Literature from the past 168 years has been filtered to provide a unified summary of the regional distribution of cutaneous water and electrolyte losses. The former occurs via transepidermal water vapour diffusion and secretion from the eccrine sweat glands. Daily insensible water losses for a standardised individual (surface area 1.8 m2) will be 0.6–2.3 L, with the hands (80–160 g.h−1) and feet (50–150 g.h−1) losing the most, the head and neck losing intermediate amounts (40–75 g.h−1) and all remaining sites losing 15–60 g.h−1. Whilst sweat gland densities vary widely across the skin surface, this same individual would possess some 2.03 million functional glands, with the highest density on the volar surfaces of the fingers (530 glands.cm−2) and the lowest on the upper lip (16 glands.cm−2). During passive heating that results in a resting whole-body sweat rate of approximately 0.4 L.min−1, the forehead (0.99 mg.cm−2.min−1), dorsal fingers (0.62 mg.cm−2.min−1) and upper back (0.59 mg.cm−2.min−1) would display the highest sweat flows, whilst the medial thighs and anterior legs will secrete the least (both 0.12 mg.cm−2.min−1). Since sweat glands selectively reabsorb electrolytes, the sodium and chloride composition of discharged sweat varies with secretion rate. Across whole-body sweat rates from 0.72 to 3.65 mg.cm−2.min−1, sodium losses of 26.5–49.7 mmol.L−1 could be expected, with the corresponding chloride loss being 26.8–36.7 mmol.L−1. Nevertheless, there can be threefold differences in electrolyte losses across skin regions. When exercising in the heat, local sweat rates increase dramatically, with regional glandular flows becoming more homogeneous. However, intra-regional evaporative potential remains proportional to each local surface area. Thus, there is little evidence that regional sudomotor variations reflect an hierarchical distribution of sweating either at rest or during exercise.
The purpose of this review is to describe the unique anatomical and physiological features of the hands and feet that support heat conservation and dissipation, and in so doing, highlight the importance of these appendages in human thermoregulation. For instance, the surface area to mass ratio of each hand is 4-5 times greater than that of the body, whilst for each foot, it is ~3 times larger. This characteristic is supported by vascular responses that permit a theoretical maximal mass flow of thermal energy of 6.0 W (136 W m(2)) to each hand for a 1 °C thermal gradient. For each foot, this is 8.5 W (119 W m(2)). In an air temperature of 27 °C, the hands and feet of resting individuals can each dissipate 150-220 W m(2) (male-female) of heat through radiation and convection. During hypothermia, the extremities are physiologically isolated, restricting heat flow to <0.1 W. When the core temperature increases ~0.5 °C above thermoneutral (rest), each hand and foot can sweat at 22-33 mL h(-1), with complete evaporation dissipating 15-22 W (respectively). During heated exercise, sweat flows increase (one hand: 99 mL h(-1); one foot: 68 mL h(-1)), with evaporative heat losses of 67-46 W (respectively). It is concluded that these attributes allow the hands and feet to behave as excellent radiators, insulators and evaporators.
Thermally induced eccrine sweating is cholinergically mediated, but other neurotransmitters have been postulated for psychological (emotional) sweating. However, we hypothesized that such sweating is not noradrenergically driven in passively heated, resting humans. To test this, nine supine subjects were exposed to non-thermal stimuli (palmar pain, mental arithmetic and static exercise) known to evoke sweating. Trials consisted of the following four sequential phases: thermoneutral rest; passive heating to elevate (by ∼1.0• C) and clamp mean body temperature and steady-state sweating (perfusion garment and footbath); an atropine sulphate infusion (0.04 mg kg −1 ) with thermal clamping sustained; and following clamp removal. Sudomotor responses from glabrous (hairless) and non-glabrous skin surfaces were measured simultaneously (precursor and discharged sweating). When thermoneutral, these non-thermal stimuli elicited significant sweating only from the palm (P < 0.05). Passive heating induced steady-state sweating ranging from 0.20 ± 0.04 (volar hand) to 1.40 ± 0.14 mg cm −2 min −1 (forehead), with each non-thermal stimulus provoking greater secretion (P < 0.05). Atropine suppressed thermal sweating, and it also eliminated the sudomotor responses to these nonthermal stimuli when body temperatures were prevented from rising (P > 0.05). However, when the thermal clamp was removed, core and skin temperatures became further elevated and sweating was restored (P < 0.05), indicating that the blockade had been overcome, presumably through elevated receptor competition. These observations establish the dependence of both thermal and non-thermal eccrine sweating from glabrous and non-glabrous surfaces on acetylcholine release, and challenge theories concerning the psychological modulation of sweating. Furthermore, no evidence existed for the significant participation of non-cholinergic neurotransmitters during any of these stimulations.
Thermal sweating from the human torso accounts for about half of the whole-body sweat secretion, yet its intra-segmental distribution has not been thoroughly examined. Therefore, the aim of the current study was to provide a detailed description of the distribution of eccrine sweating within the torso during passively-induced (water-perfusion garment: 40 degrees C) and progressively increasing, exercise-related thermal strain (36 degrees C, 60% relative humidity). Sudomotor function was measured in ten males using ventilated sweat capsules (3.16 cm(2)) attached to twelve sites on the ventral (four), lateral (three) and dorsal (four) torso, and upper shoulder surfaces. Sweating increased asymptotically in all sites, with the final core temperature averaging 39.7 degrees C (+/-0.1) and heart rates being 181 b min(-1) (+/-2). During exercise, the mean torso sweat rate averaged 1.35 mgcm(-2)min(-1), with sweating from the lateral torso surfaces generally being the lowest. Each of the between-site comparisons with the lateral torso differed significantly (P < 0.05), except for comparisons with the chest (P = 0.051) and shoulder (P > 0.05). The intra-segmental differences between the lateral torso and the chest, abdomen, upper- and lower-back areas were significantly accentuated during exercise. From these data, it is evident that the torso is another region that does not have a uniform distribution of thermally-induced sweating. Thus, it is no longer acceptable for researchers, modellers, sweating manikins engineers or clothing manufacturers to assume that the sweat rates for all local sites within any body segment are equivalent.
The importance of the head in dissipating body heat under hot conditions is well recognised, although very little is known about local differences in sweat secretion across the surface of the head. In this study, we focused on the intra-segmental distribution of head sweating. Ten healthy males were exposed to passive heating and exercise-induced hyperthermia (36 degrees C, 60% relative humidity, water-perfusion suit: 46 degrees C), with ventilated sweat capsules (3.16 cm(2)) used to measure sweat rates from the forehead and nine sites inside the hairline. Sweat secretion from both non-hairy (glabrous) and hairy areas of the head increased linearly with increments in work rate and core temperature, with heart rate and core temperature peaking at 175 b min(-1) (+/-6) b min(-1) and 39.2 degrees C (+/-0.1). The mean sweat rate during exercise for sites within the hairline was 1.95 mg cm(-2) min(-1). However, the evolution of this secretion pattern was not uniformly distributed within the head, with the average sweat rate for the top of the head being significantly lower than at the anterior lateral aspect of the head (P < 0.05), and representing only 30% of the forehead sweat rate (P < 0.05). It is hypothesised that these intra-segmental observations may reflect variations in the local adaptation of eccrine glands to differences in local evaporation associated either with bipedal locomotion, which will influence forehead sweating, or the hidromeiotic suppression of sweating, which impacts upon sweat glands within the hairline.
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