Fuel oxidation at the walk-to-run-transition in humans
Kathleen J. Ganley
a
, Anthony Stock
b
, Richard M. Herman
c
, Marco Santello
b
, Wayne T. Willis
b,
⁎
a
Department of Physical Therapy and Athletic Training, Northern Arizona University, Flagstaff, AZ, USA
b
Department of Kinesiology, Arizona State University, Tempe, AZ, USA
c
Harrington Department of Bioengineering, Arizona State University, Tempe, AZ, USA
Received 18 September 2009; accepted 7 June 2010
Abstract
Multiple factors (including anthropometric, kinetic, mechanical, kinematic, perceptual, and energetic factors) are likely to play a role in the
walk-to-run transition in humans. The primary purpose of the present study was to consider an additional factor, the metabolic fuel source.
Indirect calorimetry was used to measure fuel oxidation, and perception of effort was recorded as 10 overnight-fasted adults locomoted on a level
treadmill at speeds progressing from 1.56 to 2.46 m s
-1
in increments of 0.11 m s
-1
and 10.0 minutes under 3 conditions: (1) unconstrained
choice of gait, (2) walking at all speeds, and (3) running at all speeds. The preferred transition speed was 2.08 ± 0.03 m s
-1
. Gait transition from
walking to running increased oxygen consumption rate, decreased the perception of effort, and decreased the rate of carbohydrate oxidation. We
propose that, in an evolutionary context, gait transition, guided by the perception of effort, can be viewed as a carbohydrate-sparing strategy.
© 2011 Elsevier Inc. All rights reserved.
1. Introduction
Quadrupeds can adopt multiple terrestrial gait patterns,
whereas natural bipedal gait patterns in humans are limited to
walking and running. A preferred speed to transition from
one gait to another is commonly observed in most
mammalian species, including humans. In humans, preferred
transition speed (PTS) from walking to running occurs
within the delimited range of 1.8 to 2.2 m s
-1
[1,2].
Factors other than physical limitations trigger human gait
transition because walking at higher speeds and running at
lower speeds than the PTS are well tolerated. Previous
investigations of gait transition have explored anthropomet-
ric [3,4], kinetic [1,5], mechanical [6], kinematic [4,7],
perceptual [2,8], and energetic [1,7,9-11] factors.
Locomotion across the speed range relevant to the PTS is
dependent upon energy derived from the combustion of fat
and carbohydrate, 2 fuels stored at vastly different levels.
Because of the profound evolutionary implications regarding
the need to defend the extremely limited carbohydrate stores
of the human body [12-15], the primary purpose of the
present study was to consider the potential role of metabolic
fuel selection in the walk-to-run transition in humans.
From preferred walking speed up to PTS and beyond,
the energy cost of walking (in kilocalories per kilometer per
kilogram) steadily rises, revealing the right upward leg of
this familiar U-shaped curve. In contrast, the energy cost of
running remains relatively constant across this range of
speed [1]. Prilutsky and Gregor [2] showed that summed
lower extremity electromyography (EMG) conformed to
this same pattern. Thus, they proposed that the steadily
rising neural input to muscle and energy consumption
required to increase walking speed toward PTS might elicit
the transition to running, where the slope of these changes
would flatten.
At the PTS, gait transition from walking to running
increases the rate of energy expenditure [1,7,10,16], despite
the reasonable, and perhaps more intuitive, prediction that it
should have the opposite effect [9,11,17,18]. Moreover, this
elevated energy cost is curiously attended by decreased
perception of effort [2,8,10] and greater neural input into
muscle as assessed from the summed EMG of active muscle
[2]. These published observations present an interesting
paradox: the walk-to-run transition increases energy expen-
diture and the apparent neural input into muscle, whereas it
decreases the perception of effort.
Available online at www.sciencedirect.com
Metabolism Clinical and Experimental 60 (2011) 609 – 616
www.metabolismjournal.com
This study was approved by the Institutional Review Board at Arizona
State University, where the work was performed.
⁎
Corresponding author.
E-mail address: waynewillis@asu.edu (W.T. Willis).
0026-0495/$ – see front matter © 2011 Elsevier Inc. All rights reserved.
doi:10.1016/j.metabol.2010.06.007