J Appl Physiol 94: 2350–2357, 2003.
First published February 21, 2003; 10.1152/japplphysiol.01107.2002.
Muscle temperature transients before, during, and after
exercise measured using an intramuscular multisensor probe
G. P. Kenny,1 F. D. Reardon,1 W. Zaleski,2
M. L. Reardon,2 F. Haman,3 and M. B. Ducharme1,4
1
Faculty of Health Sciences, School of Human Kinetics, 2Faculty of Medicine, and 3Faculty
of Sciences, University of Ottawa, Ottawa K1N 6N5; and 4Human Protection and Performance
Group, Defence Research and Development Canada-Toronto, Toronto, Ontario, Canada M3M 3B9
Submitted 3 December 2002; accepted in final form 7 February 2003
Kenny, G. P., F. D. Reardon, W. Zaleski, M. L. Rear-
don, F. Haman, and M. B. Ducharme. Muscle tempera-
ture transients before, during, and after exercise measured
using an intramuscular multisensor probe. J Appl Physiol 94:
2350–2357, 2003. First published February 21, 2003; 10.1152/
japplphysiol.01107.2002.—Seven subjects (1 woman) per-
formed an incremental isotonic test on a Kin-Com isokinetic
apparatus to determine their maximal oxygen consumption
during bilateral knee extensions (V̇O2 sp). A multisensor ther-
mal probe was inserted into the left vastus medialis (middi-
aphysis) under ultrasound guidance. The deepest sensor (tip)
was located ⬃10 mm from the femur and deep femoral artery
(Tmu 10), with additional sensors located 15 (Tmu 25) and 30
mm (Tmu 40) from the tip. Esophageal temperature (Tes) was
measured as an index of core temperature. Subjects rested in
an upright seated position for 60 min in an ambient condition
of 22°C. They then performed 15 min of isolated bilateral
knee extensions (60% of V̇O2 sp) on a Kin-Com, followed by 60
min of recovery. Resting Tes was 36.80°C, whereas Tmu 10,
Tmu 25, and Tmu 40 were 36.14, 35.86, and 35.01°C, respec-
tively. Exercise resulted in a Tes increase of 0.55°C above
preexercise resting, whereas muscle temperature of the ex-
ercising leg increased by 2.00, 2.37, and 3.20°C for Tmu 10,
Tmu 25, and Tmu 40, respectively. Postexercise Tes showed a
rapid decrease followed by a prolonged sustained elevation
⬃0.3°C above resting. Muscle temperature decreased gradu-
ally over the course of recovery, with values remaining sig-
nificantly elevated by 0.92, 1.05, and 1.77°C for Tmu 10,
Tmu 25, and Tmu 40, respectively, at end of recovery (P ⬍ 0.05).
These results suggest that the transfer of residual heat from
previously active musculature may contribute to the sus-
tained elevation in postexercise Tes.
heat load; thermoregulation; hyperthermia; heat content;
heat balance
A NUMBER OF STUDIES HAVE EXAMINED muscle temperature
(Tmu) profiles for resting conditions under different
thermal conditions (4, 7, 8, 17, 20–22, 24, 27, 30, 31).
The study by Ducharme and Tikuisis (8), however, was
the only study to present a mean Tmu profile for a group
of subjects, as opposed to single-depth measurements.
Despite the large number of studies, there is still no
consistent description of resting Tmu profile. There also
Address for reprint requests and other correspondence: G. P.
Kenny, School of Human Kinetics, Univ. of Ottawa, Rm. 372, Mont-
petit Hall, P.O. Box 450 Station A, 125 Univ., Ottawa, Ontario,
Canada K1N 6N5 (E-mail: gkenny@uottawa.ca).
2350 8750-7587/03 $5.00 Copyright © 2003 the American Physiological Society
have been a number of studies that have reported Tmu
response during exercise (1, 2, 5, 6, 19, 23–27), of which
the study by Saltin et al. (25) seems to be the only one
to examine changes in Tmu profile (i.e., Tmu measured
at multiple depths). Although, these experiments were
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not designed to show the time course change in tissue
temperature gradients, their measurement of individ-
ual intramuscular temperature response did show
large variations in the temperature at the superficial,
mid-, and deep muscle sites. Furthermore, they showed
significant variation in the rates of temperature
change during muscle activity.
There are no studies that have examined changes in
Tmu during the postexercise period. Several have re-
ported postexercise Tmu response (1, 23, 25); however,
none has specifically addressed these responses. In
short, there remains a lack of information regarding
the kinetics of heat exchange between muscle and the
core of the body and within a given mass of muscle
tissue. This information is critical to our understand-
ing of the underlying mechanism responsible for the
sustained, postexercise elevation in core temperature.
In previous work, core temperature has been shown to
remain elevated by ⬃0.4°C for a prolonged period after
cessation of exercise (16, 28) performed in different
thermal environments. However, the actual mecha-
nism responsible for this increase in core temperature
remains unresolved.
Tissue temperature at any given time is ultimately
determined by the relative rates of heat production and
heat loss. For example, regional Tmu at any point in
time is the result of regional differences in metabolic
rate (9), conductive heat loss to adjacent tissue (9, 10),
and deep and peripheral convective blood flow (9, 29).
As such, it would be expected that both regional tem-
perature profile and the rate of temperature change
would differ during resting, exercise, and postexercise
recovery. The following study was designed to measure
intramuscular temperature profile during rest, exer-
cise, and postexercise recovery. In contrast to the find-
ings of previous studies, we hypothesized that the
tissue temperature profile will be consistent among
The costs of publication of this article were defrayed in part by the
payment of page charges. The article must therefore be hereby
marked ‘‘advertisement’’ in accordance with 18 U.S.C. Section 1734
solely to indicate this fact.
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 TISSUE TEMPERATURE TRANSIENTS 2351

subjects as the probe position is standardized within
the muscle of all subjects. Furthermore, in conjunction
with a postexercise decrease in heat loss, subsequent to
a decrease in skin blood perfusion, and an attenuation
of sudomotor activity during the postexercise recovery
(14), we hypothesized that convective heat transfer
between muscle and core will significantly influence
postexercise core temperature response.
METHODS
Subjects. Subsequent to approval of the project by the
University Human Research Ethics Committee, seven
healthy subjects (6 men, 1 woman) consented to participate
in the study. Mean values (⫾SD) of the subjects’ age, height,
body mass, maximal oxygen consumption during bilateral
concentric knee extensions (V̇O2 sp), and body fat content
were 24 ⫾ 5 yr, 1.8 ⫾ 0.5 m, 85.6 ⫾ 6.1 kg, 2.1 ⫾ 0.9 l/min,
and 10.9 ⫾ 2.3%, respectively.
In each trial, esophageal temperature (Tes) was measured
final position of the probe was verified by using the ultra-
sound imaging. The average depth of the probe from the
surface was 47.2 mm and within 11.0 mm of both the deep
femoral artery and femur. The probe assembly was secured
to the skin with sterile, waterproof transparent dressing (3M
1622W Tegaderm transparent dressing) and tape (total sur-
face coverage ⬃25 cm2). The Tegaderm transparent dressing
consists of a thin polyurethane membrane coated with a layer
of an acrylic adhesive. The dressing, which is permeable to
both water vapor and oxygen, is impermeable to microorgan-
isms, and, once in position, it provides an effective barrier to
external contamination.
The temperature probe was a sterile Teflon-coated multi-
sensor probe (model IT-17:3, Physitemp Instruments; ther-
mal constant of 0.25 s). Each probe had three thermocouples:
one positioned at the tip, one at 15 mm, and the third at 30
mm from the tip. The deepest temperature sensor (tip) was
located ⬃10 mm from the femur and deep femoral artery
(Tmu 10), with two sensors located at 15 (Tmu 25) and 30 mm
(Tmu 40) from the tip (Table 1). The internal position of the
temperature sensor relative to the skin surface was calcu-

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by using a thermocouple temperature probe (Mallinckrodt
Medical) inserted through a nostril, into the esophagus, to
the level of the heart. Regional Tmu of the vastus medialis
was measured by using a flexible multithermocouple temper-
ature probe (Physitemp Instruments, Clifton, NJ; model IT-
17:3) inserted into the vastus medialis. Ultrasound imaging
was used to discern the best perpendicular insertion tract
and subsequently to place the probe at a position 10 mm and
equidistant from the deep femoral artery and the femur. The
implant site was approximately midway between, and medial
to, a line joining the anterior superior iliac spine and the
superior aspect of the center of the patella.
With the use of aseptic technique and under ultrasound
guidance, the skin, subcutaneous tissue, and muscle were
anesthetized to a maximum depth of 50 mm by infiltrating
⬃2 ml of 1% lidocaine without epinephrine. The tip of this
25-gauge needle was placed at the proposed site for the deep
temperature probe. Under full ultrasound imaging and with
the use of the anesthetic needle as a guide, an 18-gauge,
50-mm polyethylene catheter (Cathlon and Critikon Canada,
Markham, Ontario) was then inserted into the anesthetized
tract to the required depth. The anesthetic needle and the
catheter stylet were then withdrawn, and the temperature
probe was inserted in the catheter shaft. When the probe was
fully inserted, the catheter was carefully withdrawn, leaving
the tip of the temperature probe ⬃10 mm from the femur and
deep femoral artery. Once the catheter was withdrawn, the
lated based on the ratio of the known depth of the probe
(radius r) from the skin surface measured by ultrasound
imaging and the radius of the thigh (rsk). Thus r/rsk is the
relative radius (8). Although it was not possible to verify the
final position of the probe after the completion of the exper-
imental trial, the length of the probe within the limb tissue
was measured during the removal of the probe. The depth of
the probe was verified with the preexperiment depth, as
determined by ultrasound imaging.
Skin temperature was monitored at 12 sites by using type
T thermocouples integrated into heat-flow sensors (Concept
Engineering, Old Saybrook, CT). The area-weighed mean
skin temperature (T៮ sk) and heat flux (H៮ Fsk) were calculated
by assigning the following regional percentages: 6% head, 9%
upper arm, 6% forearm, 2% finger, 19% chest, 9.5% upper
back, 9.5% lower back, 10% anterior thigh, 10% posterior
thigh, 9.5% anterior calf, and 9.5% posterior calf (12). Tem-
perature and heat flux data were collected and digitized
(Hewlett Packard data-acquisition module, model 3497A) at
5-s intervals and simultaneously displayed and recorded in
spreadsheet format on a hard disk (Hewlett Packard, model
PC-312, 9000).
Oxygen consumption (V̇O2) was determined by open-circuit
analysis by using an automated gas-collection system (Quin-
ton Instrument, Seattle, WA; model Q-Plex 1 cardiopulmo-
nary exercise system). Skin blood flow was measured by

Table 1. Mean (⫾SD) and individual data relating to the placement of the intramuscular multisensor thermal
probe of the upper leg
Insertion Depth From Skin
Surface§
Subject Thigh Thigh Inguinal Base of Medial Probe Tip to Probe Tip
No. Circumference* Length Crease† Femur† Deviation‡ Tmu 10 Tmu 25 Tmu 40 Femoral Artery§ to Femur§

1 58 52 24 28 5.5 4.7 3.2 1.7 0.9 1.0

2 55 47 22 25 5.4 4.8 3.3 1.8 1.0 1.0
3 65 48 21 27 5.9 5.0 3.5 2.0 1.2 1.0
4 56 46 20 26 5.1 4.4 2.9 1.4 1.3 0.9
5 56 47 23 24 6.1 4.7 3.2 1.7 1.0 1.3
6 67 44 21 23 5.7 5.3 3.8 2.3 1.3 1.3
7 61 45 22 25 5.8 5.0 3.5 2.0 1.1 1.2
Means 60 47 22 25 5.6 4.8 3.3 1.8 1.1 1.1
SD 8 4 1 2 0.2 0.3 0.3 0.3 0.2 0.1
Values are means ⫾ SD in cm. Tmu 10, Tmu 25, and Tmu 40, temperature sensor tip 10, 15, and 30 mm, respectively, from the femur and deep
femoral artery. * Values represent the circumference of thigh at level of probe insertion. †Values represent the distance to probe insertion in
relation to long axis of the thigh. ‡ Medial deviation from the thigh long axis. § Values determined by ultrasound imaging.

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2352 TISSUE TEMPERATURE TRANSIENTS
laser-Doppler velocimetry (PeriFlux System 5000, main con-
trol unit; PF5010 LDPM, operating unit; Perimed, Stock-
holm, Sweden) from the left midanterior forearm. The laser-
Doppler flow probes (PR 401 angled probe, Perimed) were
taped to cleaned skin, in an area that superficially did not
appear to be highly vascular and from where consistent
readings were noted (18). Sweat rate was estimated from a
5.0-cm2 ventilated capsule placed on the upper back. Anhy-
drous compressed air was passed through the capsule over
the skin surface at a rate of 1 l/min. Water content of the
effluent air was measured at known barometric pressure by
using the readings from an Omega HX93 humidity and
temperature sensor (Omega Engineering, Stamford, CT).
Sweat rate was calculated from the product of the difference
in water content between effluent and influent air and the
flow rate. This value was normalized for the skin surface area
under the capsule and expressed in milligrams per minute
per centimeter squared.
Subjects performed an incremental isotonic test (constant
angular velocity, increases in force output) on the Kin-Com
isokinetic apparatus to determine their V̇O2 sp. The exercise
consisted of bilateral, concentric knee extension over a range
of 70° from perpendicular, with the subject sitting (hip angle
between 90 and 110°) and the long axis of the thigh in the
horizontal plane. The force output was increased by 15 N
every 2 min until fatigue, whereas the angular velocity was
maintained at 58.3°/s throughout the test. The results of the
test were used to establish the work rate for the experimental
trial. The experimental trial was conducted in the morning
after a 24-h period without heavy or prolonged physical
activity. On arrival at the laboratory at 0800, subjects were
appropriately instrumented. Subjects then rested in a semi-
recumbent position for 60 min at an ambient temperature of
22°C, of which the final 20 min were recorded as represen-
tative of the baseline resting values. At 2 min before exercise,
the subjects were secured to the Kin-Com isokinetic appara-
tus at the level of the torso and ankles. Subjects then per-
formed 15 min of exercise, as described above, consisting of
bilateral, concentric knee extension over a range of 70° from
perpendicular against a dynamic resistance sufficient to
elicit a heat load of 4.78 kJ/kg. Exercise was followed by 60
min of seated rest.
The total energy expended (Mtotal) as a result of exercise,
during the period from onset of exercise until the time at
which V̇O2 returned to preexercise values, was calculated
from the sum of the energy expended by using the following
equation (expressed in kJ)
M total ⫽ 兺 Ṁ Ex/rest
冊冉 冊册冎
兺 再 V̇ 冋冉 (RER ⫺ 0.7) (1 ⫺ RER)
⫽ O 2䡠 ec ⫹ ef
0.3 0.3
where ṀEx/rest is the rate of energy expenditure during exer-
cise and recovery, RER is the respiratory exchange ratio, ec is
the caloric equivalent (in kJ/l O2) for carbohydrates, and ef is
the caloric equivalent (in kJ/l O2) for fat.
The minute values were summed for the entire period as
described above.
The mechanical work (W) done during each contraction of
the exercise phase was measured and recorded by using the
Kin-Com isokinetic machine. This was calculated from the
force exerted and the angular displacement during the knee
extension
W ⫽ T␪
where T is rotational force or torque and ␪ is the angular
displacement.
J Appl Physiol • VOL
The total work done (Wtotal) was the sum of the work
accomplished during each of the contractions during the 15
min of exercise.
Mechanical efficiency (ME) was defined as the Wtotal com-
pleted during the 15-min exercise period divided by the Mtotal
minus the energy expended under resting conditions (Mrest)
(Mtotal ⫺ Mrest). Thus
W total
ME ⫽ (3)
M total ⫺ M rest
The Mrest was calculated from the average rate of V̇O2 during
the 5 min preceding the exercise bout. These values were
calculated and expressed in kilojoules by using the aforemen-
tioned equation.
The total heat load generated by the exercise (HLex) for
each subject was calculated by subtracting the Mrest and the
energy equivalent of the total mechanical work done (Wtotal)
from the Mtotal. Values are expressed in kilojoules
HL ex ⫽ M total ⫺ (M rest ⫹ W total) (4)
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The total dry heat loss by radiation, conduction, and convec-
tion from the skin surface during exercise (Hsk ex) and during
recovery (Hsk rec) was estimated by subtracting the area-
weighted H ៮ Fsk (as above) corrected for body surface area (AD)
during rest from those values recorded during exercise and
recovery, respectively. Thus
៮ F sk 䡠 AD)ex ⫺ (H
H sk ex ⫽ (H ៮ Fsk 䡠 AD)rest (5)
and
៮ F sk 䡠 AD)rec ⫺ (H
H sk rec ⫽ (H ៮ Fsk 䡠 AD)rest (6)
where Hsk ex and Hsk rec are the total dry heat lost during the
15-min exercise and 60-min recovery periods, respectively.
Statistical analyses for Tes, Tmu, T៮ sk, and H
៮ Fsk were per-
formed by ANOVA for repeated measures to compare values
for preexercise, end-exercise, and at 10-min intervals during
postexercise recovery. Data are presented as means ⫾ SD.
RESULTS
Baseline Tes and T៮ sk were 36.80 ⫾ 0.30 and 31.66 ⫾
0.89°C, respectively. Resting Tmu was significantly
lower than Tes (i.e., 36.14 ⫾ 0.29, 35.86 ⫾ 0.31, and
35.01 ⫾ 0.33°C for Tmu 10, Tmu 25, and Tmu 40, respec-
tively) (Fig. 1). It should be noted that the increase in
muscle tissue temperature before the onset of exercise
was likely due to the preparation of the subject for the
exercise portion of the experimental trial.
(1)
The Tmu profiles, expressed as a function of the
position of the placement of the temperature relative to
the radius of the thigh (r/rsk), show a parabolic profile
for mean resting tissue temperature profile (Fig. 2).
This parabolic form of Tmu profile was observed consis-
tently in the data of all seven subjects (Fig. 3). During
resting, the deep (36.14°C) and mid-Tmu (35.86°C) were
significantly different from superficial (35.01°C) Tmu.
As depicted in Fig. 4, the greatest tissue temperature
difference (1.13°C) was between the deep and superfi-
cial sections of the muscle, with a 0.84°C temperature
gradient between the mid- and superficial muscle.
(2) Mean tissue temperature difference between deep and
midmuscle was only 0.28°C. Furthermore, the muscle-
to-core temperature gradient was equal to ⫺0.66,
94 • JUNE 2003 • www.jap.org
 TISSUE TEMPERATURE TRANSIENTS 2353

Fig. 1. Mean (⫾SE) muscle [temperature sensor located 10 mm

(Tmu 10; 䊐), 15 mm (Tmu 25; E), and 30 mm (Tmu 40; ƒ) from femur and
deep femoral artery] and esophageal ({) temperature response dur-

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ing rest, exercise (Ex), and postexercise recovery. Vertical dotted
lines represent the start (time ⫽ 0 min) and end (time ⫽ 15 min) of
exercise. * Significant difference from baseline resting values, P ⬍
0.05. 1 Value for esophageal temperature not significantly elevated
from baseline.

⫺0.94, and ⫺1.79°C in relation to Tmu 10, Tmu 25, and

Tmu 40, respectively (P ⬍ 0.05).
Exercise tissue temperature response. After the onset
of exercise, Tes increased gradually, reaching a maxi-
mum rate of increase of 0.05 ⫾ 0.02°C/min between 6
and 9 min of exercise, and, subsequently, the rate
decreased over the balance of the exercise period (Table
2). In contrast, Tmu at all measured points increased
rapidly during the initial period of exercise followed by
a gradual reduction in the rate over the balance of the
exercise period. Superficial muscle (Tmu 40) showed the
greatest rate of temperature increase (0.61 ⫾ 0.19°C/
min). This value was significantly higher than the rate

Fig. 3. Mean (⫾SE) (F) and individual muscle temperature profiles

during resting (A), end-exercise (B), and at 60 min postexercise
resting (C). Note: individual subjects are represented by different
symbols, and these symbols are the same for each time period in
A–C. § Significantly different from superficial muscle, P ⬍ 0.05.

measured in deep muscle (0.22 ⫾ 0.09°C/min). After

Fig. 2. Mean (⫾SE) muscle temperature profile during baseline
resting (F), Ex (䊐), end-exercise (End Ex; ■), and postexercise (post-
Ex; ‚) recovery at selected periods as a function of the placement of
the temperature sensors relative to the radius of the thigh. r, Radius
(cm); rsk, radius of the thigh (cm); r/rsk, relative radius.
J Appl Physiol • VOL 94 • JUNE 2003 •
the initial 3 min of exercise, the rate of Tmu change
decreased gradually and was similar at all three intra-
muscular sites until the end of exercise. Exercise re-
sulted in a 0.55°C (end-exercise Tes of 37.35°C) in-
crease in core temperature above baseline resting val-
ues. In contrast, Tmu increased by 2.09, 2.37, and
3.20°C above baseline resting values for Tmu 10, Tmu 25,
and Tmu 40 measurement points, respectively, with
end-exercise values similar at all three measured sites
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2354 TISSUE TEMPERATURE TRANSIENTS

Fig. 5. Mean (⫾SD) heat load (J) and dry heat loss (ß) responses

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during baseline resting, exercise, and postexercise recovery. Vertical
dotted lines represent the start (time ⫽ 0 min) and end (time ⫽ 15
min) of exercise.

Fig. 4. Mean (⫾SE) core-to-muscle temperature gradient (E, deep
muscle to core; 䊐, midmuscle to core; ‚, superficial muscle to core) (A)
and intramuscular temperature gradients (✳, deep muscle to mid-
muscle; {, deep muscle to superficial muscle; ƒ, midmuscle to superficial
muscle) (B). Vertical dotted lines represent the start (time ⫽ 0 min) and
end (time ⫽ 15 min) of exercise. P ⬍ 0.05: ‡significantly different from
the deep to midmuscle temperature gradient; §significantly different
from baseline resting; *superficial-to-core temperature gradient signif-
icantly different from deep-to-core temperature gradient.
(i.e., 38.23, 38.23, and 38.21°C for Tmu 10, Tmu 25, and
៮ sk increased continuously during
Tmu 40, respectively). T
exercise to an end-exercise value that was significantly
elevated above baseline rest (32.79°C, P ⬍ 0.05). The
increase in T៮ sk was paralleled by an increase in non-
evaporative heat loss (i.e., Fig. 5). Forearm skin blood
flow increased continuously during the course of the
exercise.
As depicted in Fig. 2, the tissue temperature profile
evolved from a parabolic form typical of resting to a
linear profile during the early stages of exercise. By the
end of exercise, Tmu values across the radial axis were
homogenous. As such, the large temperature gradient
between the deep and superficial muscle was rapidly
reduced such that, by the end of exercise, the temper-
ature at all sites was similar. Furthermore, the mus-
cle-to-core temperature gradient was reversed from

Table 2. Mean (⫾SD) rate of change of esophageal and muscle (Tmu 10, Tmu 25, and Tmu 40) temperatures
during exercise and postexercise recovery
Muscle Temperature
Esophageal
Temperature Tmu 10 Tmu 25 Tmu 40

Exercise
0–3 min 0.022 ⫾ 0.008 0.220 ⫾ 0.091 0.331 ⫾ 0.108 0.547 ⫾ 0.195
3–6 min 0.041 ⫾ 0.009 0.134 ⫾ 0.071 0.137 ⫾ 0.084 0.245 ⫾ 0.099
6–9 min 0.051 ⫾ 0.016 0.059 ⫾ 0.017 0.069 ⫾ 0.023 0.068 ⫾ 0.026
9–12 min 0.038 ⫾ 0.011 0.054 ⫾ 0.015 0.046 ⫾ 0.018 0.056 ⫾ 0.020
12–15 min 0.027 ⫾ 0.009 0.042 ⫾ 0.016 0.043 ⫾ 0.020 0.054 ⫾ 0.021
Postexercise
0–5 min ⫺0.035 ⫾ 0.010 ⫺0.107 ⫾ 0.046 ⫺0.085 ⫾ 0.035 ⫺0.093 ⫾ 0.033
5–10 min ⫺0.006 ⫾ 0.002 ⫺0.034 ⫾ 0.012 ⫺0.041 ⫾ 0.017 ⫺0.045 ⫾ 0.029
10–20 min ⫹0.001 ⫾ 0.007 ⫺0.022 ⫾ 0.010 ⫺0.028 ⫾ 0.009 ⫺0.028 ⫾ 0.011
20–30 min ⫹0.001 ⫾ 0.006 ⫺0.014 ⫾ 0.009 ⫺0.017 ⫾ 0.008 ⫺0.018 ⫾ 0.010
30–40 min ⫺0.003 ⫾ 0.007 ⫺0.005 ⫾ 0.003 ⫺0.006 ⫾ 0.008 ⫺0.010 ⫾ 0.009
40–50 min ⫺0.003 ⫾ 0.002 ⫺0.002 ⫾ 0.001 ⫺0.006 ⫾ 0.004 ⫺0.009 ⫾ 0.007*
50–60 min ⫺0.001 ⫾ 0.005 ⫺0.002 ⫾ 0.004 ⫺0.006 ⫾ 0.005 ⫺0.009 ⫾ 0.007*
Values are means ⫾ SD in °C/min. * Significant difference from deep muscle, Tmu 10 (P ⬍ 0.05).

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 TISSUE TEMPERATURE TRANSIENTS
resting, such that Tmu values at all sites were signifi-
cantly elevated above Tes by 0.90, 0.90, and 0.89°C for
Tmu 10, Tmu 25, and Tmu 40, respectively (P ⬍ 0.05).
Postexercise tissue temperature response. Tes de-
creased rapidly (⫺0.04°C/min) during the initial min-
utes after the cessation of exercise (Table 2), after
which there was a rapid decrease in the rate of tem-
perature decrease to negligible values. At ⬃5 min of
recovery, Tes reached an elevated value 0.35°C above
baseline resting values (P ⬍ 0.05). For the duration of
recovery, the rate of decrease of Tes remained at
⬃0.001°C/min. Tes decreased to 37.11°C by the end of
the 60-min recovery period (⬃0.3°C above baseline).
Tmu showed a similar high rate of temperature de-
crease during the initial 5 min of exercise recovery,
although the rates were ⬃2 to 2.7 times greater than
the rate measured for Tes. Unlike the response in Tes,
Tmu for all measured sites decreased continuously dur-
ing the initial 30 min of recovery. However, the rates of
Tmu decay were reduced for the duration of recovery. In
the final 15 min of recovery, superficial muscle demon-
strated an elevated rate of temperature decrease above
deep muscle (P ⬍ 0.05). Muscle tissue temperature at
the end of the postexercise recovery period remained
significantly elevated above baseline resting values
by 0.92, 1.05, and 1.77°C for Tmu 10, Tmu 25, and Tmu 40,
respectively (P ⬍0.05) (Fig. 1). T៮ sk and whole body
nonevaporative heat loss decreased to baseline resting
values within ⬃20–25 min of recovery. Similarly, fore-
arm skin blood flow decreased to baseline resting val-
ues within 10 min of the termination of the exercise. In
contrast, both thigh nonevaporative heat loss and
thigh skin temperature remained significantly ele-
vated from preexercise values for the duration of the
recovery period (P ⬍ 0.05).
The temperature gradient between the different
depths of muscle remained relatively unchanged dur-
ing the postexercise recovery period, despite a slow
decay in Tmu. The muscle-to-core temperature gradient
decreased gradually over the course of the recovery. At
⬃25 min into recovery, Tmu at all depths achieved
similar values to that of Tes. For the duration of the
recovery period, the muscle-to-core temperature gradi-
ent was increased, with superficial muscle demonstrat-
ing the largest temperature gradient by the end of
recovery (⬃0.3°C; P ⬍ 0.05) compared with that tem-
perature gradient between deep muscle and core. Of
note, the deep muscle-to-core temperature gradient
remained relatively unchanged for the duration of the
recovery period (⬃0.02°C).
Heat load and heat loss response. The workload re-
sistance was adjusted for each subject, according to the
individual heat load and mechanical efficiency of the
leg-extension exercise. The mechanical efficiency var-
ied from 5.98 to 15.96%, whereas the average for the
group was 9.93 ⫾ 1.32%. The average actual workload
was one at which the heat production was 4.78 ⫾ 0.38
kJ/kg. The Mtotal and Wtotal were 585.87 ⫾ 53.72 and
41.48 ⫾ 4.68 kJ, respectively. Thus the average heat
load generated as a result of the exercise was 391.73 ⫾
38.93 kJ. The average total dry heat loss during the
J Appl Physiol • VOL
2355
exercise period was 15.93 ⫾ 5.98 kJ, whereas, during
the 60 min of postexercise recovery, this value was
37.90 ⫾ 18.80 kJ.
The kinetics of heat load generation at rest, over the
15 min of exercise, and over the first 10 min of recovery
are presented in Fig. 5. The corresponding evolution of
dry heat loss is also shown. Thus during the 5 min
preceding exercise, the dry heat loss defined relative to
heat load, that is, minus the resting levels, was essen-
tially zero. During exercise, the dry heat loss increased
at a rate of 0.14 kJ/min to a maximal level of 2.19
kJ/min after 15 min of exercise. At cessation of exer-
cise, the sensitive heat loss dropped exponentially to a
level ⬃1.0 kJ/min above initial resting values and
remained elevated for the next 10 min. The heat pro-
duction, on the other hand, increased to 17.08 kJ/min
after 2 min of exercise and continued to rise at a rate of
⬃0.78 kJ/min for the next 13 min to a maximum heat
production of 27.71 kJ/min. Immediately postexercise,
Downloaded from http://jap.physiology.org/ by 10.220.32.246 on September 30, 2017
the heat load returned exponentially to resting levels
within 5 min.
DISCUSSION
In this study, an attempt was made to specifically
evaluate the kinetics of heat exchange in muscle tissue
during and after exercise by using a multisensor ther-
mal probe positioned at a predetermined internal
marker. In contrast to previous studies, we observed
similar individual and group Tmu profiles during rest-
ing, exercise, and subsequent resting recovery. Fur-
thermore, we observed a sustained elevation of core
temperature for the duration of the recovery period
that is consistent with previous findings (16, 28). Spe-
cifically, Tes showed a rapid decrease in the first min-
utes of exercise recovery followed by a prolonged sus-
tained elevation of ⬃0.3°C. Of particular importance
was the observation that deep Tmu decreased during
the early stages of exercise recovery to values equal to
that of Tes. Subsequently, deep Tmu remained rela-
tively unchanged from Tes for the duration of recovery.
This supports the hypothesis that the postexercise
recovery of core temperature may be, to a large degree,
influenced by the residual heat load of muscle.
Tissue temperature response: resting. Different
shapes of limb temperature profile have been reported
for resting conditions during different thermal stresses
between individuals, whereas we noted a consistent
parabolic profile of Tmu in all subjects (4, 7, 8, 17,
20–22, 24, 27, 31). This could arise from inconsistency
in the specific placement of the internal probe in the
muscle. There is a wide variation in recorded muscle
tissue temperature due to the proximity of the probe to
the surface and to such structures as large arteries and
bone (10). This could have been a source of significant
variation in the recorded internal temperature. The
differences in the specific heat of these tissues, as well
as the differing blood flow and hence the convective
effect within these structures, influence the rates of
temperature change in adjacent regions of the muscle.
Therefore, consistent placement of the probe is critical.
94 • JUNE 2003 • www.jap.org
2356 TISSUE TEMPERATURE TRANSIENTS
Thus an attempt was made to minimize the variation
in temperature recording resulting from individual an-
atomic differences.
Tmu response: exercise. From the onset of exercise
until the late phases of exercise, there is a gradual
change in the Tmu profile from one parabolic in the
form seen at rest to a zero gradient or homogenous
temperature profile across the muscle. As shown in
Fig. 4, the large temperature gradient that existed
between the deep and midportions of the muscle and
the superficial muscle was rapidly eliminated during
the first 5 min of exercise. By the end of exercise, the
gradient between the different muscle depths was non-
existent, as temperatures across the radial axis of the
muscle became homogenous. This effect was not ob-
served earlier by Saltin et al. (24). In that case, it was
noted that both mid- and superficial Tmu remained
generally lower than deep Tmu. Similarly, superficial
Tmu remained lower than mid-Tmu, whereas the tem-
perature gradient between mid- and deep muscle
seemed to remain relatively constant throughout exer-
cise. The gradient between the superficial muscle in-
creased relative to that between mid- and deep muscle.
The different response to that observed in our study
may be attributed to a number of factors, which may
include the following: 1) differences in the measure-
ment site (i.e., vastus lateralis); 2) differences in the
muscle mass implicated in the exercise activity; 3)
differences in ambient conditions; and 4) difference in
work intensity. For example, it would be expected that
the temperature profile of the vastus lateralis would be
different from that of the medialis, as it is less affected
by the circulation in the femoral artery and vein. The
relative influence of convective heat exchange would be
considerably different between these muscles.
The high rate of Tmu increase in the early stages of
exercise is consistent with previous studies (1, 3, 24,
25). Aulick et al. (3) showed, at the beginning of exer-
cise, that heat gained by the leg (local metabolic heat
production plus vascular heat delivery from the vis-
cera) exceeded heat loss, and femoral vein blood tem-
perature rose rapidly. In this study, the superficial
regions of the muscle demonstrated the largest rate of
Tmu increase (0.53°C/min) that was 1.4 and 1.6 times
the rate of temperature increase for the deep and
midmuscle, respectively. On the other hand, based on
visual observation of the work by Saltin and coworkers
(24), it would seem that the rate of temperature in-
crease was greater in deep muscle compared with su-
perficial muscle.
During the course of exercise, the muscle-to-core
temperature gradient increased progressively (Fig. 4),
from ⫺1.15°C at rest to ⫹0.90°C by the end of exercise.
Also, despite the rapid increase in muscle heat content
(as represented by increased Tmu) to values exceeding
that for core, the rate of temperature increase of core
remained consistently lower than that of muscle.
Therefore, this would suggest that the rate of heat
accumulation within the core region is attenuated to a
large degree by an increase in the rate of whole body
heat loss (i.e., evaporative and nonevaporative heat
J Appl Physiol • VOL
loss). For example, Aulick et al. (3) previously noted
that, as limb sweat rate, cutaneous blood flow, and
muscle-to-skin temperature differences increased dur-
ing exercise, the active leg became a more effective
vehicle for heat dissipation, and that femoral venous
temperature eventually reached a plateau during
steady state. Furthermore, Gisolfi and Robinson (11)
showed that much of the heat produced by active leg
muscles is rapidly transported to surface veins and
that this muscle heat is potentially lost across the leg
surface. In this study, muscle-to-skin temperature gra-
dient remained elevated during the course of the exer-
cise by ⬃5.2°C, and skin blood flow and sweat rate
increased gradually during the course of the exercise.
Furthermore, it has previously been shown that, dur-
ing leg work, the inactive upper limbs also act as an
avenue for vascular heat loss from the central circula-
tion (15), which would further attenuate the increase
in core temperature.
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Tmu response: postexercise. Few studies have graph-
ically presented muscle tissue temperature response
during the postexercise period, and, even so, no specific
discussion was presented with regard to these data (1,
23, 25). It is clear in this study that, during the tran-
sition from exercise to postexercise resting recovery,
the Tmu profile across the radial axis of the muscle
remains constant (i.e., linear profile, see Figs. 2 and 3).
During the course of the 60-min recovery, all three
sites showed a similar rate of temperature change,
although superficial muscle showed a significantly
greater rate of temperature decrease toward the later
stages of recovery (P ⬍ 0.05).
Deep Tmu decreased during the early stages of exer-
cise recovery to values equal to that of Tes, after which
deep Tmu remained relatively unchanged from Tes for
the duration of recovery, with the deep muscle-to-core
temperature gradient no greater than ⬃0.02°C. The
lack of a difference in temperature gradient between
muscle and core suggests equilibration of heat distri-
bution within the body. Thus changes in surface heat
loss (i.e., evaporative and nonevaporative heat loss)
will change the rate of whole body cooling. Therefore,
the rate of core temperature decay is limited by the
rate at which heat is lost at the skin-air interface.
All Tmu values remained significantly elevated above
baseline resting values at the end of recovery. That was
paralleled by a significant increase in Tes of ⬃0.3°C
(P ⬍ 0.05). Aikas et al. (1) have shown a similar
postexercise increase in Tmu of the previously active
muscle, although Tes showed a rapid decrease to values
below baseline rest within a short period (⬃15 min)
after cessation of exercise. This is more difficult to
explain if one believes that the convective arterial flow
plays a major role in muscle cooling postexercise. Our
results are more consistent with those of Saltin et al.
(25), who, on the other hand, did observe a sustained
increase in postexercise Tes, whereas Tmu remained
significantly elevated above baseline resting values.
Thoden et al. (28) previously showed a prolonged
postexercise elevation (0.4–0.5°C) in Tes after dynamic
exercise. It was subsequently shown that an increase
94 • JUNE 2003 • www.jap.org
 TISSUE TEMPERATURE TRANSIENTS
in the postexercise hypotensive response, induced by
exercise of increasing intensity, was paralleled by an
increase (⬃0.4°C) in the magnitude of the postexercise
elevation in Tes (13). It was suggested that the postex-
ercise Tes response may be defined to a large degree by
the gradient between the periphery and core and that
the convective transfer of residual heat from previously
active musculature may contribute to the sustained
elevation in postexercise Tes. Our observation of a
prolonged elevation in Tmu at values elevated above
esophageal provides further evidence to support this
conclusion. Thus, in the absence of a postexercise in-
crease in heat loss response (14), Tes would remain
elevated as long as 1) the heat content of muscle
remains higher or equal to that of core; 2) the postex-
ercise hypotensive effect persists; or 3) a combination
of the two.
Summary. In the present study, exercise was per-
formed such that the dynamic resistance during the
bilateral knee-extension exercise was sufficient to elicit
a heat load of 4.78 kJ/kg. Thus it can be assumed that
the rate of heat production and accumulation in muscle
was comparable between subjects. Thus the variation
in Tmu profile observed between the transition from
rest to exercise and exercise to resting recovery was not
only the result of the change in metabolic heat produc-
tion but also the result of changes in the convective
heat transfer between blood and muscle and conduc-
tive heat transfer within the muscle and skin surface.
Furthermore, as with previous studies that have
shown that tissue heat content and compartmental
heat exchange are significantly influenced by convec-
tive heat exchange during rest (9) and exercise (15),
our findings suggest that postexercise core tempera-
ture response (and the rate of temperature decay) is
significantly influenced by convective heat transfer be-
tween muscle and core.
We acknowledge the technical support of Carolyn Proulx and
Normand Boulé.
This research was supported by the Natural Sciences and Engi-
neering Research Council of Canada (to G. P. Kenny).
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