Orexin (hypocretin) mediates light-dependent fluctuation of hippocampal function in a diurnal rodent.

Environmental lighting conditions play a central role in cognitive function, but the underlying mechanisms remain unclear. Utilizing a diurnal rodent model, the Nile grass rat (Arvicanthis niloticus), we previously found that daytime light intensity affects hippocampal function in this species in a manner similar to its effects in humans. Compared to animals housed in a 12:12 h bright light-dark (brLD) cycle, grass rats kept in a 12:12 h dim light-dark (dimLD) cycle showed impaired spatial memory in the Morris water maze (MWM) and reduced CA1 apical dendritic spine density.

Daytime Light Intensity Modulates Spatial Learning and Hippocampal Plasticity in Female Nile Grass Rats (Arvicanthis niloticus).

Light has pervasive effects on the physiology and behavior of mammals. Several human studies have shown that light modulates cognitive functions; however, the mechanisms responsible for the effects of light remain unclear. Our previous work using diurnal male Nile grass rats (Arvicanthis niloticus) revealed that reduced illuminance during the day leads to impairments in hippocampal-dependent spatial learning/memory, reduced CA1 dendritic spine density, and attenuated hippocampal brain-derived neurotrophic factor (BDNF) expression in males.

Circadian and photic modulation of daily rhythms in diurnal mammals.

The temporal niche that an animal occupies includes a coordinated suite of behavioral and physiological processes that set diurnal and nocturnal animals apart. The daily rhythms of the two chronotypes are regulated by both the circadian system and direct responses to light, a process called masking. Here we review the literature on circadian regulations and masking responses in diurnal mammals, focusing on our work using the diurnal Nile grass rat (Arvicanthis niloticus) and comparing our findings with those derived from other diurnal and nocturnal models.

Light as a modulator of emotion and cognition: Lessons learned from studying a diurnal rodent.

Light profoundly affects the behavior and physiology of almost all animals, including humans. One such effect in humans is that the level of illumination during the day positively contributes to affective well-being and cognitive function. However, the neural mechanisms underlying the effects of daytime light intensity on affect and cognition are poorly understood.

Distributions of GABAergic and glutamatergic neurons in the brains of a diurnal and nocturnal rodent.

Light influences the daily patterning of activity by both synchronizing internal clocks to environmental light-dark cycles and acutely modulating arousal states, a process known as masking. Masking responses are completely reversed in diurnal and nocturnal species. In nocturnal rodents, masking is mediated through a subset of intrinsically photosensitive retinal ganglion cells (ipRGCs) whose projections are similar in diurnal and nocturnal rodents.

The Cost of Activity during the Rest Phase: Animal Models and Theoretical Perspectives.

For humans, activity during the night is correlated with multiple pathologies that may reflect a lack of harmony among components of the circadian system; however, it remains difficult to identify causal links between nocturnal activity and different pathologies based on the data available from epidemiological studies. Animal models that use forced activity or timed sleep deprivation provide evidence of circadian disruptions that may be at the core of the health risks faced by human night and shift workers. One valuable insight from that work is the importance of changes in the distribution of food intake as a cause of metabolic imbalances associated with activity during the natural rest phase.

Light modulates hippocampal function and spatial learning in a diurnal rodent species: A study using male nile grass rat (Arvicanthis niloticus).

The effects of light on cognitive function have been well-documented in human studies, with brighter illumination improving cognitive performance in school children, healthy adults, and patients in early stages of dementia. However, the underlying neural mechanisms are not well understood. The present study examined how ambient light affects hippocampal function using the diurnal Nile grass rats (Arvicanthis niloticus) as the animal model.

Normal behavioral responses to light and darkness and the pupillary light reflex are dependent upon the olivary pretectal nucleus in the diurnal Nile grass rat.

The olivary pretectal nucleus (OPT) is a midbrain structure that receives reciprocal bilateral retinal projections, is involved in the pupillary light reflex, and connects reciprocally with the intergeniculate leaflet (IGL), a retinorecipient brain region that mediates behavioral responses to light pulses (i.e., masking) in diurnal Nile grass rats.

Suprachiasmatic Nucleus and Subparaventricular Zone Lesions Disrupt Circadian Rhythmicity but Not Light-Induced Masking Behavior in Nile Grass Rats.

The ventral subparaventricular zone (vSPVZ) receives direct retinal input and influences the daily patterning of activity in rodents, making it a likely candidate for the mediation of acute behavioral responses to light (i.e., masking).

Acute effects of light on the brain and behavior of diurnal Arvicanthis niloticus and nocturnal Mus musculus.

Photic cues influence daily patterns of activity via two complementary mechanisms: (1) entraining the internal circadian clock and (2) directly increasing or decreasing activity, a phenomenon referred to as "masking". The direction of this masking response is dependent on the temporal niche an organism occupies, as nocturnal animals often decrease activity when exposed to light, while the opposite response is more likely to be seen in diurnal animals. Little is known about the neural mechanisms underlying these differences.

Circadian modulation of memory and plasticity gene products in a diurnal species.

Cognition is modulated by circadian rhythms, in both nocturnal and diurnal species. Rhythms of clock gene expression occur in brain regions that are outside the master circadian oscillator of the suprachiasmatic nucleus and that control cognitive functions, perhaps by regulating the expression neural-plasticity genes such as brain derived neurotrophic factor (BDNF) and its high affinity receptor, tyrosine kinase B (TrkB). In the diurnal grass rat (Arvicanthis niloticus), the hippocampus shows rhythms of clock genes that are 180° out of phase with those of nocturnal rodents.

Intergeniculate leaflet lesions result in differential activation of brain regions following the presentation of photic stimuli in Nile grass rats.

The intergeniculate leaflet (IGL) plays an important role in the entrainment of circadian rhythms and the mediation of acute behavioral responses to light (i.e., masking).

Behavioral Masking and cFos Responses to Light in Day- and Night-Active Grass Rats.

Light not only entrains the circadian system but also has acute effects on physiology and behavior, a phenomenon known as masking. Behavioral masking responses to bright light differ in diurnal and nocturnal species, such that light increases arousal in the former and decreases it in the latter. Comparisons made within a species that displays both diurnal and nocturnal patterns of behavior may provide insight into how masking differs between chronotypes and the association between mechanisms controlling masking and the circadian drive for activity.

Lesions of the Intergeniculate Leaflet Lead to a Reorganization in Circadian Regulation and a Reversal in Masking Responses to Photic Stimuli in the Nile Grass Rat.

Light influences the daily patterning of behavior by entraining circadian rhythms and through its acute effects on activity levels (masking). Mechanisms of entrainment are quite similar across species, but masking can be very different. Specifically, in diurnal species, light generally increases locomotor activity (positive masking), and in nocturnal ones, it generally suppresses it (negative masking).

Pregnancy affects FOS rhythms in brain regions regulating sleep/wake state and body temperature in rats.

Circadian rhythms in behavior and physiology change substantially as female mammals undergo the transition from a non-pregnant to a pregnant state. Here, we examined the possibility that site-specific changes in brain regions known to regulate the sleep/wake cycle and body temperature might reflect altered rhythms in these overt functions. Specifically, we compared daily patterns of immunoreactive FOS in early pregnant and diestrous rats in the medial septum (MS), vertical and horizontal diagonal bands of Broca (VDB and HDB), perifornical lateral hypothalamus (LH), and ventrolateral, medial, and median preoptic areas (VLPO, MPA, and MnPO, respectively).

Site-specific changes in brain extra-SCN oscillators during early pregnancy in the rat.

The influence of circadian timekeeping systems on behavior and physiology can change substantially as female mammals undergo the transition from a nonpregnant to a pregnant state. Here, we examined the possibility that site-specific changes in extra-SCN oscillators and in local rhythms might coincide with the emergence of new patterns of temporal organization among various behavioral and physiological rhythms. Specifically, we compared daily patterns of immunoreactive FOS and PER2 in 3 brain regions of pregnant and diestrous rats.

Fos expression in arousal and reward areas of the brain in grass rats following induced wakefulness.

In the diurnal grass rat nocturnal voluntary wakefulness induces Fos expression in specific cellular populations of arousal and reward areas of the brain. Here, we evaluated whether involuntary wakefulness would result in similar patterns of Fos expression. We assessed this question using male grass rats that were sleep deprived for 6h by gentle stimulation (SD group), starting 2h before lights off (12:12 LD cycle).

Projections of the suprachiasmatic nucleus and ventral subparaventricular zone in the Nile grass rat (Arvicanthis niloticus).

The phases of many circadian rhythms differ between diurnal and nocturnal species. However, rhythms within the hypothalamic suprachiasmatic nucleus (SCN), which contains the central circadian pacemaker, are very similar, suggesting that the mechanisms underlying phase preference lie downstream of the SCN. Rhythms in Fos expression in the ventral subparaventricular zone (vSPVZ), a major target of the SCN, differ substantially between diurnal Nile grass rats and nocturnal lab rats, raising the possibility that the vSPVZ modulates the effects of SCN signals at its targets.

Characterization of copulatory behavior in female mice: evidence for paced mating.

In this study we characterized female mouse sexual behavior using a pacing paradigm similar to that used to evaluate sexual behavior in female rats. A pacing chamber was designed for use with mice and we compared the sexual behavior of female mice that were tested in both pacing and nonpacing paradigms and under different hormone conditions. We found that, like rats, female mice do pace their copulatory behavior by altering the temporal sequence of copulatory events.

Rhythms in expression of PER1 protein in the amygdala and bed nucleus of the stria terminalis of the diurnal grass rat (Arvicanthis niloticus).

In the diurnal rodent Arvicanthis niloticus (grass rats) the pattern of expression of the clock genes and their proteins in the suprachiasmatic nucleus (SCN) is very similar to that seen in nocturnal rodents. Rhythms in clock gene expression have been also documented in several forebrain regions outside the SCN in nocturnal Ratus norvegicus (lab rats). To investigate the neural basis for differences in the circadian systems of diurnal and nocturnal mammals, we examined PER1 expression in the oval nucleus of the bed nucleus of the stria terminalis (BNST-OV), and in the basolateral (BLA) and the central (CEA) amygdala of male grass rats kept in a 12:12 light/dark cycle.

Effects of sex hormones on associative learning in spontaneously hypertensive rats.

Pavlovian conditioning of a visual stimulus paired with food was examined in spontaneously hypertensive rats (SHR), which are a commonly used model for Attention-Deficit/Hyperactivity Disorder (ADHD), and in Wistar rats (normoactive control). In gonadally intact rats of both strains, males spent more time in the food cup following onset of the light than did females, indicating a stronger association of the conditioned stimulus (CS) with reward. Gonadectomy carried out in adulthood affected conditioning differently in the two strains.

Cholinergic projections to the suprachiasmatic nucleus and lower subparaventricular zone of diurnal and nocturnal rodents.

In nocturnal species cholinergic agonists alter circadian rhythm phase when injected intraventricularly or directly into the suprachiasmatic nucleus (SCN), but the phase shifts obtained differ depending upon the site being injected. Cholinergic projections reach the SCN of nocturnal laboratory rats, however, nothing is known about these projections in diurnal rodents. The first objective of this study was to evaluate the hypothesis that cholinergic projections to the SCN are only present in nocturnal species.

Temporal and spatial distribution of immunoreactive PER1 and PER2 proteins in the suprachiasmatic nucleus and peri-suprachiasmatic region of the diurnal grass rat (Arvicanthis niloticus).

The suprachiasmatic nucleus (SCN) of the hypothalamus contains the primary circadian pacemaker in both diurnal and nocturnal mammals. The lower subparaventricular zone (LSPV) immediately dorsal to the SCN may also play an important role in the regulation of circadian rhythms. The SCN contains a multitude of oscillator cells that generate circadian rhythms through transcriptional/translational feedback loops involving a set of clock genes including per1 and per2.

Analysis of the prokineticin 2 system in a diurnal rodent, the unstriped Nile grass rat (Arvicanthis niloticus).

Prokineticin 2 (PK2) is a putative output molecule from the SCN. PK2 RNA levels are rhythmic in the mouse SCN, with high levels during the day, and PK2 administration suppresses nocturnal locomotor activity in rats. The authors examined the PK2 system in a diurnal rodent, Arvicanthis niloticus, to determine whether PK2 or PK2 receptors differ between diurnal and nocturnal species.

Mammalian diurnality: some facts and gaps.

A major factor contributing to the evolution of mammals was their ability to be active during the night, a niche previously underused by terrestrial vertebrates. Diurnality subsequently reemerged multiple times in a variety of independent lineages. This paper reviews some recent data on circadian mechanisms in diurnal mammals and considers general themes that appear to be emerging from this work.