Influence of neutral salts on the hydrothermal stability of acid-soluble collagen

The thermal stability of acid-soluble collagens was studied by circular dichroism (CD) spectroscopy. Adult bovine dermal collagen (BDC), rat-tail tendon collagen (RTC), and calf skin collagen (CSC) were compared. Despite some variability in amino acid composition and apparent molecular weight, the CD spectra for helical and unordered collagen structures were essentially the same for all the sources. The melting of these collagens occurs as a two-stage process characterized by a pretransition (T _p) followed by complete denaturation (T _d). The characteristic temperatures vary with the source of the collagen; for mature collagens (BDC, RTC) T _p = 30°C and T _d = 36deg;C, and for CSC T _p = 34°C and T _d = 40°C. Neutral salts, NaCl or KCl, at low concentrations (0.02–0.2 M) appear to bind to the collagens and shift the thermal transitions of these collagens to lower temperatures.     

Missense Mutations That Cause Bruck Syndrome Affect Enzymatic Activity, Folding, and Oligomerization of Lysyl Hydroxylase 2

Bruck syndrome is a rare autosomal recessive connective tissue disorder characterized by fragile bones, joint contractures, scoliosis, and osteoporosis. The telopeptides of bone collagen I are underhydroxylated in these patients, leading to abnormal collagen cross-linking. Three point mutations in lysyl hydroxylase (LH) 2, the enzyme responsible for the hydroxylation of collagen telopeptides, have been identified in Bruck syndrome. As none of them affects the residues known to be critical for LH activity, we studied their consequences at the molecular level by analyzing the folding and catalytic properties of the corresponding mutant recombinant polypeptides. Folding and oligomerization of the R594H and G597V mutants were abnormal, and their activity was reduced by >95% relative to the wild type. The T604I mutation did not affect the folding properties, although the mutant retained only ∼8% activity under standard assay conditions. As the reduced activity was caused by a 10-fold increase in the K_m for 2-oxoglutarate, the mutation interferes with binding of this cosubstrate. In the presence of a saturating 2-oxoglutarate concentration, the activity of the T604I mutant was ∼30% of that of the wild type. However, the T604I mutant did not generate detectable amounts of hydroxylysine in the N-terminal telopeptide of a recombinant procollagen I chain when coexpressed in insect cells. The low activity of the mutant LH2 polypeptides is in accordance with the markedly reduced extent of collagen telopeptide hydroxylation in Bruck syndrome, with consequent changes in the cross-linking of collagen fibrils and severe abnormalities in the skeletal structures.     

Regulation of local sweating in sleep-deprived exercising humans

Thermoregulatory sweating [total body (m _sw,b), chest (m _sw,c) and thigh (m _sw,t) sweating], body temperatures [oesophageal (T _oes) and mean skin temperature (T _sk)] and heart rate were investigated in five sleep-deprived subjects (kept awake for 27 h) while exercising on a cycle (45 min at approximately 50% maximal oxygen consumption) in moderate heat (T _air andT _wall at 35° C. Them _sw,c andm _sw,t were measured under local thermal clamp (T _sk,1), set at 35.5° C. After sleep deprivation, neither the levels of body temperatures (T _oes,T _sk) nor the levels ofm _{sw, b},m _{sw, c} orm _{sw, t} differed from control at rest or during exercise steady state. During the transient phase of exercise (whenT _sk andT _sk,1 were unvarying), them _{sw, c} andm _{sw, t} changes were positively correlated with those ofT _oes. The slopes of them _{sw, c} versusT _oes, orm _{sw, t} versusT _oes relationships remained unchanged between control and sleep-loss experiments. Thus the slopes of the local sweating versusT _oes, relationships (m _{sw, c} andm _{sw, t} sweating data pooled which reached 1.05 (SEM 0.14) mg·cm^–2·min^–1°C^–1 and 1.14 (SEM 0.18) mg·cm^–2·min^–1·°C^–1 before and after sleep deprivation) respectively did not differ. However, in our experiment, sleep deprivation significantly increased theT _oes threshold for the onset of bothm _{sw, c} andm _{sw, t} (+0.3° C,P<0.001). From our investigations it would seem that the delayed core temperature for sweating onset in sleep-deprived humans, while exercising moderately in the heat, is likely to have been due to alterations occurring at the central level.     

Sex- and age-related changes in the biophysical properties of cuticular lipids of the housefly,Musca domestica

We examined the biophysical properties of cuticular lipids isolated from the housefly, Musca domestica. Melting temperatures (T_m) of surface lipids isolated from female houseflies decreased from 39.3 °C to 35.3 °C as the females attained sexual maturity and produced sex pheromone, whereas those prepared from males did not change with age. Lipids melted over a 10–25 °C temperature range, and their physical properties were a complex function of the properties of the component lipids. The T_m of total cuticular lipids was slightly below that of cuticular hydrocarbons (HC), the predominant lipid fraction. Hydrocarbons were further fractionated into saturated, unsaturated, and methyl-branched components. The order of decreasing T_m was total alkanes > total HCs > methyl-branched alkanes > alkenes. For 1-day-old flies, measured T_ms of hydrocarbons were 1.3–5.5 °C lower than T_ms calculated from a weighted average of T_ms for saturated and unsaturated components. For 4-day-old flies, calculated T_ms underestimated T_m by 11–14 °C. © 1995 Wiley-Liss, Inc.     

Difference in thermal stability of type-I and type-III collagen from rat skin

Type-I and type-III collagens were obtained by differential salt fractionation of neutral-salt-soluble collagen from rat skin. Their thermal stabilities were determined by u.v. difference spectroscopy. The `melting' temperature (T_m) in 5mm-acetic acid of type-III collagen was almost 2°C above that of type-I collagen. Intramolecular covalent cross-linking had no effect on the thermal stability.     

Gas exchange characteristics and temperature relations of two desert annuals: A comparison of a winter-active and a summer-active species

Summary The gas exchange characteristics of two C₃ desert annuals with contrasting phenologies, Geraea canescens T. & G. (winter-active) and Dicoria canescens T. & G. (summer-active), both Asteraceae, were determined for plants grown under a moderate (25°/15° C, day/night temperature) and a high (40°/27° C) growth temperature regime. Both species had high photosynthetic capacities; maximum net photosynthetic rates were 38 and 48 mol CO₂ m^-2 s^-1 for Geraea and Dicoria, respectively, and were not influenced by growth temperature regime. However, the temperature optima of net photosynthesis shifted from 26° C for Geraea and from 28° C for Dicoria when grown under the moderate temperature regime to 31° C for both species when grown under the high temperature regime. Although the shifts in temperature optima were smaller than those observed for many desert perennials, both species showed substantial increases in photosynthetic rates at high temperatures when grown at 40°/27° C. In general, the gas exchange characteristics of Geraea and Dicoria were very similar to each other and to those reported for other C₃ desert annuals. Geraea and Dicoria experienced different seasonal patterns of change in several environmental variables. For Geraea, maximum daily air temperature (T _a) increased from 24° to 41° C over its growing season while Dicoria experienced maximum T _a at midseason (45° C). At points during their respective growing seasons when midday T _a ranged between 35° and 40° C, leaf temperatures (T ₁) of both species were below T _a and, therefore, were closer to the photosynthetic temperature optima measured in the laboratory. Leaf conductances to water vapor (g ₁) and water potentials () were high at these times, but later in their growing seasons Dicoria maintained high g ₁ and while Geraea showed large decreases in these quantities. The ability of Dicoria to successfully growth through the hot, dry summers of the California deserts may be related to its ability to acquire the available water in locally mesic habitats.     

Changes in body core temperatures and heat balance after an abrupt release of lower body negative pressure in humans

Changes in body core temperature (T _cor) and heat balance after an abrupt release of lower body negative pressure (LBNP) were investigated in 5 volunteers under the following conditions: (1) an ambient temperature (T _a) of 20 °C or (2) 35 °C, and (3)T _a of 25 °C with a leg skin temperature of 30°C or (4) 35°C. The leg skin temperature was controlled with water perfusion devices wound around the legs. Rectal (T _re), tympanic (T _ty) and esophageal (T _es) temperatures, skin temperatures (7 sites) and oxygen consumption were measured. The intensity of LBNP was adjusted so that the amount of blood pooled in the legs was the same under all conditions. When a thermal balance was attained during LBNP, application of LBNP was suddenly halted. The skin temperatures increased significantly after the release of LBNP under all conditions, while oxygen consumption hardly changed. The release of LBNP caused significant falls inT _cor s under conditions (1) and (3), but loweredT _cor s very slightly under conditions (2) and (4). The changes inT _es were always more rapid and greater than those ofT _ty andT _re. The falls inT _ty andT _re appeared to be explained by changes in heat balance, whereas the sharp drop ofT _es could not be explained especially during the first 8 min after the release of LBNP. The results suggest that a fall inT _cor after a release of LBNP is attributed to an increase in heat loss due to reflexive skin vasodilation and is dependent on the temperature of venous blood returning from the lower body. It is presumed thatT _es may not be an appropriate indicator forT _cor when venous return changes rapidly.     

Effect of ambient temperature and E. coli endotoxin upon the plasma iron level in wild house mice in winter season

Summary The effects of ambient temperatures of 10°C and 30°C and of E. coli endotoxin on brain temperature and plasma iron level were investigated in unrestrained wild house mice, Mus musculus. In control animals (i.p. saline-injected) exposed to cold environmenta the brain temperature decreased and plasma iron levels were lower than those observed under thermoneutral conditions (30°C). Animals injected i.p. with endotoxin (0.5 g·kg^-1) and placed at 30°C showed a drop in plasma iron level during the fever episode. The results provide strong evidence for a relationship between brain temperature and plasma iron level in control mice under thermoneutral conditions, and show that during cold exposure or after injection of endotoxin, there is no linear correlation between brain temperature and plasma iron. Moreover, it was found that cold stress influences plasma iron level and that this influence is not mediated by changes in brain temperature.Abbreviations EP endotoxin pyrogen - T _A ambient temperature - T _Br brain temperature - T _Br change in T _Br in relation to its initial value in feverish or control mice - T _Br difference between T _Br in feverish and control mice     

Collagen synthesis by cultured skin fibroblasts from siblings with hydroxylysine-deficient collagen

It has been previously shown that dermis from subjects with hydroxylysine-deficient collagen contains approximately 5% of normal levels of hydroxylysine and sonicates of skin fibroblasts contain less than 15% of normal levels of collagen lysyl hydroxylase activity. However, cultures of dermal fibroblasts from two siblings with hydroxylysine-deficient collagen (Ehlers-Danlos Syndrome Type VI) compared to fibroblasts from normal subjects synthesize collagen containing approximately 50% of normal amounts of hydroxylysine. The lysyl hydroxylase deficient cultures synthesize both Type I and Type III collagen in the same proportion as control cultures. Both α1(I) and α2 chains are similarly reduced in hydroxylysine content. Collagen prolyl hydroxylation by normal and mutant cells is severely depressed without ascorbate but in all cultures collagen lysyl hydroxylation is the same with or without ascorbate supplementation. In mutant cells the rate of prolyl hydroxylation measured after release of inhibition by α,α′-dipyridyl is the same as in control cells. The rate of lysyl hydroxylation is reduced in mutant cells but only to approximately 50% of normal.     

Collagen synthesis by cultured skin fibroblasts from siblings with hydroxylysine-deficient collagen.

It has been previously shown that dermis from subjects with hydroxylysine-deficient collagen contains approximately 5% of normal levels of hydroxylysine and sonicates of skin fibroblasts contain less than 15% of normal levels of collagen lysyl hydroxylase activity. However, cultures of dermal fibroblasts from two siblings with hydroxylysine-deficient collagen (Ehlers-Danlos Syndrome Type VI) compared to fibroblasts from normal subjects synthesize collagen containing approximately 50% of normal amounts of hydroxylysine. The lysyl hydroxylase deficient cultures synthesize both Type I and Type III collagen in the same proportion as control cultures. Both alpha 1(I) and alpha 2 chains are similarly reduced in hydroxylysine content. Collagen prolyl hydroxylation by normal collagen lysyl hydroxylation is the same with or without ascorbate supplementation. In mutant cells the rate of prolyl hydroxylation measured after release of inhibition by alpha, alpha'-dipyridyl is the same as in control cells. The rate of lysyl hydroxylation is reduced in mutant cells but only to approximately 50% of normal.     

Engineering approach to mixing quantification in bioreactors

Homogeneity-time is defined and introduced as the criterion for mixing quality in bioreactors. The criterion could replace the mixing time, in the case, when more than one measuring point (sensors) is included in the measuring system. Results based on the homogeneity-time and the temperature pulse method, achieved in stirred tank reactors under aerated conditions as well as in a jet-mixed tank, are presented.List of Symbols C _p,p kJ/kg K Heat capacity of the pulse medium - C _p,s kJ/kg K Heat capacity of the reactor-medium - F m³/s Flow rate of the pulse-input - i Inhomogeneity - I _N Inhomogeneity-number - M (t) °C Ideal response curve - m deNumber of combinations for certain number of sensors acc. to Table 1 - n Number of sensor - _p kg/m³ Density of the pulse medium - kg/m³ Density of the tank medium - s ₁ °C Mean absolute deviation of the sensor temperatures related on the ideal response curve s₂ s Mean absolute deviation of the homogeneity-times related on the time achieved with 6 sensors - t s Time - t (i) s Homogeneity-time - t _ps s Starting time of tracer injection - t _PE s End time of tracer injection - T _E °C Mean medium temperature at the end of experiment - T _k °C Temperature at k-th sensor position - T _p °C Pulse temperature - T _s °C Mean medium temperature before the tracer injection - V _s m³ Tank volume before pulse input     

Proteolytic activity in human skin fibroblasts harvested with trypsin and its effect on measurements of collagen hydroxylase activities.

Lysates of human skin fibroblasts harvested without the use of trypsin do not contain detectable proteolytic activity, but when trypsin is used, lysates may contain activity equal to 10 ng of trypsin/10⁷ cells. The amount of cell lysate ordinarily examined for collagen prolyl and lysyl hydroxylase activity is sufficiently small that such amounts of trypsin have no observable effect on the unhydroxylated collagen substrate. Larger amounts of trypsin cause proteolysis of the unhydroxylated collagen substrate and a reduction of both prolyl and lysyl hydroxylation with lysyl hydroxylation more affected at low trypsin concentration than prolyl hydroxylation.     

The effect of deuterium oxide on the stability of the collagen model peptides H‐(Pro‐Pro‐Gly)10‐OH,H‐(Gly‐Pro‐4(R)Hyp)9‐OH,and Type I collagen

The collagen triple helix has a larger accessible surface area per molecular mass than globular proteins, and therefore potentially more water interaction sites. The effect of deuterium oxide on the stability of collagen model peptides and Type I collagen molecules was analyzed by circular dichroism and differential scanning calorimetry. The transition temperatures (T_m) of the protonated peptide (Pro‐Pro‐Gly)₁₀ were 25.4 and 28.7°C in H₂O and D₂O, respectively. The increase of the T_m of (Pro‐Pro‐Gly)₁₀ measured calorimetrically at 1.0°C min^?1 in a low pH solution from the protonated to the deuterated solvent was 5.1°C. The increases of the T_m for (Gly‐Pro‐4(R)Hyp)₉ and pepsin‐extracted Type I collagen were measured as 4.2 and 2.2°C, respectively. These results indicated that the increase in the T_m in the presence of D₂O is comparable to that of globular proteins, and much less than reported previously for collagen model peptides [Gough and Bhatnagar, J Biomol Struct Dyn 1999, 17, 481–491]. These experimental results suggest that the interaction of water molecules with collagen is similar to the interaction of water with globular proteins, when the ratio of collagen to water is very small and collagen is monomerically dispersed in the solvent. © 2009 Wiley Periodicals, Inc. Biopolymers 93: 93–101, 2010. This article was originally published online as an accepted preprint. The “Published Online” date corresponds to the preprint version. You can request a copy of the preprint by emailing the Biopolymers editorial office at biopolymers@wiley.com     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: Recovery and its dependence on temperature

Recovery of photoinhibition in intact leaves of shade-grown kiwifruit was followed at temperatures between 10° and 35° C. Photoinhibition was initially induced by exposing the leaves for 240 min to a photon flux density (PFD) of 1 500 mol·m^-2·s^-1 at 20° C. In additional experiments to determine the effect of extent of photoinhibition on recovery, this period of exposure was varied between 90 and 400 min. The kinetics of recovery were followed by chlorophyll fluorescence at 77K. Recovery was rapid at temperatures of 25–35° and slow or negligible below 20° C. The results reinforce those from earlier studies that indicate chilling-sensitive species are particularly susceptible to photoinhibition at low temperatures because of the low rates of recovery. At all temperatures above 15° C, recovery followed pseudo first-order kinetics. The extent of photoinhibition affected the rate constant for recovery which declined in a linear fashion at all temperatures with increased photoinhibition. However, the extent of photoinhibition had little effect on the temperature-dependency of recovery. An analysis of the fluorescence characteristics indicated that a reduction in non-radiative energy dissipation and repair of damaged reaction centres contributed about equally to the apparent recovery though biochemical studies are needed to confirm this. From an interpretation of the kinetics of photoinhibition, we suggest that recovery occurring during photoinhibition is limited by factors different from those that affect post-photoinhibition recovery.Abbreviations and symbols F _o, F _m, F _v instantaneous, maximum, variable fluorescence - K _D, K _F, K _P, K _T rate constants for non-radiative energy dissipation, fluorescence, photochemistry, transfer to photosystem I - K(PI), k(R) rate constants for photoinhibition and recovery - PFD photon flux density - PSI, II photosystem I, II - _i photon yield of photosynthesis (incident light)     

Effects of ambient temperature and altitude on ventilation and gas exchange in deer mice (Peromyscus maniculatus)

Summary The effects of different ambient temperatures (T _a) on gas exchange and ventilation in deer mice (Peromyscus maniculatus) were determined after acclimation to low and high altitude (340 and 3,800 m).At both low and high altitude, oxygen consumption ( ) decreased with increasingT _a atT _a from –10 to 30 °C. The was 15–20% smaller at high altitude than at low altitude atT _a below 30 °C.Increased atT _a below thermoneutrality was supported by increased minute volume ( ) at both low and high altitude. At mostT _a, the change in was primarily a function of changing respiration frequency (f); relatively little change occurred in tidal volume (V _T) or oxygen extraction efficiency (O₂EE). AtT _a=0 °C and below at high altitude, was constant due to decliningV _T and O₂EE increased in order to maintain high .At high altitude, (BTP) was 30–40% higher at a givenT _a than at low altitude, except atT _a below 10 °C. The increased at high altitude was due primarily to a proportional increase inf, which attained mean values of 450–500 breaths/min atT _a below 0 °C. The (STP) was equivalent at high and low altitude atT _a of 10 °C and above. At lowerT _a, (STPD) was larger at low altitude.At both altitudes, respiratory heat loss was a small fraction (<10%) of metabolic heat production, except at highT _a (20–30 °C).Abbreviations EHL evaporative heat loss - f respiration frequency - HL _a heat loss from warming tidal air - HL _e evaporative heat loss in tidal air - HL total respiratory heat loss - MHP metabolic heat production - O ₂ EE oxygen extraction efficiency - RQ respiratory quotient - T _a ambient temperature - T _b body temperatureT _lc lower critical temperature - carbon dioxide production - evaporative water loss - oxygen consumption - minute volume - V _T tidal volume     

Type I procollagen in the severe non-lethal form of osteogenesis imperfecta

Summary We have screened type I procollagen synthesized in vitro by skin fibroblasts from several patients with the severe non-lethal form of osteogenesis imperfecta. Cells from one patient synthesized and secreted both normal and a larger amount of abnormal type I procollagen. The abnormal alpha chains are larger in size due to post-translational overmodifications involving the whole triple helical domain. Abnormal collagen heterotrimers had a melting temperature 2.5°–3°C lower than normal ones or from controls. Chemical analysis of collagen in the medium showed a greater degree of both lysyl hydroxylation and hydroxylysyl glycosylation, the major increase in molecular mass of overmodified alpha chains being due to the higher hydroxylysine-bound hexose content. The proband's cells modify proteoglycan metabolism and mineral proband's cells modify proteoglycan metabolism and mineral crystals form in the dermis, possibly a response to abnormal collagen-proteoglycan interactions. These findings can be explained by a small defect in the product of one allele for pro-1(I) chains: three-quarters of the synthesized type I procollagen molecules are composed of trimers containing one or two chains defective near the C-terminus of the triple helix or in the C-propeptide. The data obtained for this patient confirmed that the severity of clinical manifestations in osteogenesis imperfecta strongly depends on the location and nature of the mutations, and that the phenotype could be a consequence of a collagen defect(s) and its influence on collagen-collagen interactions and collagen interactions with other connective tissue components.     

Growth of the thermotolerant yeast,Candida valida,on ethanol: Dependences of maximal growth rate and cell biomass yield on temperature

Summary Growth of Candida valida on ethanol in pH-auxostat and chemostat has been studied. Maximal growth rate, _m, and cell biomass yield, Y _s, display the Arrhenius dependence on temperature within the ranges 18°–30° C and 30°–36° C and an abrupt fall above 36° C. The temprature dependence of both parameters has breaks at 30° C and 36° C. Activation energies have been measured for both _m and Y _s. The reason for a weaker effect of temperature on Y _s than on _m is discussed.     

Thermoregulation,gas exchange,and ventilation in Adelie penguins (Pygoscelis adeliae)

Summary Adelie penguins (Pygoscelis adeliae) experience a wide range of ambient temperatures (T _a) in their natural habitat. We examined body temperature (T _b), oxygen consumption ( ), carbon dioxide production ( ), evaporative water loss ( ), and ventilation atT _a from –20 to 30 °C. Body temperature did not change significantly between –20 and 20°C (meanT _b=39.3°C).T _b increased slightly to 40.1 °C atT _a=30°C. Both and were constant and minimal atT _a between –10 and 20°C, with only minor increases at –20 and 30°C. The minimal of adult penguins (mean mass 4.007 kg) was 0.0112 ml/[g·min], equivalent to a metabolic heat production (MHP) of 14.9 Watt. The respiratory exchange ratio was approximately 0.7 at allT _a. Values of were low at lowT _a, but increased to 0.21 g/min at 30°C, equivalent to 0.3% of body mass/h. Dry conductance increased 3.5-fold between –20 and 30°C. Evaporative heat loss (EHL) comprised about 5% of MHP at lowT _a, rising to 47% of MHP atT _a=30°C. The means of ventilation parameters (tidal volume [VT], respiration frequency [f], minute volume [I], and oxygen extraction [ ]) were fairly stable between –20 and 10°C (VT did not change significantly over the entireT _a range). However, there was considerable inter- and intra-individual variation in ventilation patterns. AtT _a=20–30°C,f increased 7-fold over the minimal value of 7.6 breaths/min, and I showed a similar change. fell from 28–35% at lowT _a to 6% atT _a=30°C.Abbreviations C thermal conductance - EHL evaporative heat loss - oxygen extraction - f respiratory frequency - MHP metabolic heat production - evaporative water loss - LCT lower critical temperature - RE respiratory exchange ratio - T _a ambient temperature - T _b body temperature - rate of oxygen consumption - rate of carbon dioxide production - I inspiratory minute volume - VT tidal volume     

Solar heat gain in a desert rodent: unexpected increases with wind speed and implications for estimating the heat balance of free-living animals

We quantified metabolic power consumption as a function of wind speed in the presence and absence of simulated solar radiation in rock squirrels, Spermophilus variegatus, a diurnal rodent inhabiting arid regions of Mexico and the western United States. In the absence of solar radiation, metabolic rate increased 2.2-fold as wind speed increased from 0.25 to 4.0 m·s^-1. Whole-body thermal resistance declined 56% as wind speed increased over this range, indicating that body insulation in this species is much more sensitive to wind disruption than in other mammals. In the presence of 950 W·m^-2 simulated solar radiation, metabolic rate increased 2.3-fold as wind speed was elevated from 0.25 to 4.0 m·s^-1. Solar heat gain, calculated as the reduction in metabolic heat production associated with the addition of solar radiation, increased with wind speed from 1.26 mW·g^-1 at 0.25 m·s^-1 to 2.92 mW·g^-1 at 4.0 m·s^-1. This increase is opposite to theoretical expectations. Both the unexpected increase in solar heat gain at elevated wind speeds and the large-scale reduction of coat insulation suggests that assumptions often used in heat-transfer analyses of animals can produce important errors.Abbreviations absorptivity of coat to solar radiation - kinematic viscosity of air (mm²·s^-1) - reflectivity of coat to solar radiation - a r _B expected at zero wind speed (s·m^-1) - A _P projected surface area of animal on plane perpendicular to solar beam (cm²) - A _SKIN skin surface area (cm²) - b Coefficient describing change in r _B with change in square-root of wind speed (s^1.5·m^1.5) - d hair diameter (m) - d characteristic dimension of animal (m) - D _H thermal diffusivity of air (m²·s^-1) - E evaporative heat loss (W·m^-2) - I probability per unit coat depth that photon will strike hair - k constant equalling 1200 J·m^-3·°C^-1 - l _C coat depth m) - l _H hair length (m) - M metabolic rate (W·m^-2) - n density of hairs of skin (m^-2) - Q _A solar heat gain to animal (W·m^-2) - Q _I solar irradiance intercepted by animal (W·m^-2) - RQ respiratory quotient - r _A thermal resistance of boundary layer (s·m^-1) - r _B whole-body thermal resistance (s·m^-1) - r _E thermal resistance between animal surface and environment s·m^-1) - r _R radiative resistance (s·m^-1) - r _S sum of r _B and r _E at 0.25 m·s^-1 (s·m^-1) - r _T tissue thermal resistance s·m^-1) - T _AIR air temperature (°C) - T _B body temperature (°C) - T _E operative temperature of environment (°C) - T _ES standard operative temperature of environment (°C) - u wind speed (m·s^-1)