Shivering and nonshivering thermogenesis in skeletal muscles

Humans have inherited complex neural circuits which drive behavioral, somatic, and autonomic thermoregulatory responses to defend their body temperature. While they are well adapted to dissipate heat in warm climates, they have a reduced capacity to preserve it in cold environments. Consequently, heat production is critical to defending their core temperature. As in other large mammals, skeletal muscles are the primary source of heat production recruited in cold-exposed humans. This is achieved voluntarily in the form of contractions from exercising muscles or involuntarily in the form of contractions from shivering muscles and the recruitment of nonshivering mechanisms. This review describes our current understanding of shivering and nonshivering thermogenesis in skeletal muscles, from the neural circuitry driving their recruitment to the metabolic substrates that fuel them. The presence of these heat-producing mechanisms can be measured in vivo by combining indirect respiratory calorimetry with electromyography or biomedical imaging modalities. Indeed, much of what is known regarding shivering in humans and other animal models stems from studies performed using these methods combined with in situ and in vivo neurologic techniques. More recent investigations have focused on understanding the metabolic processes that produce the heat from both contracting and noncontracting mechanisms. With the growing interest in the potential therapeutic benefits of shivering and nonshivering skeletal muscle to counter the effects of neuromuscular, cardiovascular, and metabolic diseases, we expect this field to continue its growth in the coming years.