Altitudinal variation in parental energy expenditure by white-crowned sparrows.

We used the doubly labeled water technique to measure daily energy expenditure (DEE) during the incubation and feeding nestling stages in two populations of white-crowned sparrows (Zonotrichia leucophrys) - one montane and migratory, the other coastal and sedentary - that differ in thermal environment and clutch size. We assessed the birds' thermal environment by continuously monitoring (among other variables) operative temperature and wind speed both in the open and within bushes and willow thickets occupied by sparrows. From these measurements, we derived several estimates of the birds' thermal environment, including standard operative temperature (T(es)). Shade air temperature and T(es) averaged 6.6 and 10.3 degrees C lower, respectively, at the montane study site during DEE measurements. The montane population's DEE averaged 24% higher than that of the sea-level population (103.6+/-12.2 versus 83.7+/-9.6 kJ day(-1); means +/- S.D., N=31 and 22, respectively), reflecting both its larger brood size (3.7 versus 2.9) and the colder environment. The DEE:BMR ratio was lowest in the sea-level population (2.1 versus 2.6), but neither population worked to their physiological capacity to produce young. DEE was significantly correlated with temperature across populations, with T(es) explaining 42% of the variation in DEE. Statistically removing the effect of temperature by adjusting DEE to a common temperature reduced the difference in DEE between populations by 34% to 87.7 and 100.8 kJ day(-1), respectively, for sea-level and montane populations. Basal and resting metabolic rates were similar in both populations, implying that greater activity in the montane population accounted for its higher temperature-adjusted DEE. Our results indicate that the thermal context within which behavior occurs can significantly affect interindividual variation in DEE. Attempts to assess reproductive effort by measuring DEE should therefore account explicitly for the effect of temperature.

Effects of temperature on photoperiodically induced reproductive development, circulating plasma luteinizing hormone and thyroid hormones, body mass, fat deposition and molt in mountain white-crowned sparrows, Zonotrichia leucophrys oriantha.

The mountain white-crowned sparrow, Zonotrichia leucophrys oriantha, breeds in subalpine meadows throughout many mountainous regions of western North America. Mathematical analysis of 20 years of egg-laying dates at Tioga Pass, California (3030m elevation) indicated a highly predictable breeding season suggesting that precise environmental cues such as the annual change in day length were important for regulating reproductive function. Additionally, it appeared that there was sufficient yearly variation in the timing of breeding to suggest that other environmental cues may also be important for regulating adjustments in reproductive development and regression. Captive populations of Z. l. oriantha showed strong responses in gonadal development following transfer to longs days (15L 9D) and low temperature (5 degrees C) slowed down photoperiodically induced gonadal growth and subsequent regression, in both males and females. High temperature of 30 degrees C tended to accelerate gonadal development and regression whereas gonadal development was intermediate in a group exposed to 20 degrees C. Prior exposure to these temperature regimes while on short days (9L 15D) had no effect on body mass, fat, or plasma levels of luteinizing hormone (LH) and thyroid hormones. Curiously there was no effect of temperature on photoperiodically induced rises in LH in either sex despite marked effects on gonadal growth. Brood patch development was also enhanced in females exposed to 30 degrees C. Corticosterone levels measured in a subset of plasma samples from this experiment indicated no effect of temperature suggesting that the retarded gonadal development at 5 degrees C was not a result of thermal stress. Although there was a robust effect of photostimulation on thyroid hormone levels in blood of both sexes, temperature treatment had no effect on tri-iodothyronine (T3) concentrations. However, plasma levels of thyroxine (T4) were lower initially at 5 degrees C versus 20 and 30 degrees C treatments. This may be related to the protracted gonadal cycle at 5 degrees C versus the truncated gonadal cycle at 30 degrees C. Molt score, an indication of post-reproductive state and onset of photorefractoriness, was delayed in birds exposed to 5 degrees C. Body mass, and to a lesser extent fat score, tended to be lowest in birds exposed to 5 degrees C compared with those at 20 and 30 degrees C. These results demonstrate that ambient temperature significantly affected photoperiodically induced gonadal development and regression in these birds. The endocrine mechanisms underlying these effects require further study.