During three breeding seasons, variation in the timing of estrus was monitored in wild red squirrels (Tamiasciurus hudsonicus) in a jack pine forest in central Alberta. The proximate environmental factors that affect the timing of the onset of the breeding season in female squirrels, and variation in dates of estrus, were studied in parallel laboratory and field manipulations. Wild squirrels given ad libitum sunflower seed experienced a positive energy balance (weight gains) earlier in the season than did females on a normal diet. Seed addition advanced dates of estrus by an average of 3 weeks. Three potential cuing mechanisms for the seasonal onset of estrus were proposed and tested: onset of estrus is cued by (i) a simple response to net energy gain, (ii) an interaction between an endogenous circannual rhythm and net energy gain, or (iii) an interaction between annual changes in photoperiod and net energy gain in the absence of a circannual rhythm. Our results refute the first and third mechanisms, and support the idea that red squirrels have a circannual reproductive cycle entrained by photoperiod. Net energy gain interacts with this annual cycle to fine tune the time of ovulation. In laboratory experiments, female squirrels exposed to increasing day length experienced estrus within the normal season. Squirrels kept in constant short-day conditions (< 12 h), had delayed estrus cycles. Short-day conditions did not prevent the annual cycle of reproduction, indicating that an endogenous rhythm exists and is entrained by increasing day length. Comparisons with hibernating squirrels are discussed.
Net energy gain in direct laser acceleration due to enhanced dephasing induced by an applied magnetic field
