Nocturnal activity in a diurnal rodent (Arvicanthis niloticus): the importance of masking

It is known that day-active Nile grass rats, Arvicanthis niloticus, increase the amount of activity in the night relative to that in the day when provided with running wheels. This was confirmed in the present study. Animals without a wheel displayed 69.0% of their general activity in the L phase of a 12:12 h light-dark cycle; animals provided with wheels had only 48.6% of their wheel revolutions in the light. The contribution of direct (masking) responses to light to the increased nocturnality of animals with wheels was examined in two experiments. In experiment 1, masking was tested by exposing the animals to repeated cycles of 30 min of entraining light and 30 min of a different, usually dimmer light, during the L phase of a 12:12 h light-dark cycle. For animals with wheels, there was more running during the 30-min pulses of dim light or darkness than during the 30-min periods of entraining light. In contrast, for animals without wheels, there was more general activity during the 30-min periods of entraining light than during the 30-min pulses of dim light or darkness. In experiment 2, the animals were first exposed to a 12:12 h light-dark cycle and then put on a 1:10:1:12 h LDLD skeleton photoperiod. Animals with wheels increased their running during the subjective day of the skeleton photoperiod compared to that in the actual day of the 12:12 h light-dark cycle. Animals without wheels showed similar levels of general activity during the subjective day of the skeleton photoperiod and the actual day of the 12:12 h cycle. These experiments demonstrate that when Nile rats have running wheels, their increased nocturnal activity is associated with an increased suppression of locomotion in direct response to light. It is possible that changes in masking responses to light may be an essential and integral component of switching between diurnal and nocturnal activity profiles.     

Long-term infection with Schistosoma mansoni (Gezira strain--Sudan) in the Nile rat (Arvicanthis niloticus)

The long-term infection with Schistosoma mansoni (Gezira strain--Sudan) was studied in the Nile rat (Arvicanthis niloticus) to investigate the 'self cure' phenomenon. The results indicated that while a considerable number of worms and eggs were still recovered by week 28 post-infection, elimination of some of the worms occurred by week 20.     

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.     

Acute Behavioral Responses to Light and Darkness in Nocturnal Mus musculus and Diurnal Arvicanthis niloticus

The term masking refers to immediate responses to stimuli that override the influence of the circadian timekeeping system on behavior and physiology. Masking by light and darkness plays an important role in shaping an organism's daily pattern of activity. Nocturnal animals generally become more active in response to darkness (positive masking) and less active in response to light (negative masking), and diurnal animals generally have opposite patterns of response. These responses can vary as a function of light intensity as well as time of day. Few studies have directly compared masking in diurnal and nocturnal species, and none have compared rhythms in masking behavior of diurnal and nocturnal species. Here, we assessed masking in nocturnal mice (Mus musculus) and diurnal grass rats (Arvicanthis niloticus). In the first experiment, animals were housed in a 12:12 light-dark (LD) cycle, with dark or light pulses presented at 6 Zeitgeber times (ZTs; with ZT0 = lights on). Light pulses during the dark phase produced negative masking in nocturnal mice but only at ZT14, whereas light pulses resulted in positive masking in diurnal grass rats across the dark phase. In both species, dark pulses had no effect on behavior. In the 2nd experiment, animals were kept in constant darkness or constant light and were presented with light or dark pulses, respectively, at 6 circadian times (CTs). CT0 corresponded to ZT0 of the preceding LD cycle. Rhythms in masking responses to light differed between species; responses were evident at all CTs in grass rats but only at CT14 in mice. Responses to darkness were observed only in mice, in which there was a significant increase in activity at CT 22. In the 3rd experiment, animals were kept on a 3.5:3.5-h LD cycle. Surprisingly, masking was evident only in grass rats. In mice, levels of activity during the light and dark phases of the 7-h cycle did not differ, even though the same animals had responded to discrete photic stimuli in the first 2 experiments. The results of the 3 experiments are discussed in terms of their methodological implications and for the insight they offer into the mechanisms and evolution of diurnality.     

Rhythms in Fos expression in brain areas related to the sleep-wake cycle in the diurnal Arvicanthis niloticus

Most mammals show daily rhythms in sleep and wakefulness controlled by the primary circadian pacemaker, the suprachiasmatic nucleus (SCN). Regardless of whether a species is diurnal or nocturnal, neural activity in the SCN and expression of the immediate-early gene product Fos increases during the light phase of the cycle. This study investigated daily patterns of Fos expression in brain areas outside the SCN in the diurnal rodent Arvicanthis niloticus. We specifically focused on regions related to sleep and arousal in animals kept on a 12:12-h light-dark cycle and killed at 1 and 5 h after both lights-on and lights-off. The ventrolateral preoptic area (VLPO), which contained cells immunopositive for galanin, showed a rhythm in Fos expression with a peak at zeitgeber time (ZT) 17 (with lights-on at ZT 0). Fos expression in the paraventricular thalamic nucleus (PVT) increased during the morning (ZT 1) but not the evening activity peak of these animals. No rhythm in Fos expression was found in the centromedial thalamic nucleus (CMT), but Fos expression in the CMT and PVT was positively correlated. A rhythm in Fos expression in the ventral tuberomammillary nucleus (VTM) was 180 degrees out of phase with the rhythm in the VLPO. Furthermore, Fos production in histamine-immunoreactive neurons of the VTM cells increased at the light-dark transitions when A. niloticus show peaks of activity. The difference in the timing of the sleep-wake cycle in diurnal and nocturnal mammals may be due to changes in the daily pattern of activity in brain regions important in sleep and wakefulness such as the VLPO and the VTM.     

Female prostate in Arvicanthis niloticus and Meriones libycus.

A female prostate is reported in Arvicanthis niloticus (field rat) and Meriones libycus for the first time, as a normal, constant and well developed bilateral sex organ.     

A morning surge in plasma luteinizing hormone coincides with elevated Fos expression in gonadotropin-releasing hormone-immunoreactive neurons in the diurnal rodent, Arvicanthis niloticus.

Arvicanthis niloticus is a diurnal murid rodent from sub-Saharan Africa. Here we report on processes associated with mating in this species in an attempt to elucidate how the neural mechanisms governing temporal organization differ in nocturnal and diurnal species. First, we systematically mapped the distribution of GnRH neurons in adult females. Second, we tested the hypothesis that Arvicanthis differ from nocturnal murid rodents with respect to the timing of the LH surge and the associated increase in Fos expression in GnRH-immunoreactive (IR) neurons. We examined these events around a postpartum estrus. When parturition occurred between zeitgeber time (ZT) 2 and 17 (lights on at ZT 0 and off at ZT 12; there are 24 ZT units a day, each equivalent to 1 standard hour), we collected blood and perfused females at ZT 17, 20, 23, or 2. A sharp peak in plasma LH occurred at ZT 20, and a 10-fold increase in the percentage of GnRH-IR neurons that expressed Fos-IR occurred between ZT 17 and 20. By contrast, this rise occurs in nocturnal rodents during the last few hours of the light period. This is the first indication of a difference between nocturnal and diurnal animals with respect to neural mechanisms associated with a precisely timed event of known significance.     

Thermoregulation and diurnal rhythms in 1-week-old rat pups

This paper reviews the ontogeny of thermoregulation and diurnal rhythmicity in rats. Additionally, original data are presented that indicate the emergence of an endogenous circadian core temperature rhythm during the first postnatal week. Despite neurological immaturity, newborn rats display autonomic and behavioral thermoregulatory responses within 24 h of birth. Their "biological clock" is already running before birth. The thermal environment of pups changes cyclically owing to diurnal variations in maternal behavior, but the core temperatures of 1-week-old pups huddling in the absence of the dam also show marked diurnal fluctuations. Five- to 8-day-old lean Zucker rat pups artificially reared in the absence of 24-h cycles of ambient temperature and food intake show diurnal changes in core temperature similar to those in huddling mother-reared pups. Diurnal core temperature changes, evident only when regulatory effectors are not overwhelmed, are one of the first self-maintained diurnal rhythms to appear. Because thermoregulation and circadian rhythmicity both appear before maturation of the neural networks believed to be critical for their control in adult animals, studying the immature rat might increase our understanding of the control of these processes in the more complex mature central nervous system.     

Clock-controlled endogenous melatonin rhythms in Nile tilapia (Oreochromis niloticus niloticus) and African catfish (Clarias gariepinus)

The purpose of this work was to investigate the circadian melatonin system in two tropical teleost species characterized by different behavioral habits, Nile tilapia (diurnal) and African catfish (nocturnal). To do so, fish were subjected to either a control photoperiod (12L:12D), continuous light (LL) or darkness (DD), or a 6L:6D photoperiod. Under 12L:12D, plasma melatonin levels were typically low during the photophase and high during the scotophase in both species. Interestingly, in both species, melatonin levels significantly decreased prior to the onset of light, which in catfish reached similar basal levels to those during the day, demonstrating that melatonin production can anticipate photic changes probably through circadian clocks. Further evidence for the existence of such pacemaker activity was obtained when fish were exposed to DD, as a strong circadian melatonin rhythm was maintained. Such an endogenous rhythm was sustained for at least 18 days in Nile tilapia. A similar rhythm was shown in catfish, although DD was only tested for four days. Under LL, the results confirmed the inhibitory effect of light on melatonin synthesis already reported in other species. Finally, when acclimatized to a short photo-cycle (6L:6D), no endogenous melatonin rhythm was observed in tilapia under DD, with melatonin levels remaining high. This could suggest that the circadian clocks cannot entrain to such a short photocycle. Additional research is clearly needed to further characterize the circadian axis in teleost species, identify and localize the circadian clocks, and better understand the environmental entrainment of fish physiology.     

Circadian regulation of gonadotropin-releasing hormone neurons and the preovulatory surge in luteinizing hormone in the diurnal rodent, Arvicanthis niloticus, and in a nocturnal rodent, Rattus norvegicus

Daily rhythms in the timing of the preovulatory surge and the display of reproductive behavior are reversed in diurnal and nocturnal rodents, but little is known about the neural mechanisms underlying these differences. We examined this issue by comparing a diurnal murid rodent, Arvicanthis niloticus (the grass rat), to a nocturnal one, Rattus norvegicus (the lab rat). In the first study, we established that sequential estradiol and progesterone treatment induces a proestrous-like rise in LH secretion and in the percentage of GnRH neurons that express Fos in grass rats, as is the case in lab rats. Next, we tested the hypothesis that differences in the timing of estrus-related events in diurnal and nocturnal species are caused by differences in rhythms in responsiveness to steroid hormones. We found rhythms in GnRH neuron activity, as indicated by Fos, that were 12 hours out of phase in grass rats and lab rats. These patterns persisted in both species when animals were housed in constant darkness for 5 days, suggesting that they are driven by an endogenous circadian mechanism. These results indicate that steroid-primed grass rats and lab rats are similar with respect to the temporal relationship among estrus-related events, but that the timing of these events relative to the light-dark cycle is dramatically different and that this difference is caused by endogenous circadian mechanisms.     

Effects of photoperiod on the reproductive condition of Nile grass rats (Arvicanthis niloticus) from an equatorial population

We evaluated the effects of photoperiod on the reproductive condition of male and female Nile grass rats (Arvicanthis niloticus) descended from members of an equatorial population trapped 2°S of the equator. Study animals housed in 12:12 light:dark (LD) cycles were transferred either to short photoperiod (9:15) or long photoperiod (15:9) for 9 weeks (males) or 11 weeks (females), and various reproductive parameters were assessed. We observed no differences between short‐ and long‐day males with respect to plasma concentration of testosterone, testicular mass, seminal vesicle mass, or spermatogenesis. Similarly, we observed no differences between short‐ and long‐day females with respect to oestrous cycles, uterine mass, follicle size, or presence of corpora lutea. Reproductive parameters of male and female A. niloticus housed in short‐ and long photoperiods were similar to those typically observed among animals descended from the same equatorial population and housed in LD 12:12. Thus, photoperiod appears not to elicit changes in reproductive condition among A. niloticus from populations whose native habitat lies within 2° of the equator. These data contrast with the results of other studies indicating that photoperiod alters reproductive condition in A. niloticus populations living >10° from the equator.     

Daily rhythms after vaccination on specific and non-specific responses in Nile tilapia (Oreochromis niloticus)

ABSTRACT

We evaluated the daily changes in immunological and hematological factors in tilapia (Oreochromis niloticus) after an immunization period with a subsequent challenge. Experiments were divided into two phases: Phase 1 (immunization): 144 fish were distributed into two groups with 72 fish in six tanks. One group (T1) was immunized, comprising six vaccination time points (ZT schedule = ZT2 h, ZT6 h, ZT10 h, ZT14 h, ZT18 h, ZT22 h). The same schedule was applied to the other group, but with saline solution (non-vaccinated: T2). Both groups remained in the laboratory for 30 days (considered the immunization period). Phase 2 (challenge): on day 30, both vaccinated and non-vaccinated groups were challenged with Streptococcus agalactiae (2.0 × 10⁷ CFU mL^?1) following the same ZT schedule to stimulate the immune response without leading to widespread infection and mortality. On day 45, blood and head kidney samples were collected during the same ZT schedule. The variations in time of the following parameters within each group were evaluated: hematology, peroxidase activity, IgM, tnf-α3, tgf-β1, il-1β and il-12 gene expression. No significant mortality was observed for the groups or the ZT schedule (p > 0.05). Daily rhythms with diurnal acrophases were found in T2 for il12, tnf-α3 and tgf-β1 expression gene, while the acrophases of the peroxidase level, hematocrit and thrombocytes were at nighttime (p < 0.05). In contrast, most of the parameters in the vaccinated tilapia showed no daily rhythms (p > 0.05), except IgM. For all the parameters, the interaction effect between time and treatment (vaccinated and non-vaccinated groups) depended on ZT. Our results reveal that the humoral and non-specific immune system displayed a circadian rhythm based on the light-dark cycle, which could be affected by the vaccination procedure in tilapia.     

Nitrite-induced methemoglobinemia in Nile tilapia, Oreochromis niloticus

Exposure of Nile tilapia, Oreochromis niloticus (mean weight, 55.72 ± 4.30 g), to two sublethal NO₂–N concentrations was studied for 24 and 48 h in a static test. In nitrite exposure tests, the percentages of methemoglobin, external nitrite, plasma nitrite, hemoglobin and hematocrit were assessed. Nitrite exposure in the range of 0.50 and 1.38 mg l⁻¹ NO₂–N caused an increase in methemoglobin levels; however, methemoglobin percentages ranging from 16% to 42% represented a mild methemoglobinemia. Levels of methemoglobin were unrelated to environmental and plasmatic nitrite concentrations. The nitrite concentration in the blood did not seem to be linked to time of exposure. Nitrite exposure in Nile tilapia was associated with a marked reduction in hemoglobin and hematocrit.     

Effects of climate and local aridity on the latitudinal and habitat distribution of Arvicanthis niloticus and Arvicanthis ansorgei (Rodentia, Murinae) in Mali

Introduction The genus Arvicanthis (Lesson 1842) (Rodentia: Murinae), usually referred to as the unstriped grass rat, is mainly distributed in savanna and grassland habitats of Sub-Saharan Africa. Among the four chromosomal forms of Arvicanthis recently differentiated in Western and Central Africa, the one with a diploid chromosomal number (2n) of 62 and an autosomal fundamental number (NFa) of 62 or 64 is ascribed to Arvicanthis niloticus (Demarest 1822), while the one with 2n = 62 and a NFa between 74 and 76 is referred to A. ansorgei (Thomas 1910). Despite the broad area of sympatry recently uncovered along the inner delta of the Niger river in Mali [details in Volobouev et al. (2002) Cytogenetics and Genome Research, 96, 250–260], the distribution of the two species is largely parapatric and follows the latitudinal patterns of the West-African biogeographical domains, which are related to the latitudinal patterns of annual rainfall in this region. Here, we analyse the suggestion that the two species show specific adaptations to differences in climate aridity. Methods Karyologically screened animals were sampled in 19 localities in seasonally flooded regions located along the ‘Niger’ river in Mali and extending from 1100 to 200 mm of mean annual rainfall. The analysis of trapping success (TS) data allowed us to investigate the respective effects of climate (i.e. annual rainfall) and local (i.e. duration of the green herbaceous vegetation) aridity on the latitudinal and habitat distribution of the two species. Conclusions The broad zone of sympatry was found to correspond to a northward expansion of the recognized distribution area of A. ansorgei. TS values indicated that the two species responded very differently to climatic and local conditions of aridity. Arvicanthis ansorgei decreased in TS as regional conditions became more arid; a similar trend was also observed within regions where habitat occupancy decreased with local aridity. The higher TS observed in the most humid habitat relative to the others persisted throughout the latitudinal rainfall gradient. In contrast, TS of A. niloticus increased with latitudinal aridity. This species was present in more arid habitats than A. ansorgei from 1000 mm down to 400 mm of mean annual rainfall where a shift to the most humid habitat occurred. These opposite trends in TS distribution between species suggest that A. ansorgei is less adapted than A. niloticus to arid environments at both a regional and habitat level; thus, A. ansorgei would be able to invade dry regions only along the extensive floodplains bordering the inner delta of the ‘Niger’ river. Several biological traits that may be involved in limiting the southward distribution of A. niloticus are discussed.