Muscular changes accompanying and/or promoting the rapid postnatal improvement of the thermogenic efficiency of shivering were investigated in piglets. Animals were obtained at birth or killed after 5 days at thermoneutrality (34-30 degrees C) or in the cold (24-15 degrees C), to stimulate intense shivering thermogenesis. Fast-twitch-glycolytic (longissimus lumborum) and slow-twitch-oxidative (rhomboid) muscles were prepared for electron microscopic examination and chemical measurements. Muscle-specific changes in energy stores and metabolism were observed after birth, including the switch from glycogen to lipids and variation of the lactate/pyruvate ratio corresponding to the progressive acquisition of the metabolic type of the mature muscles. There was major age-related and/or cold-induced development of the structures involved in excitation-contraction coupling (triadic profiles, +80% in the cold), oxidative metabolism (number of lipid droplets, +81% with age in the cold; number of mitochondria, +29% with age or cold; surface of mitochondrial inner membranes, +18% with age and +32% in the cold) and contraction potential (myofibril volume, +62% with age). In contrast, neither age nor cold affected capillary volume density and capillary-to-fibre ratio. The observed changes reflect the immaturity and remarkable plasticity of piglet skeletal muscle and are likely to underlie its enhanced capacity for shivering thermogenesis after birth.

SUMMARYExposure to a temperature of 14°C was used to induce a progressive hypothermia in fourteen conscious newborn piglets. Heat production, body (rectal) and skin (between the shoulders) temperatures and shivering intensity assessed as the electromyographic activity (EMG) of longissimus thoracis muscle were measured until body temperature reached 30°C and during a recovery period of 2 h at an ambient temperature of 24°C (n = 7) or 34°C (n = 7). During body cooling, heat production increased up to 9.67 + 1 28 W (kg BW)-1, but started to decrease below a body temperature threshold of 34-4 + 0-7 'C. EMG activity increased (P < 0.023) curvilinearly during body cooling; the main increase occurred between body temperatures of 38 and 33 'C (+142 %, P < 0.001), and changes in EMG activity between 33 and 30°C were not significant (+18%, P > 0 1). A marked increase in circulating levels of glucose (+312 %, P < 0.001), glucagon (+76%, P < 0.05), adrenaline (+172 %, P < 0.05) and noradrenaline (+113 %, P < 0.05) occurred during body cooling. Insulin levels were not detectable at 2 h of life and increased during body cooling. During 2 h of rewarming at 24°C, heat production and EMG activity remained elevated, changes in carbohydrate metabolism were not completely reversed and the final body temperature was only 35-6 + 0-9 'C. Rewarming of the piglets was faster at 34 'C. There was a net influx of heat into the animals and heat production and shivering activity decreased when body temperature reached 33-9 + 0-5 'C; the final body temperature was 37.5 + 0-2 'C. Circulating levels of lactate, glucagon and catecholamines returned to control levels. These results show that in conscious piglets exposed to a constant cold temperature there is an inverse relationship between EMG activity and body temperature during moderate hypothermia and that the thermoregulatory response and carbohydrate metabolism of the piglet are seriously impaired below a body temperature of 34°C.

SUMMARYNewborn pigs rely essentially on shivering thermogenesis in the cold. In order to understand the rapid postnatal enhancement of thermogenic capacities in piglets, the oxygen and nutrient uptake of hindquarters was measured in vivo in 1-(n = 6) and 5-day-old (n = 6) animals at thermal neutrality and during cold exposure. The hindquarters were considered to represent a skeletal muscle compartment. Indirect calorimetry and arterio-venous techniques were used. The cold challenge (23°C at 1 day old and 15°C at 5 days old for 90 min) induced a similar increase (+90 %) in regulatory heat production at both ages. Hindquarters blood flow was higher at 5 days than 1 day old at thermal neutrality (26 + 3 vs. 17 + 1 ml min-' (100 g hindquarters)-') and its increase in the cold was much more marked (+65 % at 5 days old vs. +25 % at 1 day old). Oxygen extraction by the hindquarters rose from 30-35 % at thermal neutrality to 65-70 % in the cold at both ages. The calculated contribution of skeletal muscle to total oxygen consumption averaged 34-40% at thermal neutrality and 50-64% in the cold and skeletal muscle was the major contributor to regulatory thermogenesis. Based on hindquarters glucose uptake and lactate release, carbohydrate appeared to be an important fuel for shivering. However, net uptake of fatty acids increased progressively during cold exposure at 5 days old. The enhancement in muscular blood supply and fatty acid utilization during shivering is probably related to the postnatal improvement in the thermoregulatory response of the piglet.

Early after birth, piglets rely almost exclusively on muscular shivering thermogenesis to produce heat in the cold and this can possibly modulate skeletal muscle development. An experiment involving 10 individually housed piglets was conducted to determine the influence of cold (24-15 degrees C, D5C group) vs. thermoneutrality (34-30 degrees C, D5TN group) between birth and 5 days on myosin heavy chain (MyHC) polymorphism and metabolic characteristics of longissimus lumborum (LL) and rhomboideus (RH) muscles. Five additional piglets were sacrificed at birth. Piglets exposed to cold received 43% more artificial milk on a liveweight basis in order to achieve similar growth rates. D5C piglets produced 93% more heat and exhibited intense shivering during the whole experiment. Contractile and metabolic characteristics of muscles were determined by immunocytochemistry, electrophoresis and enzyme activities. At least eight MyHC isoforms were detected, including atypical expressions of the alpha-cardiac and extraocular isoforms. Dramatic changes in MyHC composition, myofiber cross-sectional area (CSA) and energy metabolism occurred between birth and 5 days. Cold exposure did not affect either the total number of fibers or the CSA, but it did influence muscle maturation. In particular, it increased the expression of alpha-cardiac and type I MyHC, and decreased that of fetal MyHC, confirming an acceleration in the rate of postnatal maturation. An increase in oxidative enzyme activities was observed in both muscles in the cold, whereas the activity of a glycolytic enzyme, lactate dehydrogenase, remained unchanged. Cold exposure also induced an increase in T3 plasma levels. The extent to which these changes are the result of sustained shivering or are due to the action of hormonal factors, such as thyroid hormones, are discussed.

To determine whether newborn pigs are able to display adequate cardiovascular adjustments favouring shivering thermogenesis in skeletal muscles soon after birth, regional blood flow and fractional distribution of cardiac output were determined in 1-day-old (n = 6) and 5-day-old (n = 6) conscious piglets at thermal neutrality and during cold exposure, using coloured microspheres. Five-day-old piglets stayed with the sow before the experiment. The cold challenge was designed to induce a similar increase (approximately +90%) in heat production at both ages. Skeletal muscle blood flow increased with both age (p < 0.05) and cold exposure (p < 0.001), with the effect of cold being more pronounced in 5-day-old piglets than in 1-day-old piglets (+60%, p < 0.05). The difference between individual muscles increased with age, with fractional blood flow being 41% higher in rhomboideus than in longissimus thoracis muscle during cold exposure in 5-day-old piglets (p < 0.05). Cardiac output was similar at both ages and increased by 23% in the cold (p < 0.001). At 1 day of age, there was no redistribution of cardiac output among the internal organs during the cold challenge, while at 5 days of age, the increase in muscle fractional blood flow was associated with a reduction (p < 0.05) in the fraction of cardiac output reaching the skin (-24%), the small intestine (-21%), and the liver (-20%). In conclusion, these results suggest that there is a rapid postnatal improvement of cardiovascular adjustments favouring blood perfusion and probably heat production during cold-induced shivering in the most oxidative muscles studied. This cardiovascular response may play a role in the postnatal enhancement of thermoregulation in piglets.