1. According to risk-sensitive foraging theory, animals should make foraging decisions that balance nutritional costs and gains to promote fitness. Human disturbance is a form of perceived risk that can prompt avoidance of risky habitat over acquisition of food. Consequently, behavioural responses to perceived risk could induce nutritional costs.
2. Population declines often coincide with increases in human disturbance, which likely is associated with direct and indirect habitat loss. Nevertheless, behavioural and physiological responses to perceived risks associated with human disturbance could be an added nutritional deficit with population-level repercussions.
3. Using GPS-collar data from three populations of migratory mule deer Odocoileus hemionus exposed to a gradient of established industrial energy development on winter ranges where direct and indirect habitat loss were well documented, we evaluated whether exposure and behavioural responses to human disturbance alter changes in nutritional condition (i.e. fat reserves) over winter.
4. Although animals exhibited behaviours indicative of perceived risk of human disturbance, such as increased movement rates and avoidance of infrastructure, exposure and behavioural responses to human disturbance had little to no measurable effect on fat loss over a winter. Instead, catabolism of fat reserves occurred primarily as a function of the amount of fat animals had entering winter, suggesting that, in the short term, animals were able to mitigate the energetic costs of perceived risk of human disturbance over winter. Animals, however, did not appear to overcome persistent food limitations, and animals with less food availability lost more fat over winter.
5. Our findings heed caution in using short-term behavioural and physiological responses to inform long-term nutritional consequences of human disturbance. Although animals appear to mitigate the energetic costs of perceived risk, food limitations exacerbated by broader-scale avoidance of food near human disturbance may be the primary pathway causing the frequently observed population declines following human disturbance to pristine landscapes.

中文翻译：

---

对人类改变的景观的短期反应不会影响迁徙有蹄类动物的脂肪动态

1. 根据风险敏感觅食理论，动物应该做出平衡营养成本和收益的觅食决定，以促进健康。人为干扰是一种感知风险的形式，可以促使人们避开危险的栖息地而不是获取食物。因此，对感知风险的行为反应可能会导致营养成本。
2. 人口减少通常与人类干扰增加同时发生，这可能与直接和间接栖息地丧失有关。然而，对与人类干扰相关的感知风险的行为和生理反应可能是一种额外的营养缺乏，对人群产生影响。
3. 使用来自三个迁徙骡鹿Odocoileus hemionus种群的 GPS 项圈数据，它们在冬季范围内暴露于已建立的工业能源发展梯度，其中直接和间接栖息地丧失得到充分记录，我们评估了暴露和对人类干扰的行为反应是否会改变营养变化冬季状况（即脂肪储备）。
4. 尽管动物表现出的行为表明人类干扰的感知风险，例如增加的运动速率和避免基础设施，但暴露和对人类干扰的行为反应对整个冬天的脂肪损失几乎没有或没有可测量的影响。相反，脂肪储备的分解代谢主要作为进入冬季的脂肪动物数量的函数而发生，这表明，在短期内，动物能够减轻冬季人类干扰风险的能量成本。然而，动物似乎并没有克服持续的食物限制，食物供应量较少的动物在冬天失去了更多的脂肪。
5. 我们的研究结果注意谨慎使用短期行为和生理反应来告知人类干扰的长期营养后果。尽管动物似乎可以减轻感知风险的能量成本，但由于在人类干扰附近更广泛地避免食物而加剧的食物限制可能是导致人类干扰原始景观后经常观察到的人口下降的主要途径。