Regulation of fat metabolism during resistance exercise

in sedentary lean and obese men

Michael J. Ormsbee, Myung Dong Choi, Justin K. Medlin, Gabriel H. Geyer,
Lauren H. Trantham, Gabriel S. Dubis and Robert C. Hickner
J Appl Physiol 106:1529-1537, 2009. First published 5 March 2009;
doi:10.1152/japplphysiol.91485.2008

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Regulation of fat metabolism during resistance exercise in sedentary lean
and obese men
Michael J. Ormsbee,1 Myung Dong Choi,1 Justin K. Medlin,1 Gabriel H. Geyer,1 Lauren H. Trantham,1
Gabriel S. Dubis,1 and Robert C. Hickner1,2
1
Human Performance Laboratory, Department of Exercise and Sport Science, College of Health and Human Performance;
and 2Department of Physiology, Brody School of Medicine, East Carolina University, Greenville, North Carolina
Submitted 13 November 2008; accepted in final form 3 March 2009
Ormsbee MJ, Choi MD, Medlin JK, Geyer GH, Trantham
LH, Dubis GS, Hickner RC. Regulation of fat metabolism dur-
ing resistance exercise in sedentary lean and obese men. J Appl
Physiol 106: 1529 –1537, 2009. First published March 5, 2009;
doi:10.1152/japplphysiol.91485.2008.—The effect of acute resistance
exercise (RE) on whole body energy expenditure (EE) and ␣2-
adrenergic receptor (␣2-AR) regulation of lipolysis in subcutaneous
abdominal adipose tissue (SCAAT) was determined in sedentary lean
(LN) and obese (OB) men. Lipolysis was monitored using microdi-
alysis in 10 LN [body mass index (BMI) 20.9 ⫾ 0.6] and 10 OB (BMI
36.2 ⫾ 2.7) men before, during, and for 24 h after RE. EE was
measured before and immediately after RE for 40 min. Changes in
interstitial glycerol were measured in SCAAT with three microdialy-
sis probes perfused with a control solution, phentolamine (␣2-AR
antagonist), or propranolol (␤-AR antagonist). EE and fat oxidation
(FOX) were significantly (P ⬍ 0.001) elevated immediately post-RE
compared with pre-RE in LN and OB subjects, with no differences
between groups. RE-induced increases in SCAAT glycerol concen-
trations from rest to peak exercise were greater in LN than in OB men
in the control (LN 142.1 ⫾ 30.8 vs. OB 65.4 ⫾ 14.2%, P ⫽ 0.03) and
phentolamine probes (LN 187.2 ⫾ 29.6 vs. OB 66.7 ⫾ 11.0%, P ⫽
0.002). Perfusion of propranolol had no effect on interstitial glycerol
concentrations over the time course of the experiment in either group.
Plasma insulin concentrations were significantly lower (P ⫽ 0.002)
and plasma growth hormone (GH) was significantly higher (P ⫽ 0.03)
in LN compared with OB men. The mechanism behind RE contrib-
uting to improved body composition may in part be due to enhanced
SCAAT lipolysis and improved EE and FOX in response to RE in LN
and OB men. The blunted SCAAT lipolytic response to RE in OB
compared with LN men is unrelated to RE-induced catecholamine
activation of the antilipolytic ␣2-ARs and may be due to depressed
GH in OB subjects.
microdialysis; lipolysis; fat oxidation
OBESITY AND DIABETES are at the forefront of health concerns in
the world today, and their prevalence continues to grow at an
alarming rate. However, evidence that resistance exercise (RE)
can improve body composition is mounting (8, 57), and it is
now recommended as a necessary component to any exercise
program (46).
RE has been shown to decrease resting respiratory exchange
ratio (RER) after exercise (6, 38, 44), indicating elevated
postexercise fat oxidation. Moreover, RE has been shown to
have a similar effect on 24-h EE and fat oxidation as a bout of
aerobic exercise (AE) with a comparable energy expenditure
(37). Recently, Chatzinikolaou et al. (8), using gluteal fat
biopsies, reported that 30 min of circuit-style RE increased
Address for reprint requests and other correspondence: R. C. Hickner,
Human Performance Laboratory, 363 Ward Sports Medicine Bldg., East
Carolina Univ., Greenville, NC 27858 (e-mail: hicknerr@ecu.edu).
http://www. jap.org 8750-7587/09 $8.00 Copyright © 2009 the American Physiological Society
J Appl Physiol 106: 1529–1537, 2009.
First published March 5, 2009; doi:10.1152/japplphysiol.91485.2008.
gluteal triacylglycerol lipase activity in lean (LN) and obese
(OB) men. Moreover, we recently demonstrated, using the
microdialysis technique, that an acute bout of RE results in
elevated subcutaneous abdominal adipose tissue (SCAAT)
lipolysis in healthy, young trained males (44). Whether RE has
the same lipolytic effect in sedentary LN or OB individuals
remains unknown.
The lipolytic response in fat cells is a function of the
Downloaded from jap.physiology.org on March 2, 2012
interplay between the opposing effects of the stimulatory
␤-ARs and the inhibitory ␣-ARs that are expressed in human
adipocytes (16, 29, 33, 34, 54). In vitro studies using isolated
human adipocytes show that ␤-adrenergic stimulation of lipol-
ysis is attenuated by activation of ␣2-ARs by epinephrine and
norepinephrine (2, 33, 34, 53). Catecholamines regulate lipol-
ysis in adipose tissue by interacting with both the ␣-ARs and
␤-ARs (2, 4, 10, 16, 47). Interestingly, it appears that the
antilipolytic ␣-ARs predominate in SCAAT, as opposed to
femoral and gluteal adipocytes, and that epinephrine has a
higher affinity for ␣-ARs than ␤-ARs (5). AE elevates cate-
cholamine release and activates ␣-ARs, resulting in inhibition
of stimulated lipolysis (53) in nonobese untrained subjects (9,
53, 55) and particularly in untrained obese subjects (52). This
result has been confirmed using isolated human fat cells (36).
The microdialysis technique allows for a relatively nonin-
vasive method to directly monitor lipolysis in SCAAT, and for
local perfusion of pharmacological agents such as ␣-AR inhib-
itors, to investigate the mechanistic properties of adipose tissue
lipolysis. In fact, pharmacologically blocking ␣-ARs with
phentolamine has been reported to elevate AE-induced lipoly-
sis, especially in OB subjects (52). Due to the paucity of data
regarding the effects of RE on lipolysis and fat oxidation and
the importance of the antilipolytic ␣2-AR pathway seen during
AE, further study investigating fat metabolism and the role of
the ␣2-AR pathway during and after RE is warranted.
Adipose tissue was studied because it is a major source of
endogenous fatty acids that are oxidized by skeletal muscle
over prolonged periods of inactivity and activity. In addition,
SCAAT appears to be more responsive to catecholamine-
stimulation than femoral and gluteal subcutaneous fat depots
(2, 5, 35, 58). Wahrenberg et al. (58) concluded that the
lipolytic effect of catecholamines was four- to fivefold more
marked in abdominal adipocytes than in other depots. Further-
more, android obesity is common among men and is linked to
many chronic diseases; therefore it is the critical fat depot to
investigate.
Therefore, the purpose of the present investigation was to
determine the effect of acute RE on lipolysis in SCAAT and
the role of ␣- and ␤-ARs in the regulation of lipolysis in
sedentary LN and OB men. In addition, the effect of RE on
1529
1530 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY
whole body EE and fat oxidation was explored. We hypothe-
sized that 1) strenuous whole body RE would increase lipoly-
sis, as measured with microdialysis, and whole body fat oxi-
dation immediately following RE, but to a greater extent in
LN than OB men in both cases; and 2) pharmacologically
blocking the ␣2-ARs in SCAAT with phentolamine would
increase lipolysis to a greater extent in OB than LN men in
response to RE.
MATERIALS AND METHODS
Participants. Ten lean [LN; body mass index (BMI) 20.9 ⫾ 0.6
kg/m2; body fat%, 13.6 ⫾ 1.5] and 10 obese (OB; BMI 36.2 ⫾ 2.7
kg/m2, body fat%, 37.8 ⫾ 2.2) sedentary (exercise ⬍2 days/wk, ⬍30
min/day) men (18 – 40 yr old) were recruited for this study. All
subjects were free of existing acute or chronic illness, known cardio-
vascular or metabolic disease, did not take any medications or dietary
supplements, and were nonsmokers. A preparticipation health history
and physical activity questionnaire were used to screen all volunteers
for inclusion in this study. All participants were informed both orally
and in writing of the purpose, risks, and benefits of the research and
gave their written informed consent to participate before beginning
the investigation. This study was approved by the East Carolina
University Medical Center Institutional Review Board. Subject char-
acteristics are presented in Table 1.
Study design. Participants reported to the Human Performance
Laboratory at East Carolina University on two separate occasions. The
first visit was used to gather baseline information including height,
weight, body composition, waist and hip circumferences, and 10-
repetition maximum (10RM) lifts. Informed consent was obtained
from all participants before any data collection.
Before the second laboratory visit, all participants abstained from
vigorous activity, alcohol, and caffeine intake for 24 h. The experi-
mental timeline for day 2 is shown in Fig. 1. All participants remained
sedentary in a semirecumbent position in the Human Performance
Laboratory for minutes 0 –200, with the exception of during the RE
session. Dialysate samples were collected every 20 min for 200 min
(the in-laboratory portion of the experiment) and, thereafter, were
taken every 60 min for the remainder of the day (away from the
laboratory in ambulatory subjects). In addition, a single overnight
dialysate collection was made for each participant for each probe to
determine interstitial glycerol concentration using the extrapolation-
to-zero flow method (described below; Refs. 4, 52). Plasma samples
were taken every 20 min between dialysate collection times for 180
min (10 collections). The prolonged period of study was performed to
monitor the 24-h effects of RE on lipolysis.
Body composition. Participants were weighed on an electronic scale
with weight recorded to the nearest 0.1 kg, and height was measured
with a standard stadiometer. Fat-free mass, fat mass, and percent body
Table 1. Subject characteristics
Variable Lean Obese
n 10 10
Age, yr 23.2⫾1.6 27.3⫾2.4
Weight, kg 69.6⫾2.9 120.5⫾8.4*
Height, cm 182.1⫾2.6 183.0⫾2.4
BMI, kg/m2 20.9⫾0.6 36.2⫾2.7*
Body fat, % 13.6⫾1.5 37.8⫾2.2*
Fat mass, kg 9.7⫾1.2 46.8⫾5.7*
Fat-free mass, kg 59.9⫾1.9 73.7⫾3.3*
Waist/hip ratio 0.81⫾0.01 0.95⫾0.03*
HOMA-IR 0.8⫾0.1 3.1⫾0.6*
Values are means ⫾ SE; n ⫽ no. of subjects. BMI, body mass index;
HOMA-IR, homeostatic model assessment of insulin resistance. *Significantly
different from lean subjects, P ⬍ 0.005.
J Appl Physiol • VOL
fat were determined using dual-energy x-ray absorptiometry (DXA;
GE Lunar Prodigy Advance, Madison, WI).
10RM strength testing and RE protocol. The participant’s 10RM
lifts for the exercises used during the strength testing period were
determined using previously described procedures (28, 56) and were
performed using the resistance training equipment (Cybex, Medway,
MA) that the subjects were familiarized to during preliminary testing.
The RE protocol consisted of the following exercises performed in
the listed order: chest press, lat pull down, leg press, shoulder press,
leg extension, and leg curl. Each exercise was performed for three
total sets: two sets of 10 repetitions and a third set to muscular
exhaustion with a load equaling 85% of the individual’s previously
established 10RM. Rest periods were kept to 90 s between all sets and
exercises, and the RE session lasted for a total of 40 min. The workout
was designed to be similar to those from other studies in which plasma
catecholamines, anabolic hormones, insulin, and lactate concentra-
tions were significantly affected (27, 28, 56). Water intake was
allowed ad libitum during the experimental period. All 10RM testing
and RE sessions were supervised by a certified strength and condi-
tioning specialist.
Experimental protocol. Participants entered the laboratory at 0700
Downloaded from jap.physiology.org on March 2, 2012
after an overnight fast. Each participant had their body weight mea-
sured and then rested in a semirecumbent position for insertion of the
microdialysis probes. Three previously sterilized microdialysis probes
(CMA 20 Elite, 10-mm membrane, 14/10 PAES, 20-kDa cutoff, CMA
Microdialysis; Stockholm, Sweden) were inserted percutaneously into
the participant’s SCAAT, at a distance of 5–10 cm from the umbili-
cus, with techniques previously described (21). Before probe inser-
tion, the skin was cleaned with iodine and the insertion site numbed
with ethyl chloride (a topical cold spray) to reduce discomfort. All
three probes were attached to portable microdialysis pumps (CMA
107, CMA Microdialysis) that were continuously perfused at 2.0
␮l/min with 0.9% sodium chloride (Braun Medical, Irvine, CA)
containing ⬃10 mM ethanol. Ethanol was added to the perfusion
medium to estimate local adipose tissue blood flow as previously
described (21, 23, 24). After a 60-min equilibration period, one
control probe was perfused with 0.9% sodium chloride and etha-
nol. A second probe was perfused with the control solution plus 0.1
mmol/l of the nonspecific ␣-AR inhibitor phentolamine (Sigma-
Aldrich, St. Louis, MO). The third probe was perfused with the
control solution plus 0.1 mmol/l of the nonspecific ␤-AR inhibitor
propranolol (Sigma-Aldrich). The perfusate was collected at the
exit end of the probe (dialysate) and stored at 4°C for analysis of
ethanol within 24 h and subsequently stored at ⫺20°C until
analyzed in batch for dialysate glycerol (an indicator of lipolysis)
according to the manufacturers’ instructions using an automated
microdialysate analyzer (CMA 600 analyzer, CMA Microdialysis,
Solna, Sweden).
An indwelling polyethylene catheter was inserted into the antecubital
vein for blood sampling. Blood samples (10 ml) were collected every 20
min, in the middle of each microdialysis sample collection period for the
duration of the laboratory portion of the experiment (180 min), and
immediately stored at ⫺80°C for later analysis of insulin, epinephrine,
norepinephrine, nonesterified fatty acids (NEFA), glucose, glycerol, and
GH concentrations. Blood from a heated hand was obtained via a
fingerstick and immediately analyzed for blood glucose (Accucheck One
Touch, Lifescan Inc, Milpitas, CA) to verify compliance with our pretest
fasting protocol and to ensure that each participant was not hyperglyce-
mic or diabetic.
EE was determined for 40 min via indirect calorimetry (Parvomed-
ics, True Max 2400, Sandy, UT) before RE. Participants were re-
quired to remain awake, quiet, and as motionless as possible while the
room was darkened and noise kept to a minimum. Substrate oxidation
was calculated using CO2 produced/O2 consumed and calculations
developed by Frayn (14). During the pre-RE EE, two 20-min baseline
dialysate and blood collections were taken.
106 • MAY 2009 • www.jap.org
 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY
Participants then engaged in the high-intensity RE session. Imme-
diately following RE, each participant laid in a semirecumbent posi-
tion while the dialysate collection vial was changed and measurement
of post-RE EE began for 40 min as described above. Dialysate
samples were collected throughout the day. Before going to bed the
perfusion pumps were changed from 2.0 to 0.3 ␮l/min for all three
probes. After a 5-min equilibration period an overnight collection vial
was placed on each probe while each participant slept overnight to
calculate glycerol recovery in each probe.
A perfusion rate of 0.3 ␮l/min in adipose tissue, in the present
study, represents 96% glycerol recovery as assessed by the methods of
Stich et al. (52). The estimated interstitial glycerol concentrations
were calculated by plotting (after log transformation) the concentra-
tion of glycerol in the dialysate measured at 0.3 and 2.0 ␮l/min against
the perfusion rates (52). Linear regression analysis was used to
extrapolate dialysate concentration at zero perfusion flow rate, corre-
sponding to the true interstitial glycerol concentration. The in vivo
recovery rate for each probe was determined as the ratio between the
dialysate glycerol concentration at a perfusion rate 2.0 ␮l/min and
calculated interstitial glycerol concentrations (4, 52). When each
probe was analyzed, there was no difference between individual probe
recovery rates, and therefore the average recovery rate (33%) was
used to calculate interstitial glycerol concentrations.
The following morning, subjects returned to the laboratory for
removal of microdialysis probes and collection of all microdialysis
samples. While away from the laboratory, participants collected and
stored all microdialysis samples in a refrigerator or on ice.
Dietary and activity control. Participants recorded their dietary
intake and physical activity for 3 days before the day of the experi-
ment. Diets were analyzed with Nutritionist Pro software (Axxya
Systems, Stafford, TX). In addition, each participant completed an
activity questionnaire and an in-person interview before participation
in this study to ensure a sedentary lifestyle . All subjects self-reported
as completely sedentary with no structured daily exercise outside
of those activities required for daily living. After leaving the
laboratory, the participants remained sedentary for the rest of the
experimental day.
Eighty minutes after completion of the RE portion of the experi-
ment, participants were fed a liquid meal (Ensure; 14.4% of total kcal
protein, 21.6% of total kcal fat, and 64% of total kcal carbohydrate)
to account for 25% of the individual’s total daily EE. This was
calculated from the before-RE resting energy expenditure (REE)
measurement and included estimated caloric cost of daily activities
(REE ⫻ 0.2), estimated thermic effect of food (REE ⫻ 0.1) (19), and
the estimated cost of our RE protocol [⬃300 kcal; estimated from
previous work by Ormsbee et al. (43)]:
total daily energy expenditure: REE ⫹ 共REE ⫻ 0.2兲 ⫹ 共REE ⫻ 0.1兲
⫹ 300
J Appl Physiol • VOL
1531
Fig. 1. Study timeline. Dialysate and blood
samples were collected before, during, and after
acute resistance exercise in 10 lean and 10 obese
sedentary men. Dialysate samples 1–10 were
collected in the laboratory, and the remaining
samples were collected at home. Subjects re-
turned to the laboratory the following morning
for removal of the microdialysis probes.
For the remainder of the day, the participants were free to leave the
laboratory and go through their normal daily activities; however, each
participant was assigned an isocaloric diet of Ensure to control for
differences in lipolysis due to dietary intake. Compliance was deter-
Downloaded from jap.physiology.org on March 2, 2012
mined by telephone and/or personal contact throughout the day,
written documentation of liquid meal ingestion, collection of empty
drink containers, and specific questioning of the participants by the
research staff.
Blood flow. Ethanol (⬃10 mM) was included with the perfusion
medium to monitor adipose tissue blood flow in the area of the
microdialysis probe (21, 23, 24). During perfusion, ethanol diffuses
over the dialysis membrane and away from the local area by the
microcirculatory blood flow in the immediate vicinity of the probe
membrane because ethanol is not metabolized in adipose tissue to any
significant extent (24). Ethanol concentrations were subsequently
measured in both the dialysate and the perfusion solutions using a
multilabel plate reader (Wallac Victor3, Perkin-Elmer, Waltham, MA)
and previously described enzymatic fluorometric methods (24). Blood
flow is expressed as the ratio of the ethanol concentration in the
dialysate (outflow) to the ethanol concentration in the perfusate
(inflow):
ethanol outflow/inflow ratio ⫽ 关ethanol dialysate兴/关ethanol perfusate兴
Therefore, the ethanol outflow/inflow ratio is related to the blood flow
in the adipose tissue in an inverse manner (24).
Blood analysis. Blood was analyzed for insulin, glucose, glycerol,
NEFA, catecholamines, and growth hormone. All blood was centri-
fuged, and aliquots of serum and plasma were immediately stored at
⫺80°C for later batch analysis to limit day-to-day assay variability.
Plasma glucose and insulin were measured with a YSI model 2300
Stat Plus (Yellow Springs Instrument, Yellow Springs, OH) and
Access Immunoassay System (Beckman Coulter, Fullerton, CA),
respectively, according to the manufacturer’s instructions. Intra-assay
coefficients of variation (CV) for glucose and insulin were 1.9% and
2.1%, respectively. Homeostasis model assessment of insulin resis-
tance (HOMA-IR) was calculated using the following equation (26):
[fasting glucose (mmol/l) ⫻ fasting insulin (␮U/ml)]/22.5.
Plasma glycerol was analyzed with a two-part flourometric assay
adapted from Wieland et al. (59). Briefly, 70 ␮l of heparanized plasma
was added to each well containing 140 ␮l of buffer solution (0.2 M
glycine, 1 M hydrazine, 2 mM magnesium chloride; Sigma-Aldrich),
5 ␮l of 20 mM NAD (Sigma-Aldrich), and 5 ␮l of 50 mM ATP
(Sigma-Aldrich). After measuring the fluorescence using a multilabel
plate reader (Wallac Victor3, Perkin-Elmer), 10 ␮l of an enzyme
solution containing 2 ␮l of glycerophosphate dehydrogenase (GDH;
Sigma-Aldrich), 2 ␮l of glycerokinase (GK; Sigma-Aldrich), and 6 ␮l
of sterile water was added to each well. After shaking the plate and a
40-min incubation period in the dark (at room temperature), fluores-
cence was measured using a 355-nm excitation filter and a 430-nm
106 • MAY 2009 • www.jap.org
1532 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY
emission filter. The concentration of plasma glycerol was determined
following the calculation of a standard curve using a known amount
of glycerol. Intra-assay CV was 4.8%.
Plasma NEFAs were analyzed with a colorimetric reagent kit
[NEFA-HR(2), Wako Chemicals, Richmond, VA] according to the
manufacturer’s instructions. Intra-assay CV was 4.2%.
Plasma catecholamine concentrations were determined by HPLC
utilizing Dionex ICS-3000 instrumentation (Sunnyvale, CA). Intra-
assay and interassay CVs were 5.2% and 5.4%, respectively, for
norepinephrine and 4.1% and 4.5%, respectively, for epinephrine.
Serum GH was analyzed using the Access Immunoassay System
(Beckman Coulter, Fullerton, CA) according to the manufacturer’s
instructions. Intra-assay CV was 7.6%.
Statistical analysis. Two-way (group ⫻ time) repeated-measures
ANOVA tests were used to determine differences in EE, RER, fat
oxidation, plasma measurements (insulin, glucose, glycerol, NEFAs,
catecholamines, and GH), and group-specific SCAAT lipolysis
changes (treatment ⫻ time). Three-way [group (LN or OB) ⫻ time ⫻
treatment (control vs. phentolamine vs. propranolol)] ANOVA tests
were used to determine differences in SCAAT lipolysis. Significance
was located with Newman-Keuls post hoc analysis. SigmaStat statis-
tical software (Systat, San Jose, CA) was used for all analyses. The
level of significance was set at P ⬍ 0.05, and all values are reported
as means ⫾ SE unless otherwise noted.
RESULTS
Subject characteristics. Subject characteristics are presented
in Table 1. With the exception of the leg press exercise (OB
significantly higher than LN, P ⫽ 0.003), there were no
differences in 10RM for any of the exercises performed (chest
press: LN 39.3 ⫾ 2.9 vs. OB 45.7 ⫾ 4.3 kg; lat pull down: LN
35.2 ⫾ 1.7 vs. OB 39.3 ⫾ 1.9 kg; leg press: LN 98.4 ⫾ 4.9 vs.
OB 123.4 ⫾ 6.6 kg; shoulder press: LN 27.5 ⫾ 2.1 vs. OB
29.3 ⫾ 2.2 kg; leg extension: LN 38.6 ⫾ 2.9 vs. OB 45.2 ⫾ 3.7
kg; leg curl: LN 39.5 ⫾ 1.9 vs. OB 44.8 ⫾ 5.5 kg). Total
volume lifted (weight lifted in kg ⫻ no. of repetitions) was
significantly greater in OB compared with LN (P ⫽ 0.01);
however, this result was driven primarily by the leg press
exercise (Table 2). In addition, no differences were reported in
nutritional intake between LN and OB men for total kilocalo-
ries, protein, carbohydrate, fat, or alcohol intake (data not
shown).
Whole body EE and fat oxidation. EE was significantly
greater at baseline in OB patients compared with LN individ-
uals when expressed in kcal/h (LN 76.1 ⫾ 3.6 vs. OB 95.6 ⫾
4.0 kcal/h, P ⬍ 0.05). However, this result was completely
abolished when expressing EE per kilogram of fat-free mass
(Fig. 2). EE was 13 ⫾ 4.4 and 22 ⫾ 4.6% greater after RE in
Table 2. Average total volume of weight lifted (weight lifted
in kg ⫻ number of repetitions) in sedentary lean and obese
men during the resistance exercise lifting protocol
Lean Obese
Chest press 1,115.9⫾98.9 1,374.3⫾148.0
Lat pull down 1,040.8⫾56.4 1,118.2⫾88.1
Leg press 3,124.5⫾179.8 5,011.4⫾545.4*
Shoulder press 681.1⫾64.8 740.2⫾99.3
Leg extension 1,056.5⫾54.9 1,304.3⫾88.7
Leg curl 1,182.5⫾411.8 1,151.6⫾887.9*
Values are means ⫾ SE. *Significantly different from lean subjects,
P ⬍ 0.05.
J Appl Physiol • VOL
Fig. 2. Energy expenditure before and after resistance exercise (RE) in 10 lean
and 10 obese sedentary men. Values are means ⫾ SE. *Significantly different
from before RE, P ⬍ 0.001.
Downloaded from jap.physiology.org on March 2, 2012
LN and OB groups (P ⬍ 0.001), respectively, but there were
no differences between groups.
RER was significantly reduced immediately following RE in
both LN and OB groups (P ⬍ 0.001; Fig. 3A). Fat oxidation
was not different at baseline between LN and OB groups.
There was a significant increase in fat oxidation for LN and OB
groups immediately following RE (LN 49 ⫾ 31.3%; OB 75 ⫾
31.1%; P ⬍ 0.001; Fig. 3B). However, there was no group
difference or group-by-time interaction for fat oxidation.
Interstitial glycerol concentrations in SCAAT. Interstitial
glycerol concentrations in SCAAT were not different between
LN and OB subjects in the control probes at rest (Fig. 4A).
Glycerol concentrations in SCAAT were two- to threefold
higher than glycerol concentrations in plasma at rest in both
groups. Basal interstitial glycerol concentrations were not dif-
ferent between any of the probes or between LN and OB men.
At rest, only the propranolol probe yielded interstitial glycerol
concentrations that were different between groups (LN
104.0 ⫾ 11.9 vs. OB 157.5 ⫾ 19.6 ␮mol/l, P ⫽ 0.03; Fig. 4C).
Throughout the entire study period (minutes 0 – 800; Fig.
4), interstitial glycerol concentrations were significantly
greater in the LN group compared the OB group in the
control and phentolamine probes (P ⬍ 0.05), but not the
propranolol probe (P ⫽ 1.0).
During the RE period (minutes 40 – 80), the peak interstitial
glycerol concentration in the control probe was significantly
greater in the LN group than the OB group (P ⫽ 0.03; Fig. 4A).
Phentolamine induced significantly greater interstitial glyc-
erol concentrations compared with the control probe (P ⬍
0.05) and resulted in significantly higher interstitial glycerol
concentrations over the entire study period in the LN group
only (control 159 ⫾ 10.4 vs. phentolamine 202.4 ⫾ 15.3
␮mol/l, P ⫽ 0.03). In OB subjects, interstitial glycerol con-
centrations were enhanced in response to RE but were not
different from the control probe at any time point over the
study period.
In the propranolol probe, RE resulted in a significant in-
crease in interstitial glycerol concentrations compared with
baseline concentrations in both groups (P ⬍ 0.001). However,
106 • MAY 2009 • www.jap.org
 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY 1533
LN group for the entirety of the study period (LN 13.1 ⫾ 2.8
vs. OB 37.3 ⫾ 6.9 ␮U/ml; P ⬍ 0.002).
Plasma glucose concentrations (Table 3) were significantly
higher in the OB group compared with the LN group over the
experimental time period (LN 94.0 ⫾ 2.6 vs. OB 102.0 ⫾ 2.6
mg/dl; P ⬍ 0.05).

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Fig. 3. A: respiratory exchange ratio before and after RE in 10 lean and 10
obese sedentary men. Values are means ⫾ SE. *Significantly different from
before RE, P ⬍ 0.001. B: fat oxidation before and after RE in 10 lean and 10
obese sedentary men. *Significantly different from before RE, P ⬍ 0.001.

no difference was detected between the propranolol and control

probes in either the LN (P ⫽ 0.5) or OB (P ⫽ 0.2) groups over
time (Fig. 4C).
Plasma catecholamines and serum GH. Plasma norepineph-
rine and epinephrine concentrations were not different at rest
between LN and OB subjects (Fig. 5). RE induced a significant
increase in both plasma catecholamines (P ⬍ 0.001) that lasted
beyond the completion of RE. There was no group difference
or group-by-time interaction for either of the catecholamines.
GH concentrations were not different between LN and OB Fig. 4. Interstitial glycerol concentrations in subcutaneous abdominal adipose
tissue (SCAAT) calculated from dialysate glycerol levels before, during, and
groups at rest. In response to RE, LN subjects had significantly after 40 min of high-intensity RE (horizontal filled bar) in 10 lean and 10 obese
(P ⬍ 0.001) elevated GH concentrations compared with resting sedentary men. Microdialysis probes were placed in SCAAT with one of the
values and compared with the OB group at 70, 90, 110, and following added to the perfusion medium: a control solution (A), 0.1 mmol/l of
130 min. Group differences were measured in the GH response the ␣2-adrenergic receptor (AR) antagonist phentolamine (B), or 0.1 mmol/l of
between LN and OB men (LN 3.3 ⫾ 0.7 vs. OB 1.0 ⫾ 0.9 the ␤-AR antagonist propranolol (C). Values are means ⫾ SE. *Significant
effect of time with glycerol concentrations greater than baseline concentra-
ng/ml, P ⫽ 0.037; Fig. 6). tions, P ⬍ 0.001. †Significantly different glycerol concentrations from obese
Plasma glucose and insulin. Plasma insulin concentrations group, P ⬍ 0.05. Lean response in B is significantly greater than the lean
(Table 3) were significantly higher in the OB group than the response in A.

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1534 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY

probe (control 0.73 ⫾ 0.0005, phentolamine 0.77 ⫾ 0.001,

propranolol: 0.78 ⫾ 0.001; P ⬍ 0.001). There was no main
effect of time or group-by-time or group-by-treatment interac-
tions (data not shown).
DISCUSSION

Microdialysis of SCAAT was used to assess lipolysis in situ

during physiological and pharmacological stimulation of adi-
pose tissue lipolysis in LN and OB men. To our knowledge, no
group has investigated whole body and SCAAT lipolysis in
sedentary LN and OB men in response to an acute bout of RE.
The major findings of the this investigation suggest that despite
a significant increase in whole body energy expenditure and fat
oxidation observed in both sedentary LN and OB men follow-
ing acute RE, SCAAT lipolysis is blunted during RE in OB
men, and this impairment is unrelated to RE-induced catechol-
amine activation of antilipolytic ␣2-ARs (Fig. 4).
In agreement with our previous work and that of others (38,
44, 50), EE (Fig. 2) and whole body fat oxidation (Fig. 3) were
significantly elevated following RE for a period of at least 40

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min; however, there were no differences in these measures
between LN and OB groups. This result confirms recent reports
of a significant increase in EE in LN and OB men with 30 min
of circuit-type RE (8). It is known that EE is elevated for up to
38 h after an acute bout of heavy RE. A reduction in RER has
also been reported by others both immediately (8, 13, 44) as
well as hours (17, 38, 50) after a single bout of RE. The
subjects in the present study may therefore have had elevated
EE and whole body fat oxidation for a period of time greater
than 40 min after RE. A limitation of the present study is not
only the lack of 24-h EE data but also the fact that whole body,
rather than pool-specific, fat oxidation was measured, as it has
been shown that a reduced oxidation of plasma fatty acids is
Fig. 5. Plasma epinephrine (A) and norepinephrine (B) concentrations before, compensated for by an increased oxidation of intramuscular
during, and after RE (horizontal filled bar) in 10 lean and 10 obese sedentary fatty acids in obese individuals during endurance exercise (40).
men. Values are means ⫾ SE. *Significantly greater than baseline value, p ⬍ We observed a significant increase in plasma glycerol during
0.001. There was no difference between groups and there was no group-by- and following RE in both groups but no difference between LN
time interaction.
and OB at any time point (Table 3). Our results verify those of
Borsheim et al. (7) who found an increase in plasma glycerol
with aerobic exercise, but no difference between LN and OB
Plasma glycerol and NEFA levels. Plasma glycerol and
NEFA concentrations are depicted in Table 3. Plasma glycerol
concentrations were significantly elevated (P ⬍ 0.001) at the
onset of RE and remained significantly elevated for the entirety
of the experiment in both groups.
Plasma NEFA concentrations were similar at rest between
LN and OB men and significantly changed over-time (P ⬍
0.001). With the onset of RE, plasma NEFA concentrations
tended to decrease in both groups; however, they only reached
significance (P ⬍ 0.02) in the LN group at minutes 90, 110,
and 130 compared with rest. Plasma NEFAs did not change
from baseline values in the OB group at any time point.
Ethanol outflow-to-inflow ratio in SCAAT. Blood flow was
unchanged in any probe before, during, or after acute RE. The
OB group had a higher (P ⬍ 0.001) ethanol outflow-to-inflow
ratio (LN 0.74 ⫾ 0.0004 vs. OB 0.78 ⫾ 0.0004, P ⬍ 0.001)
than the LN group in all probes, indicating lower SCAAT
blood flow throughout the experimental day. In addition, there
Fig. 6. Serum growth hormone concentrations before, during, and after RE
was a treatment effect in that both the phentolamine and (horizontal filled bar) in 10 lean and 10 obese sedentary men. Values are
propranolol probes significantly elevated the ethanol outflow- means ⫾ SE. * Significantly greater than baseline values, P ⬍ 0.001. †
to-inflow ratio (reduced blood flow) compared with the control Significantly different from lean group, P ⬍ 0.05.

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 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY 1535
Table 3. Plasma concentrations of glucose, insulin, glycerol, and NEFA in sedentary lean and obese men before, during,
and after resistance exercise
Time, minutes

10 30 50 70 90 110 130 150 170 190

Glucose, mg/dl
LN 85.9⫾3.0 87.5⫾2.4 93.2⫾1.5 100.7⫾3.9 94.9⫾4.5* 93.3⫾3.7 84.8⫾3.1 82.0⫾2.5 94.8⫾3.0 122.6⫾7.0*
OB 94.9⫾2.0 95.4⫾1.8 96.1⫾1.9 94.6⫾6.6 106.4⫾4.8* 105.5⫾4.7 98.1⫾4.3† 94.8⫾3.5 104.4⫾3.6 130.1⫾5.7*
Insulin, ␮U/ml
LN 3.9⫾0.6 3.5⫾0.5 6.6⫾0.8 7.1⫾1.4 10.9⫾2.3* 8.4⫾2.1 4.8⫾1.7 3.8⫾0.4 17.8⫾6.1 64.2⫾12.7*
OB 13.1⫾2.1† 13.1⫾2.2† 14.5⫾2.2† 18.1⫾3.8† 25.5⫾4.5*† 26.1⫾4.6*† 20.1⫾5.0*† 17.4⫾3.9† 46.3⫾12.8*† 178.4⫾28.0*†
Glycerol, ␮mol/l
LN 45.8⫾2.5 44.9⫾3.1 114.2⫾7.6* 137.8⫾9.3* 140.2⫾11.6* 107.7⫾9.3* 88.4⫾9.2* 81.5⫾13.2* 87.6⫾12.0* 70.6⫾6.5*
OB 57.4⫾5.1 63.9⫾8.7 114.4⫾10.4* 133.6⫾8.0* 142.2⫾9.7* 125.6⫾9.7* 95.4⫾5.7* 98.3⫾8.1* 87.9⫾6.1* 77.5⫾4.7*
NEFA, mmol/l
LN 0.5⫾0.04 0.5⫾0.04 0.4⫾0.03 0.4⫾0.03 0.4⫾0.03* 0.3⫾0.02* 0.3⫾0.03* 0.5⫾0.1 0.6⫾0.1 0.4⫾0.04
OB 0.5⫾0.1 0.5⫾0.1 0.4⫾0.02 0.4⫾0.02 0.5⫾0.1 0.4⫾0.1† 0.4⫾0.1 0.5⫾0.03 0.4⫾0.03† 0.4⫾0.02
Values are means ⫾ SE. LN, lean; OB, obese; NEFA, nonesterified fatty acids. Resistance exercise took place during minutes 40 – 80 of the experiment.
*Significantly greater than baseline values, P ⬍ 0.001. †Significantly different from LN group, P ⬍ 0.05.

groups. We found no difference in the epinephrine or norepi- elevated dialysate glycerol concentrations. Therefore, OB sub-

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nephrine response between groups at rest or with RE (Fig. 5),
which corroborates the findings of Stich et al. (52). The mean
plasma insulin level (Table 3) and HOMA-IR values were
significantly higher in OB than LN subjects, which may indi-
cate different insulin sensitivities between groups, confirming
earlier findings (25, 52). It is well known that insulin sup-
presses lipolysis (1, 22, 45), and it has been reported that the
lipolytic effect of epinephrine is blunted during a hyperinsu-
linemic-euglycemic clamp in men (45, 54). It is possible that
the discrepancy in plasma glycerol response between this study
and that of others is due to the fact that our study utilized very
obese individuals (BMI ⫽ 36.2 kg/m2) and a high-intensity RE
protocol rather than a moderate-intensity aerobic exercise
study design. The similar plasma glycerol response between
groups in the present study, despite a lower dialysate glycerol
response in OB than LN men, could be due to greater adipocyte
mass or hydrolysis of triglycerides in areas other than SCAAT
in OB subjects. In addition, the contribution of lipolysis from
visceral adipocytes was not investigated in this study and may
have influenced the systemic glycerol response (35, 36).
Plasma NEFA concentrations were not different at rest
between groups and were significantly lower than baseline
concentrations for 50 min following RE in the LN group only
(Table 3). The latter is most likely due to a combination of
upregulated fat oxidation and reesterification of fatty acids,
although without the use of stable isotope tracers we cannot be
certain of the fatty acid partitioning. Other groups confirm our
findings of no difference in fasting plasma NEFA concentra-
tions at rest between LN and OB subjects (7, 25). There were
significant differences between groups 30 min and 90 min after
RE, which may be due to some metabolic inflexibility in the
OB group. Our results conflict with Stich et al. (52), who
reported significantly greater NEFA concentrations at rest and
in response to 60 min of cycling at 50% of heart rate reserve in
OB compared with LN men. This result is likely due to the
mode and intensity of exercise used in previous studies com-
pared with our high-intensity RE protocol.
Basal interstitial glycerol concentrations were not different
between LN and OB subjects in the control probe (Fig. 4A).
However, OB individuals had significantly lower blood flow
throughout the study period, which should have resulted in
J Appl Physiol • VOL
jects most likely had lower SCAAT glycerol release at rest than
LN subjects.
RE-induced increases in interstitial glycerol were signifi-
cantly greater in LN (⫹142.1 ⫾ 30.8%) than in OB (⫹65.4 ⫾
14.2%) men in the control probe, a result that was not rectified
by the ␣-AR antagonist phentolamine in OB men (Fig. 4B). In
fact, phentolamine perfusion enhanced the RE-induced in-
crease in interstitial glycerol in LN rather than OB men,
demonstrating that catecholamine stimulation of the antilipo-
lytic ␣-ARs is not responsible for decreased lipid mobilization
in OB men during RE. This is in contrast to the ␣-AR-mediated
suppression of lipolysis in obese men during endurance exer-
cise (52).
An increase in SCAAT glycerol could be achieved by either
elevated local lipolysis or a reduced blood flow in SCAAT
(39). Indeed, OB subjects in the present study had a signifi-
cantly greater ethanol outflow-to-inflow ratio (reduced blood
flow) over the course of the experiment compared with LN
subjects, which agrees with others (7, 52). One would expect
interstitial glycerol concentrations to be artificially elevated in
OB men due to reduced blood flow. Despite this lower blood
flow in OB men, interstitial glycerol concentrations were
markedly lower in OB men compared with LN men. Therefore,
the lipolytic response to RE was lower in OB than LN men,
and it is possible that the magnitude of this difference was
underestimated.
The mechanisms of local SCAAT lipolytic regulation were
further elucidated by adding the nonselective ␤-AR blocker
propranolol to the microdialysis perfusate. In the present study,
neither LN nor OB subjects had any lipolytic alterations in
response to propranolol compared with the control probe at rest
or during RE (Fig. 4C). Our results substantiate those of
Hellstrom et al. (20) that resting dialysate glycerol levels were
not influenced by ␣- or ␤-adrenergic blockers. However, dur-
ing endurance exercise, perfusion of propranolol has previ-
ously been shown to reduce endurance exercise-induced in-
creases in interstitial glycerol concentrations (3, 20, 41). The
discrepancy between these results and our data could be the use
of different types of exercise modalities. The paradoxical
increase in SCAAT interstitial glycerol concentrations during
RE despite inhibition of adipocyte ␤-ARs is likely associated
106 • MAY 2009 • www.jap.org
1536 RESISTANCE EXERCISE, FAT METABOLISM, AND OBESITY
with alternative lipolytic regulators. One such regulator is atrial
natriuretic peptides (ANP), which has been shown to regulate
endurance exercise-induced lipolysis and works via an insulin-
independent mechanism (30, 31, 41, 42, 51). Our results
suggest that at rest and during RE, the ␤-AR stimulatory
effects on lipolysis may not be as involved in modulating
lipolysis as previously shown during aerobic exercise in LN
and OB men.
GH, another important regulator of lipolysis, is widely
recognized as a stimulator of regional and systemic lipolysis
(11, 32, 49). In the present study, GH increased markedly from
resting values in response to RE in LN but not OB subjects and
remained elevated for 50 min after the RE session (Fig. 6). An
increase in GH during and following an acute bout of high-
intensity RE, similar to that used in this study has been shown
elsewhere in healthy subjects (18, 28). Interestingly, Enevold-
sen et al. (12) reported that GH induced lipolysis following, but
not during, exercise at 50% maximal oxygen uptake (V̇O2max)
on a semirecumbent cycle ergometer in healthy, normal-weight
young men. In light of this finding, it is apparent that further
research regarding the role of GH during RE is warranted.
Moreover, it is well established that spontaneous and stimu-
lated serum GH concentrations are suppressed in human obe-
sity (15, 48) and return to normal with normalization of body
weight (48). It is possible that the blunted lipolytic response
observed in OB compared with LN subjects in SCAAT may, in
part, be related to the significantly diminished GH response to
RE in the OB.
Several methodological limitations exist in the present
study. Participants were ambulatory, and therefore without
supervision, following the in-laboratory portion of the experi-
ment. However, the participants were provided all of their daily
calories in liquid form to control for any influence of dietary
intake on the lipolytic response to RE. In addition, using stable
isotopic tracers to monitor lipolysis and fat oxidation would
add valuable information and should be included in future
investigations.
In conclusion, high-intensity RE improves whole body EE
and fat oxidation in LN and OB subjects. However, we dem-
onstrate for the first time that, in SCAAT, OB men have
suppressed RE-induced lipolysis compared with LN men and
the mechanism for this is unrelated to overstimulation of the
antilipolytic ␣2-AR system by catecholamines. In addition, we
determined that ␤-ARs are not involved in the regulation of
lipolysis during RE in sedentary LN or OB men. The blunted
lipolytic response to RE in OB subjects could be due to the
significantly lower serum GH response to RE compared with
LN subjects. It is also a possibility that an alternative lipolytic
pathway, such as that induced by ANP, is playing an important
role in lipolysis; however, ANP was not measured in the
present study. It is important to recognize that the results of the
present study are specific to the SCAAT of sedentary LN and
OB men during RE.
Nevertheless, the favorable metabolic effects resulting from
acute RE are relevant, as alternative or multiple modes of
exercise may be necessary to strategically combat the delete-
rious effects of obesity. Indeed, RE training has been shown to
improve catecholamine-mediated lipolysis in OB subjects (45).
Furthermore, because RE may be a mode of exercise more
suitable for OB individuals than cardiovascular endurance
exercise, and RE is gaining popularity as a primary mode of
J Appl Physiol • VOL
exercise, future studies are warranted to better understand the
regulation of lipolysis with acute and chronic RE training.
ACKNOWLEDGMENTS
We acknowledge the technical assistance of Pamila Allen and the Univer-
sity of Colorado Health Sciences Center for the analysis of plasma catechol-
amine concentrations, and Raymond Kraus, PhD, for insightful assessment of
this manuscript.
GRANTS
Grant support for M. D. Choi was provided by Experimental and Applied
Sciences, and M. J. Ormsbee was provided grant support by the Gatorade
Sports Science Institute and Phi Kappa Phi National Honors Society.
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