes of surface colon. AQP-3 immunolabeling, on the other hand, was absent from the epithelium but present in a subepithelial fibroblast layer, pericryptal cells, and muscularis mucosae. The hydration state did not modify the amount of immunostaining for any of the AQPs. Colon water absorption was markedly decreased by the mercurial agent p-chloromercuribenzenesulfonic acid and was not affected by water deprivation. The NHE3 isoform of Na(+)/H(+) exchanger and alpha-1 subunit of the Na(+)-K(+)-ATPase were found in apical and basolateral membranes of surface-absorptive cells, respectively. These results suggest that colon water absorption is mostly transcellular and mediated by water channels like AQP-1. Apical Na(+)/H(+) exchanger and basolateral Na(+)-K(+)-ATPase in surface cells could be part of the Na(+) absorption pathway. It is hypothesized that this transport is necessary to provide an osmotic gradient for water absorption. The roles of AQP-8 and AQP-3 in water absorption remain to be established.     

Preparation of rhinovirus antisera in the South American hystricomorph rodent, Octodon degus (38507).

Rhinovirus antisera have been prepared for rhinoviruses (RV) 7,9,26,32,67 and 87 in guinea pigs and in degus. Titers achieved were either similar in the 2 animals (RV7) somewhat higher in the degu (RV9 and RV32) or clearly higher in the degu (RV26, RV67 and RV87). Specificity of the antisera was similar in both animals. In special instances where it is difficult to prepare high-titered rhinovirus antisera in the guinea pig, the degu offers an attractive alternative source.     

Gonadal hormones provide the biological basis for sex differences in behavioral responses to cocaine

Both clinical and rodent studies show sexually dimorphic patterns in the behavioral response to cocaine in all phases of the addiction process (induction, maintenance, and relapse). Clinical and rodent studies also indicate that hormonal fluctuations during the menstrual/estrous cycle modulate cocaine-induced subjective effects in women and locomotor activity in female rats. Evidence suggests that gonadal hormones underlie these observed differences and could be the biological basis of sex-specific differences in cocaine addiction. To study the effects of gonadal hormones on cocaine-induced activity, two approaches have been used. First, studies have examined the role of endogenous hormones through gonadectomy (GDX) and side-by-side comparisons with intact rats. Second, the individual contributions of testosterone, progesterone, and estrogen have been determined by hormone replacement in GDX rats. In this review, we discuss gonadal hormones as the biological basis for the behavioral responses to cocaine, and the clinical implications of these findings.     

A daily rhythm in mating behavior in a diurnal murid rodent Arvicanthis niloticus

The time of day at which mating occurs is dramatically different in diurnal compared to nocturnal rodents. We used a diurnal murid rodent, Arvicanthis niloticus, to determine if inverted rhythms in responsiveness to hormones contribute to this difference. Male and hormone-primed female grass rats were tested for mating behavior at four different times of day (ZT 5, 11, 17, 23; ZT 0=lights-on). In females, there was considerable inter-individual variability with respect to patterns of responsiveness to hormones. Overall, the lordosis quotient (LQ) was rhythmic with a single peak just before lights-on (ZT 23); however, while roughly half of the females (7/15) exhibited this clear daily rhythm, the remaining animals (8/15) had relatively high LQs that did not change as a function of time. Males had their shortest ejaculation latencies and their highest number of ejaculations at ZT 23. Rhythms in mount frequency and post-ejaculatory refractory period were bimodal, with peaks around lights-on and -off (ZT 23 and 11). This temporal pattern of mounting behavior closely parallels previously documented patterns of general activity, whereas rhythms in the more reflexive components of sex behavior (LQ and ejaculation) had more restricted peaks that coincided with just the onset of rhythms in general activity. These rhythms in sexual behavior are essentially reversed relative to those previously documented in lab rats.     

Serum concentrations of reproductively-related circulating steroid hormones in the free-ranging lemon shark,Negaprion brevirostris

Synopsis We have examined the concentrations of reproductively-related steroid hormones in 5 species of carcharhinid sharks, marine fishes possessing the unique attribute of placental viviparity. Measurements of serum estradiol, testosterone, progesterone, dihydrotestosterone, and corticosterone have provided baseline data for these hormones in both immature and adult male and female placental sharks. Our studies include: (1) changes in hormonal levels during maturation, (2) concentrations of circulating steroid hormones during peak breeding season, and (3) hormonal levels during gestation, including the collection of serial samples through and beyond birth from free-ranging lemon sharks. Our data suggest that steroids, important in regulating reproduction in higher mammals, are also essential in these cartilaginous fishes.     

Effects of vertebrate predation on a caviomorph rodent,the degu (Octodon degus), in a semiarid thorn scrub community in Chile

The effects of vertebrate predation have been monitored since 1989 on 16 replicated 0.56 ha study plots in a semiarid thorn scrub community in north-central Chile. Using fences of different heights with and without holes and suspended game netting to alter principal predator (foxes and raptors) and large rodent herbivore (Octodon degus) access, four grids each have been assigned to the following treatments: 1) low fencing and holes allowing free access of predators and small mammals; 2) low fencing without holes to exclude degus only; 3) high fencing and netting with holes to exclude predators only; and 4) high fencing and netting without holes to exclude predators and degus. Small mammal population censuses are conducted monthly using mark-recapture techniques. Degu population trends during 1989 and 1990 showed strongly but nonsignificantly lower numbers in control plots during months when densities were characteristically low (September–November) for this seasonally reproductive species; since March 1991, differences have become persistent and increasingly significant. Predators appear to have greater numerical effects when their prey populations are low. Survival times of degus, particularly established adults, were significantly longer in predator exclusion grids during the 2 1/2 years of observation; thus, predation also affects prey population structure.