Redefining physiological responses of moose (Alces alces) to warm environmental conditions.

We tested the concept that moose (Alces alces) begin to show signs of thermal stress at ambient air temperatures as low as 14 °C. We determined the response of Alaskan female moose to environmental conditions from May through September by measuring core body temperature, heart rate, respiration rate, rate of heat loss from exhaled air, skin temperature, and fecal and salivary glucocorticoids. Seasonal and daily patterns in moose body temperature did not passively follow the same patterns as environmental variables. We used large changes in body temperature (≥1.25 °C in 24hr) to indicate days of physiological tolerance to thermal stressors. Thermal tolerance correlated with high ambient air temperatures from the prior day and with seasonal peaks in solar radiation (June), ambient air temperature and vapor pressure (July). At midday (12:00hr), moose exhibited daily minima of body temperature, heart rate and skin temperature (difference between the ear artery and pinna) that coincided with daily maxima in respiration rate and the rate of heat lost through respiration. Salivary cortisol measured in moose during the morning was positively related to the change in air temperature during the hour prior to sample collection, while fecal glucocorticoid levels increased with increasing solar radiation during the prior day. Our results suggest that free-ranging moose do not have a static threshold of ambient air temperature at which they become heat stressed during the warm season. In early summer, body temperature of moose is influenced by the interaction of ambient temperature during the prior day with the seasonal peak of solar radiation. In late summer, moose body temperature is influenced by the interaction between ambient temperature and vapor pressure. Thermal tolerance of moose depends on the intensity and duration of daily weather parameters and the ability of the animal to use physiological and behavioral responses to dissipate heat loads.

Robust Responses of Female Caribou to Changes in Food Supply.

AbstractUngulates can respond to changes in food supply by altering foraging behavior, digestive function, and metabolism. A multifaceted response to an environmental change is considered robust. Short seasons of plant growth make herbivores sensitive to changes in food supply because maintenance and production must be accomplished in less time with fewer options in a more fragile response. Caribou live at high latitudes where short summers constrain their response to changes in food supply. We measured the ability of female caribou to resist and tolerate changes in the quality and quantity of their food supply during winter and summer. Caribou resisted changes in food abundance and quality by changing food intake and physical activity with changes in daily temperature within each season. Peak food intake rose by 134% from winter pregnancy to summer lactation (98 vs. 229 g kg-0.75 d-1), as digestible requirements to maintain the body increased by 85% for energy (1,164 vs. 2,155 kJ kg-0.75 d-1) and by 266% for N (0.79 vs. 2.89 g N kg-0.75 d-1). Caribou required a diet with a digestible content of 12 kJ g-1 and 0.8% N in pregnancy, 18 kJ g-1 and 1.9% N in early lactation, and 11 kJ g-1 and 1.2% N in late lactation, which corresponds with the phenology of the wild diet. Female caribou tolerated restriction of ad lib. food intake to 58% of their energy requirement (680 vs. 1,164 kJ kg-0.75 d-1) during winter pregnancy and to 84% of their energy requirement (1,814 vs. 2,155 kJ kg-0.75 d-1) during summer lactation without a change in stress level, as indicated by fecal corticosterone concentration. Conversely, caribou can respond to increased availability of food with a spare capacity to process digestible energy and N at 123% (2,642 vs. 2,155 kJ kg-0.75 d-1) and 145% (4.20 vs. 2.89 g N kg-0.75 d-1) of those respective requirements during lactation. Robust responses to changes in food supply allow caribou to sustain reproduction, which would buffer demographic response. However, herds may decline when thresholds of behavioral resistance and physiological tolerance are frequently exceeded. Therefore, the challenge for managing declining populations of caribou and other robust species is to identify declines in robustness before their response becomes fragile.

Trapped between food, heat, and insects: Movement of moose (Alces alces) and exposure to flies in the boreal forest of Alaska.

Moose (Alces alces) in the boreal forest habitats of Alaska are unlike other northern ungulates because they tolerate high densities of flies (Diptera) even though flies cause wounds and infections during the warm summer months. Moose move to find food and to find relief from overheating (hyperthermia) but do they avoid flies? We used GPS collars to measure the rate of movement (m⋅h-1) and the time spent (min⋅day-1) by enclosed moose in four habitats: wetlands, black spruce, early seral boreal forest, and late seral boreal forest. Fly traps were used in each habitat to quantify spatio-temporal abundance. Average daily air temperatures increased into July when peak biomass of forage for moose was greatest in early seral boreal forest habitats (424.46 vs. 25.15 kg⋅ha-1 on average in the other habitats). Average daily air temperatures were 1.7°C cooler in black spruce than other habitats, but fly abundance was greatest in black spruce (approximately 4-fold greater on average than the other habitats). Moose increased their movement rate with counts of biting flies (mosquitoes, black flies, horse and deer flies), but not non-biting flies (coprophagous flies). However, as air temperature increased (above 14.7°C) moose spent more time in fly-abundant black spruce, than early seral boreal forest, showing great tolerance for mosquitoes. Warm summer temperatures appear to cause moose to trade-off foraging in fly-sparse habitats for resting and dissipating heat in shady, wet habitats with abundant flies that adversely affect the fitness of moose.

Tracking reproductive events: Hoof growth and steroid hormone concentrations in hair and hoof tissues in moose (Alces alces).

Measurements of reproductive and stress-related hormones in keratinous tissues (e.g. hair, claws, hooves, baleen) can provide a record of stress and reproductive response in wildlife. We evaluated a method to collect keratin tissue from hooves of immobilized moose (Alces alces) and validated enzyme immunoassays for measuring cortisol and progesterone in hooves and hair. We also measured the annual growth and wear rates of moose hooves. Progesterone (range: 1.0-43.7 pg/mg) and cortisol (range: 0.05-2.9 pg/mg) were measurable and showed variation among hoof samples and moose. Pregnant females had twice as high progesterone concentrations (18.00 ± 3.73 pg/mg) from hoof sample locations post breeding compared to non-pregnant moose (9.40 ± 0.25 pg/mg). Annual hoof growth differed between the front (5.58 ± 0.12 cm) and rear (4.73 ± 0.13 cm) hooves and varied by season with higher growth rates during summer which decreased into autumn and winter. Adult female hooves represented between 1.6 and 2.1 years of growth and included up to two reproductive cycles. We established a method to estimate hoof growth rate and applied this to postmortem samples and were able to detect previous pregnancies. Shoulder guard hairs grew between August and March including during late gestation; however, hair progesterone concentrations (range: 2-107.1 pg/mg) were not related to reproductive state. Hair cortisol concentrations in our study (range: 0.2-15.9 pg/mg) were within the range of values previously reported for cervids. Our study supports the use of hooves for longitudinal sampling and measuring reproductive and stress-related hormones, providing a new tool for tracking reproductive events and understanding what variables may contribute to population level changes in reproduction.

Behaviour influences thermoregulation of boreal moose during the warm season.

Management of large herbivores depends on providing habitats for forage supply and refuge from risks of temperature, predation and disease. Moose (Alces alces) accumulate body energy and nutrient stores during summer, while reducing the impact of warm temperatures through physiological and behavioural thermoregulation. Building on the animal indicator concept, we used rumen temperature sensors and GPS collars on captive moose (n = 6) kept in large natural enclosures to evaluate how behaviour and habitat selection influence the rate of change in rumen temperature during the growing season on the Kenai Peninsula, Alaska, USA. We compared movement and habitat selection of individual females during tolerance days (daily amplitude in rumen temperature was ≥1.2°C in 24 h) with those of control days (daily amplitude in rumen temperature was < 1.2°C) before and after the tolerance day. Moose moved more during tolerance days (172 m • h-1; 95% confidence intervals (CI) = 149-191 m • h-1) than on control days (151 m • h-1; 95% CI = 128-173 m • h-1). The rate of change in rumen temperature (°C • h-1) declined with low to moderate movement rates that were probably associated with foraging in all habitats. Movement only increased the rate of change in rumen temperature at high activity (~ > 500 m • h-1). Additionally, the relationship between rate of change in rumen temperature and movement rate was different during tolerance and control days in open meadow and wetland habitats. In all habitats except wetlands, the rate of change in rumen temperature increased while resting, which probably is a result of diet-induced thermogenesis. Our study demonstrates that the behavioural choices of moose on the landscape are associated with the rate of change in rumen temperature and their ability to thermoregulate. Wildlife managers must consider high-value habitats where wildlife can employ both behavioural and physiological mechanisms to tolerate warm ambient conditions in a landscape of forage, predators and pests.

Preliminary investigation of proton and helium ion radiation effects on fluorescent dyes for use in astrobiology applications.

The Specific Molecular Identification of Life Experiment (SMILE) instrument (Sims et al. Planet. Space Science 2005, 53, 781-791) proposes to use specific molecular receptors for the detection of organic biomarkers on future astrobiology missions (e.g., to Mars). Such receptors will be used in assays with fluorescently labeled assay reagents. A key uncertainty of this approach is whether the fluorescent labels used in the system will survive exposure to levels of solar and galactic particle radiation encountered during a flight to Mars. Therefore, two fluorescent dyes (fluorescein and Alexa Fluor 633) have been exposed to low-energy proton and alpha radiation with total fluences comparable or exceeding that expected during an unshielded cruise to Mars. The results of these initial experiments are presented, which show that both dyes retain their fluorescent properties after irradiation. No significant alteration in the absorption and emission wavelengths or the quantum yields of the dyes with either radiation exposure was found. These results suggest other structurally similar fluorophores will likely retain their fluorescent properties after exposure to similar levels of proton and alpha radiation. However, more extensive radiation fluorophore testing is needed before their suitability for astrobiology missions to Mars can be fully confirmed.