Diversity of Function of SCN Pacemakers in Behavior and Ecology of Three Species of Sciurid Rodents

Information on the widespread function of the suprachiasmatic (SCN) pacemaker has increased dramatically in the past decade for laboratory rodents and for humans. Interest is now also growing in the adaptive value of SCN pacemakers in wild species of free-living mammals in natural pristine habitat. The squirrel family, Sciuridae, is highly diverse in North America. Radiation into many specialized groups has evolved in response to the temporal and spatial characteristics of specific habitats. The sciurids thus offer possibilities for comparison of SCN function for closely related species from very different habitats. Results from field and laboratory investigations concerning the ecological significance of the SCN are reported here for three ground squirrel species. The semi-fossorial antelope squirrels of the American southwest deserts were arrhythmic in above ground activities after deletion of the SCN pacemaker. In a desert enclosure, predation rates rose dramatically for lesioned animals in comparison to intact controls. The semi-arborial chipmunks were prone to capture by mustelid burrow predators such as weasels after SCN-deletion, even though the chipmunks did not exhibit any above ground activity at night. The telltale cue for the predator was apparently arrhythmic restlessness within the complex home burrow. The semi-fossorial golden-mantled squirrels of the Cascade Mountains of western United States rely on profound, long-lasting hibernation for over-wintering. Periods of torpor alternate with brief euthermic arousal episodes lasting less than 24 h throughout the 5- to 8-month winter phase. Deletion of the SCN in golden-mantled squirrels resulted in aberrations of hibernation arousal patterns that were metabolically expensive and put individuals at risk of starvation immediately after emergence in spring.     

Circadian photoentrainment: parameters of phase delaying

Experiments were carried out using simulated den cages to delineate specific characteristics of phase delaying in circadian photoentrainment of a nocturnal rodent, the flying squirrel. The principal experiments entailed presentation of one to five consecutive 15-min white-light pulses per activity cycle at activity onset to animals free-running in darkness, in order to determine the immediate and final phase-shifting effect. Auxiliary experiments recorded entrainment patterns on light-dark (LD) schedules in the den cages. Phase response curves (PRCs) based on 15-min white-light pulses in standard wheel cages were also constructed for these animals as background information for interpreting the phase-delaying experiments. Exposure of a den animal to light by light sampling at the time of initial arousal from the rest state at circadian time (CT) 12, either by an LD schedule or by a 15-min light pulse, resulted in a return to the nest box for a short rest period. The phase delay occurring after a single light exposure at activity onset was equal to the induced rest, thus suggesting an immediate phase shift. The maximum delay was about 1 1/2 hr/cycle, with the amount of delay related to the number of light exposures. During the photoentrained state on an LD schedule, the activity rhythm of a den-housed animal was essentially free-running on the days following a phase delay. The data are used to expand current models for photoentrainment of circadian activity rhythms in nocturnal rodents.