RELATIONSHIP BETWEEN FUNCTIONAL HAMSTRING:
QUADRICEPS RATIOS AND RUNNING ECONOMY
IN HIGHLY TRAINED AND RECREATIONAL
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FEMALE RUNNERS
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ØYVIND H. SUNDBY AND MARK L.S. GORELICK
Department of Kinesiology, San Francisco State University, San Francisco, California
ABSTRACT
Sundby, ØH and Gorelick, MLS. Relationship between func-
tional hamstring:quadriceps ratios and running economy in
highly trained and recreational female runners. J Strength
Cond Res 28(8): 2214–2227, 2014—The purpose of this
study was to investigate the relationship between running
economy (RE), functional hamstring:quadriceps peak torque
ratios (f-H:Q), and flexibility among female runners. Seven
highly trained (HT) female runners (age: 25.7 6 4.7 years,
V_ O2peak of 62.0 6 4.8 ml$kg21$min21) and 11 recreational
female runners (age of 28.8 6 5.6 years, V_ O2peak of 49.2 6
4.6 ml$kg21$min21) were measured for maximal aerobic
power (V_ O2peak), RE, heart rate, respiratory exchange ratio,
f-H:Q (Hecc:Qcon and Hcon:Qecc), and sit-and-reach ham-
string/trunk flexibility. On 2 separate days, RE was measured
on a treadmill at 1% grade at 2 velocities (160.9 and
201.2 m$min21) for 6 minutes each, and isokinetic knee
strength was measured at 3 angular velocities (60, 120,
and 1808$s21) for both concentric and eccentric muscle
actions. The unpaired t-tests showed a consistent trend
toward higher f-H:Q ratios at all angular velocities among
the HT runners. Highly trained runners had significantly high-
er Hecc:Qcon at 1208$s21 (p # 0.05) and 1808$s21 (p #
0.05). Whole group correlations demonstrated a significant
correlation between Hcon:Qecc at 1808$s21 and RE
(ml$kg20.75$km21) at 201.2 m$min21 (R = 20.48, p # 0.05).
No significant relationships were found between flexibility, or
hamstring and quadriceps peak torque (N$m) and RE (p .
0.05). This cross-sectional analysis suggests that higher f-H:
Q torque ratios, and not muscle strength per se, are associated
with a lower metabolic cost of running. Therefore, runners
Address correspondence to Øyvind H. Sundby, oyvindheibergsundby@
gmail.com.
28(8)/2214–2227
Journal of Strength and Conditioning Research
Ó 2014 National Strength and Conditioning Association
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2214 Journal of Strength and Conditioning Research
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should consider implementing hamstring exercises to improve
their f-H:Q ratios.
KEY WORDS strength, long-distance running, females, oxygen
cost of running, flexibility
INTRODUCTION
O
xygen consumption is a key to sustain the
repeated muscle contractions required for endur-
ance exercise. There are 3 major physiological
factors that determine power output that can be
transformed into performance pace in distance running: (a)
gross energy cost of running, commonly referred to as work
economy or running economy (RE), (b) maximal aerobic
power (V_ O2max), and (c) the ability sustain a high percent
of V_ O2max (fractional utilization of V_ O2max, % V_ O2max) for
an extended period of time (14,20,23,46,54). These 3 factors
explain more than 70% of interindividual variance in long-
distance running performance (20). Among a homogenous
group of endurance athletes, RE may be a better predictor of
performance than V_ O2max (14,45,50). Indeed, variation in
RE has been found to explain 65.4% of the variation
observed in race performance in a 10-km race (14).
Research to elucidate the variables causing differences in
RE is substantial (4,60,71). Nevertheless, it is not yet clear
precisely which factors contribute to better RE. Running
economy is believed to be influenced by gender (19), type
of training (52,61), fitness level (51), peak stride forces (62),
muscle stiffness (22), nitrate supplementation (44), and
a number of other biomechanical and physiological factors
(4,60,71). It seems that concurrent strength or (36,73) plyo-
metrics training (61,67) and a distance running program may
be common elements that improve RE. Researchers specu-
late that strength training increases RE in part by altering
neuromuscular (52) and elastic components in the musculo-
skeletal system (8,61).
Turning to the relationship between RE and flexibility,
stiff tendon structures have a favorable effect on stretch-
shortening cycle exercise, such as running (3). Tendon stiff-
ness may also enhance force and power production during

dynamic muscle action (8). This may potentially increase
muscle output and propulsion force for the same or less met-
abolic cost by augmenting effective force transmission from
the contractile elements to the bone (8), thereby improving
RE and consequently, performance (18,61). Studies of sit-and-
reach flexibility among male (17,37) and female runners (66),
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as well as hamstring flexibility in untrained subjects (27),
provide evidence of a negative relationship between muscu-
loskeletal system flexibility and oxygen cost of locomotion.
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Moreover, knee extensor endurance (in the quadriceps
muscles) has been shown to significantly correlate with RE
in elite male middle-distance runners (70). Recent studies sug-
gest that local eccentric and concentric muscular endurance in
the knee flexors and hip flexors helps maintain stride mechan-
ics by delaying kinematic changes correlated with fatigue dur-
ing endurance running (29,30). Additionally, a study has
found that hamstring strength is a singular correlate to per-
formance in long-distance running (65).
Although hamstring and quadriceps strength seem to be
critical components of RE, their relative importance is
unknown. Currently, there is little research relating muscle
ratios to functional tasks that are specific to a sport.
Although research has shown that interventions of either
strength exercises or plyometric exercises increase RE, it is
still unclear by what mechanisms this occurs. Understanding
the relationship between muscular strength distribution,
flexibility, and aerobic capacity is important because such
knowledge may advance training methods and thereby
improve distance running performance.
The above findings showing a relationship between both
flexibility and muscular strength with endurance perfor-
mance may be due to both neuromuscular and elastic
components in the muscles, or to how the muscles work
together, that is, their agonist-antagonist relationship. To
date, there has been little research examining the relative
muscle strength ratios required to succeed in sports. To the
best of our knowledge, this is the first report on f-H:Q ratios
applied to functional tasks (RE and flexibility) that are
specific to sport (distance running).
Therefore, the purpose of this study was twofold: First, to
investigate the differences and the relationship between RE
(gross oxygen cost of running) and functional hamstring to
quadriceps torque ratios (f-H:Q) among highly trained (HT)
and recreational (REC) female runners. Second, to examine
RE and its relationship to differences in hamstring/trunk
flexibility, and anthropometric variables commonly known to
be associated with RE among the 2 trained states. In addition
to these 2 main purposes, a secondary purpose of the study
was to establish norms and provide recommendations for f-H:
Q ratio values for healthy female runners. We hypothesized
that (a) compared with REC runners, HT runners would
possess higher f-H:Q ratios at low, medium, and high angular
velocities; (b) that f-H:Q would correlate with better RE at
submaximal velocities; and (c) that RE would be inversely
correlated with flexibility scores on the sit-and-reach test.
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METHODS
Experimental Approach to the Problem
For testing our hypotheses, we used a cross-sectional design
to investigate the impact of relative muscle strength,
flexibility, and anthropometrics on gross oxygen cost of
running (RE). To integrate the concept of studying the
hamstring and quadriceps muscles in motion, we used
a functional method of describing the hamstring to quadri-
ceps relationship, referred to as functional hamstring to
quadriceps ratios (f-H:Q) (2). The HT and REC female run-
ners were divided according to their training status and race
experience. HT runners in this study were defined as those
who were (a) members of a running club or a university track
and field running team at the time of the study, and who had,
over the past 3 years, (b) run $40 miles per week, (c) run $4
times per week, and (d) competed in at least 3 races per year.
The REC runners were recruited based on the following
inclusion criteria: having run (a) 5–30 miles per week and
(b) 2–4 times per week over the last 3 years.
Subjects reported to the laboratory on 2 separate occa-
sions, separated by at least 48 hours. All test trials were
completed within 2 weeks for each subject. The dependent
variables in visit 1 included subjects’ anthropometrics (stat-
ure, body mass, body composition, girth measures, and leg
length), gross oxygen cost of running (RE), % of heart rate
max (% HRmax), respiratory exchange ratio (RER), % of
V_ O2peak (% V_ O2peak), and a maximal aerobic power test.
On visit 2, sit-and-reach flexibility and peak muscular ham-
string to quadriceps (H:Q) torque ratios at slow, medium,
and fast velocities were measured. Slow, medium, and fast
isokinetic velocities were defined as 60, 120, and 1808$s21,
respectively. Similar velocities have been used in previous
investigations of f-H:Q ratios in healthy male Kenyan run-
ners (40), and when examining c-H:Q ratios in healthy male
and female intercollegiate varsity athletes (57). Girth and leg
length variables were selected because of both findings from
previous studies that examined factors influencing RE and
because of their potential mechanical advantage.
The hypotheses were tested by determining the differ-
ences and correlations among the variables. The Student’s
t-test was the primary statistical tool used to determine dif-
ferences in the dependent variables across the 2 trained
groups. Pearson’s correlation coefficients were used to iden-
tify possible relevant relationships. The independent varia-
bles were the velocity of the isokinetic dynamometer with
3 factors: 60, 120, and 1808$s21, and the velocity of the
treadmill with 2 factors: 160.9 and 201.2 m$min21.
Subjects
Twenty-one female runners, 10 highly trained, and 11 REC
runners between the ages of 21–39 years (mean 6 SD: 27.4 6
5.0 years) from local running clubs and the local community
were recruited on a voluntary basis to participate in the study.
Exclusion criteria were any overt metabolic, endocrine, cardio-
vascular, neurological, or metabolic diseases; musculoskeletal
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 Functional Hamstring:Quadriceps Ratios and Running Economy

1. The HT and REC runners

reported their total training fre-
TABLE 1. Physical characteristics of the subjects in mean 6 SD (n = 18).*
quency the last year to be 9.1
Variables Highly trained (n = 7) Recreational (n = 11) (2.8) and 4.1 (1.1) times per
week, respectively. Five and 6
Age (yr) 25.7 6 4.7 28.8 6 5.6
subjects performed supplemen-
58.4 6 5.7 63.3 6 8.8
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Body mass (kg)†

Body fat (%)† 19.7 6 3.0 26.4 6 6.2z tary resistance training among
Fat-free mass (kg)† 47.3 6 4.3 46.3 6 5.2 the HT and REC runners,
Stature (cm) 169.6 6 8.1 167.4 6 7.4 respectively. The REC runners
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BMI (kg$m22) 19.6 6 0.7 22.6 6 2.8z reported that they performed
Ankle girth§ (cm) 20.9 6 1.2 21.0 6 1.3
most of their weekly running
Calf girth§ (cm) 35.0 6 1.1 36.3 6 1.7
Midthigh§ (cm) 46.0 6 4.3 46.7 6 3.5 at low intensities.
Leg lengthk (cm) 93.1 6 4.7 87.5 6 5.7z Before testing on visit 1, the
Leg lengthk/stature (%) 54.1 6 1.5 52.3 6 1.5z subjects completed a question-
Mileage (km$wk21) 104.6 6 18.5 32.0 6 15z naire on physical exercise
Total training (h$wk21) 12.1 6 5.1 6.1 6 2.3z
background and present activ-
*BMI = body mass index. ity level, demographics, medi-
†Based on BOD POD measures. cal history, and previous injury
zSignificant differences between groups at p # 0.05.
§Measured on right leg only. history. Conventional H:Q
kMeasured from anterior superior iliac spine to lateral malleole on right leg only. ratios (Hcon:Qcon) have been
reported to be much lower
among injured individuals
(57). Thus, the authors in the
injury related or unrelated to overtraining; consumption of present study screened for recent injuries in the hamstring
medications or using any drugs that may influence the energy and quadriceps muscles. Past injuries could mask “true” val-
metabolism system; and pregnancy. In addition, exclusion cri- ues because the injury might have affected the athletes’ abil-
teria included any hamstring or quadriceps injury in the past ity to strength train or run hard workouts. Consequently, the
6 months. Before participation, all the subjects were informed results in the present study are deemed to be representative
about the experimental procedures, and the possible risks and for healthy athletes.
discomforts associated with the study. The subjects then Three of the HT runners reported hamstring injuries
signed a written informed consent form before participating. during the past 6 months and were excluded from the data.
This study was conducted according to the Declaration of Table 1 presents the 18 remaining subjects’ demographic
Helsinki, and the protocol was fully approved by the San variables. The HT and REC subjects (n = 18) had been
Francisco State University Institutional Review Board for running for 11.7 6 4.3 years and 4.7 6 2.3 years, respectively,
Human Subjects (protocol number: H11-59). before the study. All of the HT and all but one of the REC
The HT subjects consisted of competitive distance runners runners had run at least 3 or more times on a treadmill
(n = 7) competing across a range of distances throughout the before participating in the study and were thus familiar with
year (5k-marathon distances). Each subject in the HT group treadmill running already. None of the participants were
regularly undertook 1–2 interval and 1 continuous running familiar with the isokinetic dynamometer before participat-
sessions per week. Subject characteristics are listed in Table ing in the study.
The study took place in the
basic training phase for the HT
runners, and all subjects were
TABLE 2. Maximal cardiovascular treadmill results (n = 18).* tested in the period between
Variables Highly trained (n = 7) Recreational (n = 11) January 2012 and March 2012.

V_ O2peak (L$min21) 3.6 6 0.4 3.1 6 0.3† Procedures

V_ O2peak (mL$kg21$min21) 62.0 6 4.8 49.2 6 4.6† Before the tests, the subjects
V_ O2peak (mL$kg20.75$min21) 171.4 6 13.3 138.5 6 11.0†
HRmax (b$min21) 181.7 6 9.6 195.1 6 8.2†
were given instructions on stan-
RERmax 1.1 6 0.0 1.1 6 0.1 dardized behavior to follow.
Subjects were instructed to
*Values are in mean 6 SD. HRmax = heart rate max; RERmax = respiratory exchange ratio arrive at the laboratory in a fully
max.
†Significant differences between groups at p # 0.05. rested and hydrated state, at
least 2 hours postprandial, and
to avoid strenuous exercise for
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as follows: The subjects were

fitted with a mouthpiece for
TABLE 3. Steady-state cardiovascular treadmill exercise results (treadmill at 18 grade).*
expired gas analysis. The
Variables Highly trained (n = 7) Recreational (n = 11) researcher described the
potential for discomfort from
160.9 m$min21
exercising with a mouthpiece
V_ O2 ml$kg21$min21 31.1 6 2.8 32.6 6 2.2
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V_ O2 ml$kg20.75$min21 86.0 6 7.5 91.8 6 6.2† and nose clip and explained
V_ O2 ml$kg21$km21 193.5 6 17.5 202.7 6 13.6 the 15-grade scale for rating
V_ O2 ml$kg20.75$km21 534.2 6 46.3 570.3 6 38.5 of perceived exertion (RPE)
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% V_ O2peak 50.2 6 3.4 66.7 6 6.9† (9). Open circuit spirometry

HR (b$min21) 128.2 6 11.0 159.1 6 18.3†
was used to calculate oxygen
% HRmax 70.6 6 6.2 81.7 6 9.4†
RPE 10.5 6 0.7 12.2 6 1.6† uptake (V_ O2) for steady state
RER 0.8 6 0.1 0.9 6 0.1 and maximal aerobic power
(V_ O2peak) measures with
Variables Highly trained (n = 7) Recreational (n = 10)z a metabolic cart (Parvo Med-
201.2 m$minute21 ics’ True One 2400 [Parvo-
_VO2 ml$kg21$min21 38.3 6 3.0 41.2 6 2.2† Medics Inc., Yorba Sandy,
V_ O2 ml$kg20.75$min21 105.8 6 9.0 115.5 6 5.7† UT, USA]) by sampling breath-
V_ O2 ml$kg21$km21 190.4 6 15.1 204.9 6 11† by-breath expired gases. Each
V_ O2 ml$kg20.75$km21 526.1 6 44.6 574.3 6 28.4†
% V_ O2peak 61.9 6 4.4 83.1 6 6.3†
subject had her nose clamped
HR (b$min ) 21 148.8 6 9.2 174.6 6 14.3† while breathing through a mouth-
% HRmax 82.0 6 5.6 89.6 6 6.7† piece connected to a 2-way non-
RPE 12.0 6 1.4 13.9 6 1.1† rebreathing valve. Ambient air
RER 0.8 6 0.0 0.9 6 0.1† was inspired through a dry hose,
*Values are in mean 6 SD. HR = heart rate; HRmax = heart rate max; RERmax = respi- and expired air was directed to
ratory exchange ratio max. a mixing chamber.
†Significant differences between groups at p # 0.05.
zOnly 10 recreational runners ran steady state at the second stage.
To minimize diurnal biolog-
ical deviations, the researchers
planned to test each subject at
the same time of day for both
visits, however, the researchers
24 hours before the test. Before visit 1, the subjects were were not able to do this for all subjects because of some of
instructed to bring their lightest shoes (racing shoes/light- the participants’ scheduling constraints. Each subject was
weight trainers) and to refrain from any eating and drinking 2 advised to maintain a regular diet the day before testing.
hours before arrival. In addition, to avoid possible confounders The procedures were administered to all subjects by the
related to ergogenic effects, the subjects were asked to refrain same investigator.
from consuming caffeine and alcohol 24 hours before testing The laboratory conditions were relatively stable (temper-
on both visits. To participate in the study, each subject had to ature: 20.1 6 0.58 C, humidity: 45.4 6 7.9%) for the duration
sign statements that they had followed these preparticipation of the study.
procedures. All subjects were instructed to maintain and
record their current training habits using a self-reported Anthropometric Measurements. Standing height was deter-
activity diary. mined without shoes using a wall-mounted stadiometer
Before each test trial, the analyzers and flow meters were (Doran Scales Inc., Batavia, IL, USA) to the nearest 0.1
calibrated strictly according to the manufacturer’s recommen- cm, and measures were taken after a voluntary deep
dations using certified commercial gas preparations of known inspiration. Girth (circumference) measurements were
oxygen and carbon dioxide concentration and a manual 3-L taken to the nearest 0.1 cm with a measuring tape using
syringe (Hans-Rudolph, Kansas City, MO, USA). Tempera- a calibrated tension device (Creative Health Products Inc.,
ture, barometric pressure, and humidity were calibrated and Ann Arbor, MI, USA) as follows: leg length (anterior
updated from an electronic device (Perception II; Davis superior iliac spine to lateral malleolus), hip girth (max-
Instruments, Hayward, CA, USA). During all V_ O2 testing, imum circumferences over the buttocks), thigh girth
heart rate data were monitored continuously using a Polar (midway between inguinal crease and the proximal border
FT1 heart rate monitor (Polar Electro OY, Kempele, Finland). of patella), leg girth (maximal circumferences between
All V_ O2 determinations were performed on a motorized knee and ankle while the leg measured was relaxed), and
Quinton Q65 Series 90 treadmill (Quinton Instruments, Q65, ankle girth (minimum circumferences just above lateral
Series 90; Seattle, WA, USA) following identical procedures malleolus).

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 Functional Hamstring:Quadriceps Ratios and Running Economy

type swimsuit or single-layer

compression shorts and jog
TABLE 4. H:Q ratios for highly trained (n = 7) and recreational female runners (n = 11)
(total: N = 18).* bra without padding or wires.
Each subject’s body mass (kg)
Velocity (8$s21) Highly trained (n = 7) Recreational (n = 11) % Differences was measured on a calibrated
electronic scale connected to
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Hecc:Qcon
60 1.10 6 0.15 0.98 6 0.20 11.5 the BOD POD. Computer
120 1.41 6 0.30 1.15 6 0.12† 20.3 software calculated FFM and
180 1.61 6 0.25 1.39 6 0.19† 14.7 percent body fat using the Siri
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Hcon:Qecc equation (60). After the BOD

60 0.37 6 0.06 0.34 6 0.07 7.1
POD test, subjects were al-
120 0.36 6 0.05 0.31 6 0.06 13.9
180 0.34 6 0.05 0.29 6 0.06 14.3 lowed to consume water ad li-
Hcon:Qcon bitum before and between the
60 0.56 6 0.08 0.51 6 0.09 9.3 treadmill tests. Body mass
120 0.68 6 0.11 0.54 6 0.07† 23.0 index (BMI) was calculated as
180 0.72 6 0.06 0.62 6 0.11† 14.9
body mass in kilograms
*Values are in mean 6 SD. divided by height in meters
†Significant differences between groups at p # 0.05. squared (kg$m22).

Running Economy Test. The

Body Composition Testing. Estimation of body composition
(fat mass, fat-free mass [FFM], and percent body fat) was
assessed through air displacement plethysmography using
the BOD POD (BOD POD; Life Measurement Instruments,
Concord, CA, USA). The BOD POD was calibrated before
each test according to the manufacturer’s instructions using
a cylinder of known volume (50.094 L) and for body mass
using two 10-kg weights. All examinations were performed
by the same researcher. Body composition was measured
using measured thoracic gas volumes (Vtgmeas) (48). Each
subject wore only a swim cap and Speedo or Lycra spandex-
RE test was preceded by
a 10-minute warm-up at 4 mph (107.3 m$min21) at 1%
grade. Running economy was assessed by having the sub-
jects run for 6-minute intervals at 6 and 7.5 mph, respectively
(160.9 and 201.2 m$min21). The submaximal running speed
was set to no higher than 201.2 m$min21 to make sure both
groups ran at their steady-state oxygen consumption (25)
and below their lactate threshold (60–90% of V_ O2max),
where RE has been shown to be independent of running
speed (32). A 1% incline was used to mimic outdoor con-
ditions (38). Subjects had a 5-minute recovery period (stand-
ing fully connected to the metabolic cart) between the 2
intervals to make sure the intervals were performed at sub-
maximal intensities. Serial, 30-second V_ O2 collections were
made for each 6-minute interval. Running economy was
defined as the steady-state oxygen consumption in
ml$kg21$min21 obtained during each 6-minute workload
and was measured as the average per minute V_ O2 during
the last 3 minutes under steady-state conditions. V_ O2 was
also normalized to mL$kg20.75$min21 (7,63). To allow for
comparison between treadmill speeds, RE was normalized
per distance covered by dividing the velocity (km$min21)
into the V_ O2 (ml$kg$min21) consumed at the submaximal
velocity (66). Metabolic rate during running was calculated
as gross metabolic cost (resting oxygen uptake plus cost of
movement) of steady-state treadmill running, divided by
oxygen consumption (ml$kg$min21).

V_O2peak Test. All participants performed a graded exercise

Figure 1. Relationship between Hcon:Qecc at 1208$s21 and respiratory
exchange ratio (RER) among both HT and REC groups (n = 17) at
201.2 m$min21. One subject did not reach steady state and is therefore not
displayed in the graph. The graph denotes HT runners with black diamonds
_ O2 = oxygen cost of running.
and recreational runners with gray squares. V
the TM
test to volitional exhaustion to determine V_ O2peak after the
last 6-minute interval at 201.2 m$min21. V_ O2peak was deter-
mined using a modified Åstrand incremental treadmill
protocol, which required subjects to run at an increased
grade increment every second minute. The subjects were
instructed to run to maximal volitional fatigue. The first

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Figure 2. Hecc:Qcon ratios divided by HT and REC groups. HT runners
(n = 7) are displayed in black. REC runners (n = 11) are displayed in
gray. *HT runners (n = 7) had significantly (p # 0.05) higher Hecc:Qcon at
120 and at 1808$s21 than REC runners (n = 11). HT = highly trained;
REC = recreational.
2 minutes of each incremental test were run at a 0% treadmill
grade. Treadmill velocity was set at a constant of either 215,
241, or 268 m$min21 (subject’s preference). The grade was
increased by 2% every 2 minutes thereafter until the subject
reached voluntary exhaustion. The running speed was indi-
vidually chosen to limit the exercise period to 6–8 minutes.
Figure 3. The relationship between Hecc:Qcon and V _ O2 (ml$kg20.75$km21)
at 201.2 m$min21 among the study population (n = 17). One subject did
not reach steady state and is therefore not displayed in the graph. The graph
denotes highly trained runners with black diamonds and REC runners with
gray squares. R = 20.30, NS.
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Figure 4. Relationship between Hcon:Qecc at 1808$s21 and
V_ O2 (ml$kg20.75$km21) among both highly trained and recreational
groups (n = 18) at 201.2 m$min21. One subject did not reach steady
state and is therefore not displayed in the graph. The graph denotes HT
runners with black diamonds and REC runners with gray squares. V_ O2 =
oxygen cost of running per unit distance.
Strong verbal encouragement was given throughout the test.
All runners in this study reached voluntary exhaustion
within 6 minutes. V_ O2peak was considered to be the highest
V_ O2 recorded during the test session. To confirm that max-
imal effort had been achieved, 2 of the following criteria had
to be attained: a change in V_ O2 of , 62.1 ml$kg$min21 with
concomitant increase in workload, an RER value $1.10,
attainment of age predicted maximal heart rate (220-age 6
5%), volitional exhaustion, or a rating of perceived exertion
$17. This measure was used for descriptive purposes only.
Flexibility Testing
On visit 2, the sit-and-reach test was conducted after
a 10-minute warm-up, running at a self-selected speed on
the treadmill. The subjects were instructed to jog at
a comfortable pace. Stretching or any kind of flexibility
exercises were prohibited before initiation of the sit-and-
reach test to minimize interindividual variability with the
standardized protocol. The sit-and-reach test was conducted
using a specially designed sit-and-reach box (Acuflex; Novel
Products Inc., Rockton, IL, USA).
The subjects were seated on the floor, knees extended and
legs together, with their bare feet against the sit-and-reach
box (908 in the ankle joints), keeping the palm of their right
hand on top of the left hand. The subjects were then in-
structed to slowly reach toward their toes (zero point on
the sit-and-reach box), and beyond if possible, while pushing
the slide as far forward on the sit-and-reach box as possible
without flexing their knees. The subjects were asked to hold
this stretch for a minimum of 2 seconds at the furthermost
point reached. The most distant point reached with their
VOLUME 28 | NUMBER 8 | AUGUST 2014 | 2219
 Functional Hamstring:Quadriceps Ratios and Running Economy

a full explanation of the procedures, subjects were seated

TABLE 5. Correlation coefficient steady-state V_ O2
(ml$kg20.75$km21) and RER at (1% grade) vs.
physiological variables for both groups (n = 17).*
R vs.
V_ O2201.2
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upright on the dynamometer with their hip joint at approx-
imately 908 flexion, and the rotation axis of the dynamometer
was visually oriented with the participant’s lateral condyle at
the right knee. The input axis of the dynamometer was
R vs. aligned with the axis of the knee. Each subject was fastened

Variables RER201.2 with restraining straps used to prevent unwanted movements

Hecc:Qcon at 608$s21 20.15 20.34 in accordance with the Biodex User Guide (Biodex Pro Man-
20.30 20.38 ual, Applications/Operations; Biodex Medical Systems).
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Hecc:Qcon at
1208$s21 The range of motion (ROM) of the knee was set to 0–908.
Hecc:Qcon at 20.28 0.10 To restrict any undesirable (lateral) movements and to main-
1808$s21
Hcon:Qecc at 608$s21 20.45 20.38 tain body posture, a thigh strap was applied over the test leg,
Hcon:Qecc at 20.33 20.66† and the subjects were instructed to grip the handles on each
1208$s21 side of the Dynamometer while performing the test. Only
Hcon:Qecc at 20.48† 20.24 the right leg was tested.
1808$s21 Gravity correction was attained by measuring the gravity
Sit-and-reach test 20.12 0.40
effect torque, that is, the flexor torque applied on the
_ O2201.2 = gross oxygen cost of running at 201.2
*V dynamometer arm with the knee in a relaxed state at full
m$min21; RER201.2 = respiratory exchange ratio at extension. The thigh strap was released between each trial to
201.2 m$min21; R = correlation.
†p # 0.05 significant correlation. ensure adequate blood flow to the thigh muscles. The testing
protocol for each subject consisted of both concentric and
eccentric muscle actions, respectively. Before the testing
intervention, subjects did a 5-repetition (Con/Con) warm-up
at 608$s21 to ensure familiarity with the isokinetic mode of
fingertips was taken as the measure for the sit-and-reach test.
muscle action.
The best score from 3 attempts was recorded and used for
Each test trial involved 5 repetitions at the following
statistical analysis.
velocities: 60, 120, and 1808$s21, respectively. The Con/Con
muscle actions were performed with extension followed by
Isokinetic Strength Testing. Muscle test analysis was measured flexion. Sixty seconds of rest between sets has been shown to
using a Biodex System 3 dynamometer (Biodex Medical be adequate for recovery following a set of isokinetic exercise
System, Shirley, NY, USA) interfaced with a computer. After (53). A 90-second rest period was required between testing
of each velocity within a set, and a 240 seconds recovery
period between trials to make sure the subjects were fully
recovered. Subjects were asked to perform the movement at
their maximal effort. Every effort was made to ensure that
the subjects made genuinely maximal contractions for each
trial. Verbal encouragement was given throughout the test-
ing session for all subjects to attain maximum performance
(49). The repetition resulting in the greatest maximal peak
torque for the hamstring and quadriceps was used for data
analysis. Peak torque was expressed in absolute values (N$m)
and relative to bodyweight (N$m$kg21).
The strength data were used to calculate both the
functional H:Q (f-H:Q) and conventional H:Q peak torque
ratios (c-H:Q). The f-H:Q was calculated as maximal
eccentric hamstring to concentric quadriceps moments
(Hecc:Qcon, representative for knee extension), or as maximal
concentric hamstring divided by maximal eccentric quadri-
ceps moments (Hcon:Qecc, representative for knee flexion)
(1,2). The f-H:Q ratio is thought to be more appropriate
for strength evaluation than the c-H:Q ratio using concentric
torques of both muscle groups, as it better reflects the true
Figure 5. Sit-and-reach flexibility by groups. HT runners displayed in black
(n = 7) and REC runners in gray (n = 11). HT = highly trained; REC = agonist-antagonist muscle interaction (1).
recreational. Isokinetic strength assessment has demonstrated high
reliability and validity for measuring muscular peak torque
the TM

2220 Journal of Strength and Conditioning Research

Copyright © National Strength and Conditioning Association Unauthorized reproduction of this article is prohibited.

at velocities below 3008$s21 (21). The isokinetic dynamom-
eter controls velocity of movement, as it provides accommo-
dating resistance throughout a joint’s ROM, and it is a tool
that can easily be used to assess dynamic muscle actions in
research and clinical settings (21).
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Allometric Scaling
Conventionally, a standard ratio of oxygen uptake has been
denoted in milliliters per kilogram per minute to compare
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individuals with different body mass. However, as neither
maximal aerobic capacity nor submaximal oxygen uptake
increases at the same rate as body mass (7,69), and to be mass
independent, allometric scaling is recommended when assess-
ing both submaximal and maximal aerobic capacities (11).
Body mass raised to the power of 1 (per kg body mass) under-
_ O2max) and overesti-
estimates maximal aerobic capacity (V
mates submaximal aerobic capacity (RE) in heavier
individuals. For lighter individuals, measurements conducted
without allometric scaling will overestimate maximal aerobic
capacity and underestimate submaximal aerobic capacity.
Thus, to reduce the influence of body mass on oxygen uptake
during running, research suggests that V_ O2 should be scaled to
body mass to the power of 0.75 (ml$kg275$min21) (7,31,58).
Accordingly, oxygen uptake values are presented to the power
of 0.75 in the present study.
Statistical Analyses
Descriptive and inferential statistics were performed. Data
are presented as mean and SD unless otherwise specified.
The Shapiro-Wilk test of normality revealed that the data
were normally distributed, and all had an observed power
greater than 0.95. Once the assumption of normality was
confirmed, parametric tests were performed. Conventional
and functional H:Q ratios were calculated for both trained
states. Student’s unpaired t-tests were used to examine pos-
sible differences between groups (HT vs. REC) on physical
and physiological variables. Pearson’s product-moment cor-
relations (r) were used to evaluate relationships between
dependent variables within and between the 2 groups: (a)
strength ratios and RE and (b) anthropometrics and RE.
Further correlation analysis between demographic and other
possible relevant physiological variables and RE were also
investigated. In all cases, an a priori alpha level of p # 0.05
was used for statistical significance in 2-tailed tests. The data
analysis was performed using the software program SPSS
(version 17.0, SPSS Inc., Chicago, IL, USA).
RESULTS
Demographics
The results of the maximal cardiovascular treadmill tests are
listed in Table 2. There were no significant differences
between groups regarding age, BMI, or body mass (p .
0.05) (Table 1). No significant differences were found
between the 2 groups in the amount of self-reported strength
training, in the number of subjects that performed
hamstring-emphasized strength exercises as part of their
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Journal of Strength and Conditioning Research | www.nsca.com
strength training, nor in the use of hormonal contraception
(p . 0.05). Comparison of anthropometric variables, using
whole group correlations, showed a positive relationship
between calf girth and submaximal V_ O2 consumption
(ml$kg20.75$min21) at 201.2 m$min21 (R = 0.57, p #
0.05). No correlations were found between the anthropo-
metric variables body mass, stature, BMI, leg length, ankle
circumference, and RE (ml$kg20.75$min21) (p . 0.05). A
negative relationship was found between the total miles
run per week and submaximal V_ O2 consumption
(ml$kg20.75$min21) at 201.2 m$min21 (R = 20.55, p #
0.05). V_ O2peak was significantly higher in the HT runners
than among the REC, when normalized per kg21 (p # 0.05)
and when expressed per kg0.75 (p # 0.05) (Table 2). The HR
at 201.2 m$min21 expressed as percentage of HR at V_ O2peak
correlated inversely with RE (R = 20.51, p # 0.05). No
significant correlation was found between V_ O2peak and
f-H:Q strength (p . 0.05).
Oxygen Cost of Running
Results of the steady-state measures are listed in Table 3. The
HT had a significantly lower submaximal V_ O2 at 160.9 and
201.2 m$min21 when expressed as ml$kg20.75$min21 (6.5
and 8.8% difference, respectively, p # 0.05). The intensity
levels for the 2 submaximal speeds such as 160.3 m$min21
and 201.2 m$min21 were on average lower for the HT, as
confirmed by significantly lower % V _ O2peak, % HRmax, and
rate of perceived exertion (RPE) (Table 3). Rating of perceived
exertion was 11.8% lower among the HT at both submaximal
treadmill velocities, confirming their higher fat utilization com-
pared with the REC runners. Whole group correlations re-
vealed a significant positive relationship between calf
circumference and miles run per week (R = 0.57, p # 0.05)
and a significant negative relationship between miles run
per week and RE at 201.2 m$min21 ml$kg20.75$min21
(R = 20.55, p # 0.05).
H:Q Ratios
H:Q ratios between groups are shown in Table 4. The aver-
age percent difference between the 2 trained states for the 3
velocities at Hecc:Qcon, Hcon:Qecc, and Hcon:Qcon were 15.5,
11.8, and 15.7%, respectively. There were no significant dif-
ferences in Hcon:Qcon ratios at 120 and 1808$s21 among the
HT vs. REC runners (p . 0.05). Hecc:Qcon was significantly
higher among the HT than among the REC subjects at 120
(p # 0.05) and at 1808$s21 (p # 0.05) (Figure 2).
Whole group correlations demonstrated a significant
negative correlation between Hcon:Qecc 1208$sec21 and
RER at 201.2 m$min21 (R = 2.66, p # 0.05) (Figure 1).
There was a negative but non-significant relationship
between Hcon:Qecc at 608$sec21 (R = 20.45, p . 0.05) and
at 1208$sec21 (R = 20.33, p . 0.05) and V_ O2 consumption at
201.2 m$min21 (mL$kg20.75$km1 ) (Figure 2). However, the
relationship between Hcon:Qecc at 1808$sec21 and V_ O2 con-
sumption at 201.2 m$min21 (mL$kg20.75$km21) was signif-
icant (R = 20.48, p # 0.05). No significant correlation was
VOLUME 28 | NUMBER 8 | AUGUST 2014 | 2221
 Functional Hamstring:Quadriceps Ratios and Running Economy
found between Hcon:Qcon and RE at 201.2 m$min21 (R =
20.07, 20.06, 20.16, p # 0.05) at 60, 120, and 1808$sec21,
respectively.
There were also no significant whole group correlations
found between Hecc:Qcon and submaximal V_ O2 (RE) at 160.9
m$min21 and 201.2 m$min21 (Figure 3; Table 5). Hecc:Qcon
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608$sec21 was significantly negatively correlated with RER
at 160.9 m$min21 (R = 0.49, p # 0.05), but not at 201.2
m$min21 (R = 20.34, p . 0.05) (Table 5).
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Whole group correlations demonstrated that RER at
201.2 m$min21 was significantly negatively correlated with
km per week of running (R = 20.65, p # 0.05) and
RE (ml$kg20.75$km21) (R = 20.55, p # 0.05). Calf girth
was positively correlated with RE (ml$kg20.75$km21) at
201.2 m$min21 (R = 20.57, p # 0.05).
Peak Torque
When expressed in absolute values (N$m), we did not find
any significant differences between the 2 groups regarding
peak torque at any of the 3 velocities, in either knee flexion
eccentric or concentric muscle actions (p . 0.05). The aver-
age knee flexion eccentric and concentric peak torque for the
isokinetic velocities were 148.8 6 24.6 vs. 134.6 6 23.0 N$m
and 71.7 6 12.0 vs. 64.6 6 13.7 N$m for HT and REC
runners, respectively. The average knee extension eccentric
and concentric peak torque values for HT and REC runners
were 205.7 6 42.8 vs. 210.1 6 52.2 N$m and 111.7 6 15.3 vs.
118.5 6 27.5 N$m, respectively.
Peak torque normalized to body weight (N$m$kg21) re-
vealed no significant differences between groups at 608$s21
(p . 0.05). However, there were significant differences
between the 2 groups (HT vs. REC) in knee flexion at
1208$s21 (1.3 6 0.2 vs. 1.0 6 0.1 N$m$kg21, p # 0.05,
respectively) and 1808$s21 (1.2 6 0.1 vs. 0.9 6
0.1 N$m$kg21, p # 0.05, respectively). There were no sig-
nificant differences in maximal isokinetic knee extension
torque normalized to body weight among the 2 groups at
any of the 3 velocities (p . 0.05). No significant relation-
ships were found between hamstring peak torque expressed
in absolute (N$m) or relative values (N$m$kg21) and RE
(p . 0.05).
Flexibility
Although the HT runners (n = 7) scored 13.2% (29.6 vs.
33.8 cm) lower on the sit-and-reach test than the REC female
runners, there were no significant differences in sit-and-reach
measures between groups (p . 0.05). There was no significant
correlation between submaximal V_ O2 and sit-and-reach flexi-
bility at any of the submaximal treadmill speeds (Table 5).
When normalized per kilometer (ml$kg20.75$km21) among
HT and REC runners (n = 17), there were weak negative
correlations between submaximal V_ O2 at 201.2 m$min21
and flexibility (R = 20.12, p . 0.05). Finally, when comparing
RER vs. flexibility among all groups at the 2 submaximal
velocities, there was a weak positive correlation (R = 0.40,
p . 0.05) (Figure 5, Table 5).
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2222 Journal of Strength and Conditioning Research
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DISCUSSION
The main findings of this study can be summarized as follows:
(a) RE was significantly positively correlated with f-H:Q ratios
(Hcon:Qecc) at both 120 and 1808$s21, (b) Hecc:Qcon was sig-
nificantly higher among the HT compared to the REC female
runners at both 120 and 1808$s21, (c) neither absolute knee
extensor torque nor knee flexor torque was significantly
correlated with RE, and (d) no statistically significant relation-
ship was found between RE and flexibility. In addition, the
HT female runners had a trend toward consistently higher
f-H:Q at all 3 angular velocities (60, 120, and 1808$s21).
As hypothesized, the runners with higher f-H:Q ratios
tended to be more economical at a submaximal velocity. Our
results indicate that relative muscle strength (higher f-H:Q
ratios) may be an important factor regulating the metabolic
cost of running, and therefore a determinant of success in
distance running. The results may have functional implica-
tions for future directions in the assessment of strength
profiles among runners, and in planning and developing
adequate training programs to improve running performance.
Commonly, the conventional ratio (c-H:Q ratio) is reported
to be 0.50–0.80 in healthy subjects (15), that is, the hamstrings
have approximately 50–80% of the quadriceps strength
depending on angular velocity, test position, population
group, and use of gravity correction. Studies have consistently
demonstrated that when tested as conventional isokinetic tor-
que, hamstring to quadriceps torque ratios in women do not
increase at velocities that approaches those of functional activ-
ities, whereas male hamstring to quadriceps torque ratios
increase as velocity increases (33). Nevertheless, the results
from c-H:Q ratios at different velocities in this study were
not consistent with the analyses by Hewett et al. (33). In
our study, all subjects’ c-H:Q ratios increased from slow to
fast velocities. In contrast to our study’s healthy cohort, the
results found in the meta-analysis by Hewett et al. (33) may
reflect the lack of noninjured, well-trained subjects in previous
investigations. Future studies should therefore investigate the
potential impact of both injury and training status on c-H:Q
and f-H:Q ratios among large cohorts to determine the rela-
tionship between these variables.
The precise mechanism whereby the results of this study
were obtained can only be speculated on, that is, significant
differences in f-H:Q ratios between the 2 trained states, and
the positive correlation between increased f-Hcon:Qecc ratios
with greater RE. Running involves muscle work, which
consists of a combination of eccentric muscle contraction
followed immediately by a concentric muscle contraction.
This process is commonly referred to as the stretch-shorten
cycle (SSC). Several studies link increased joint and muscu-
lotendinous stiffness to better RE (5,18,61,66). Because the
elastic contributions are greater during eccentric muscle
actions than concentric actions, f-H:Q (Hecc:Qcon) ratios are
more forceful (1). The force generated during a prestretched
muscle is added to the force in the subsequent concentric

contraction, without a proportionate required metabolic cost.
Thus, the SSC generates more force and is more efficient
(more work per unit of metabolic energy input) than pure
concentric contractions. Higher Hecc:Qcon among HT when
performing eccentric muscle work can be explained by pre-
vious research finding that muscle output in high-force iso-
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metric and dynamic muscle actions is positively related to the
stiffness of the tendinous structures (8). In addition, our data
indicate a trend toward differences in musculoskeletal flexibil-
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ity between the HT and REC runners. It is possible that the
sit-and-reach test in our study was not sufficient to detect any
real differences in musculoskeletal flexibility among the HT
and REC female runners. It has been suggested that inflexi-
bility in certain areas of the musculoskeletal system may lower
the aerobic demand of running by increasing storage and
return of elastic energy during the SSC, and reducing the need
for muscle-stabilizing activity (17).
In addition to the possible contribution of strain energy
among the HT runners, their significantly greater ratios
compared with the REC runners may reflect natural
selection, biomechanics, or training background. It is also
possible that the difference in running routine alone between
the 2 trained states in our study has resulted in greater f-H:Q
ratios among the HT female runners. The HT had no
differences in anatomical traits from the REC runners in the
present study, except for longer legs and greater leg length
relative to height (Table 1). This might be a biomechanical
advantage and therefore a possible explanation for the
greater torque production in the HT runners in our study.
We did not find any significant differences in absolute
muscle strength in our study between the 2 groups.
Although hamstring peak torque normalized to body weight
(N$m$kg21) was significantly different between the 2 groups
at medium and high angular velocities, no significant rela-
tionship was found between hamstring absolute or normal-
ized peak torque and RE. From our study, the low-energy
expenditure among the HT may partly be related to the
higher f-H:Q ratios of the subjects’ hamstring muscles but
not absolute strength. It is known that runners with a strong
musculoskeletal apparatus tend to exhibit more stable run-
ning styles, which seem to increase muscular efficiency,
thereby allowing the athletes to run long distances at max-
imal aerobic speed (26). At increasing speeds, the hamstring
muscles’ biceps femoris activity electromyography (EMG) is
highly correlated with energy expenditure (42). Poor tech-
nique, in turn, may partially be caused by vertical medio-
lateral and braking forces, resulting from insufficient
hamstring muscle actions (42). In the study by Kyrolainen
et al., the treadmill velocities were higher than in our study.
In addition, we did not measure EMG activity in our study.
It is possible, however, that higher treadmill speeds would
have revealed correlations between absolute knee flexor
strength and RE.
As anticipated, functional Hecc:Qcon ratios were higher
than the conventional ratios (c-H:Q) for both groups and
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Journal of Strength and Conditioning Research | www.nsca.com
for each individual. Eccentric muscle action, that is, muscle
lengthening, is an important factor in stability and energy
saving when running (59). Perhaps the higher f-H:Q ratios
in HT runners in our study can be explained by their
increased eccentric strength, because of more elasticity and
stiffness, which affects eccentric hamstring torque. This can
be explained by the force-velocity curve, which shows that
eccentric contraction generates higher forces than concen-
tric contractions. Also, increased angular velocity results in
greater force when contracting eccentrically, while the
opposite is true when contracting concentrically (1).
Previous investigations in nonathletes found that the
functional Hecc:Qcon ratio produced a 1:1 hamstring to quad-
riceps strength relationship for fast knee extension (1). To
the best of our knowledge, only 1 study has previously inves-
tigated f-H:Q ratios in runners (40). Interestingly, the study
by Kong and de Heer (40) investigating anthropometric var-
iables in 6 highly trained male, sub-elite, Kenyan runners
(40), found Hecc:Qcon ratios corresponding to the Hecc:Qcon
ratios for the HT female runners in the present study, with
f-H:Q ratios greater than 1.0 at all angular velocities. Taken
together, the results by Kong and de Heer and our study
results indicate that f-H:Q ratios are similar between the
sexes for distance runners who are at the same relative per-
formance level, and that runners exhibit a higher f-H:Q
ratios (Hecc:Qcon) than athletes in other sports (1,35). How-
ever, the study by Kong and de Heer (40) involved only 6
athletes, did not include a control group, and the authors did
not test for V_ O2 consumption, blood profile or musculoskel-
etal stiffness in any of the muscle measurements, making it
impossible to elucidate any relationship between f-H:Q and
endurance performance. In fact, an intervention study (35)
on 12 female soccer players after 6 weeks with hamstring-
emphasized strength training reported average post-test
Hecc:Qcon ratios for both legs combined to be lower than
that found for both the HT and REC runners in the present
study. However, the soccer players in the same study had
mean c-H:Q ratios higher than the HT and REC runners in
our study. It should be assumed that soccer players should
display both lower c-H:Q and lower f-H:Q ratios than run-
ners considering the soccer players’ more dominant quadri-
ceps muscles from the kicking motion. In sum, the results in
our study suggest that subjects who run long distances have
higher functional Hecc:Qcon strength ratios than subjects in
other sports, which could possibly be due to weaker quad-
riceps muscles or stronger hamstring muscles in runners, or
a combination of these two.
The lack of significant relationships between the isokinetic
measures for Hecc:Qcon and RE were not unexpected, con-
sidering the limited specificity using this type of assessment,
and the muscle mechanics during treadmill running vs. over-
ground running. The main reasons for the lack of strong
correlations for Hecc:Qcon and RE may be due to the low
treadmill speed, the fact that the testing was performed on
a treadmill, rather than overground running, or simply
VOLUME 28 | NUMBER 8 | AUGUST 2014 | 2223
 Functional Hamstring:Quadriceps Ratios and Running Economy
because the isokinetic strength tested may not reflect the
mechanical requirements experienced while running.
Running economy has been shown to be independent of
running speed, as expressed per distance covered, for well-
trained distance runners within a range of 60–90% of
V_ O2peak (32). The well-trained women in the study by
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Helgerud et al. (32) had a higher average V_ O2max (157.8 6
17.8 ml$kg20.75$min21) than reported in the present study
for the REC runners but lower than for the HT runners in
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our study. This makes it difficult to assess whether the V_ O2
intensity was valid at higher velocities for the REC runners
in the present study. The HT reached the 60–90% V_ O2 inten-
sity range at 201.2 m$min21. Hence, the submaximal V_ O2
measures in the present study are deemed to be representa-
tive for RE measured at race speeds for distances from
10,000 m and above for the HT (32).
Respiratory exchange ratio at 201.2 m$min21 was signif-
icantly negatively correlated with Hcon:Qecc at 1208$s21
when analyzed for the whole group (n = 17), possibly indi-
cating that other measures of the metabolic demand of run-
ning might have resulted in more accurate measurements of
RE than those used in the present study. Functional H:Q,
measured as Hcon:Qecc at 1808$s21, was significantly
correlated with submaximal V_ O2 (ml$kg20.75$min21) at
201.2 m$min21, suggesting that strong knee flexion may be
important for running at low oxygen cost.
Contrary to previous reports (17,27,37,66), and our initial
hypothesis, there was no significant relationship between
oxygen demand at submaximal running and flexibility in
the present study, although HT runners had lower flexibility
scores than less economical REC runners, as confirmed by
the sit-and-reach test. This result is in accordance with
a study by Craib et al. (17), which found no significant cor-
relation between RE and sit-and-reach flexibility (r = 0.12),
despite significant correlation between RE and inflexibility in
the ankle and the hip. However, the results by Craib et al. are
not in line with previous findings relating sit-and-reach flex-
ibility to better RE in both male and female collegiate dis-
tance runners (66), and among internationally competitive
male long-distance runners (37). These latter studies support
the theory that both muscle strength and stiffness are related
to improved performance. Perhaps running in and of itself
causes stiffer and more economical locomotion, or the HT
athlete’s inflexibility and outstanding RE in the present study
can be explained partly by the COL5A1 genotype, which
recently has been associated with both a subject’s ROM (13),
and endurance performance (10,55) in separate studies.
The reason for this inverse relationship possibly relates to
the stretch and recoil of tendon and muscle springs, which
adds mechanical work, while active, energy-demanding
muscle fibers produce the high forces necessary for momen-
tum (56). Thus, muscles act as active struts rather than pure
working machines (56), making this storage and return of
elastic-strain energy an important energy-saving mechanism
for running (3). Also, storage of elastic energy in the con-
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2224 Journal of Strength and Conditioning Research
Copyright © National Strength and Conditioning Association Unauthorized reproduction of this article is prohibited.
tracted muscle during eccentric muscle actions is an impor-
tant mediator to increase the total work output during
muscle contractions (6). As a result, athletes who are able
to use more of this highly efficient energy source would
clearly have an advantage as their metabolic cost of move-
ment is lower. Lending support for this theory, optimal mus-
culotendinous stiffness for maximum concentric and
isometric activities has been shown to be toward the stiff
end of the elasticity continuum (72), meaning that a stiffer
muscle may enhance the force production capabilities of the
contractile units. Thus, muscle stiffness, in addition to
strength, may be an important factor explaining differences
in endurance determinants such as RE (43).
The large differences in average distance run per week
between the HT and REC (104.6 vs. 32 km$wk21, p # 0.05)
may explain the lower oxygen demand of running among
the HT runners. It is possible that higher treadmill velocities
might have revealed even larger differences. The lack of
correlation between RE and f-H:Q ratios at the lower angu-
lar velocities among the HT and REC in the present study
can also be explained by the fact that the percentage of
energy expenditure due to fat oxidation at any given absolute
running speed is inversely related to V_ O2max (16). The per-
centage of energy expenditure due to fat oxidation at any
given common running speed increases with endurance
training (12), and a higher V_ O2max is related to higher fat
oxidation (47). Moreover, at any given submaximal speed,
runners with a higher V_ O2max may require higher V_ O2 due
to their greater reliance on fat utilization (24). Thus, the HT
females in this study may spend more V_ O2 relative to the
REC female runners at the submaximal paces, thus not re-
flecting their “true” RE for the HT. Support for this view can
be found in the lower RER and % V_ O2peak among the HT in
this study compared with the REC runners. As a result, the
HT runners’ RE may be considered “poor” simply because of
the additional oxygen required to metabolize fat as opposed
to carbohydrate. Thus, the lack of a larger difference
between the trained groups possibly indicates that submax-
imal V_ O2 was a poor indicator of RE in the present study.
The respiratory exchange ratio did not increase between
the 2 speeds for the HT, and there was a significantly lower
RER between HT and REC runners at the highest sub-
maximal speed, indicating that it was a low intensity for
trained runners. Respiratory exchange ratio increased only
among REC runners from 160.9–201.2 m$min21 (4.7%
increase in RER), indicating that intensity affected substrates
metabolized during the two 6-minute intervals among the
REC runners but not in the HT runners. Respiratory
exchange ratio at 160.9 m$min21 was significantly inversely
related to Hecc:Qcon at 608$s21, reflecting that RER might be
a better indicator of RE than submaximal V_ O2 in the present
study. The lack of larger differences in submaximal V_ O2
between groups may also explain why the correlation coef-
ficients between H:Q strength ratios and RE were higher for
the RER and f-H:Q strength relationship.

The lack of a significant correlation between Hecc:Qcon
ratios and submaximal V_ O2 was expected, as both treadmill
velocities might have been too slow to reach the threshold at
which hamstring eccentric strength contributes significantly
to the aerobic demand of running. Considering the greater
active lengthening contraction (eccentric work) of the ham-
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string during the late swing phase at higher velocities (64)
and the accompanying increased activation in the hamstring
muscles with increased treadmill speed (34,41), it is possible
CX1AWnYQp/IlQrHD3i3D0OdRyi7TvSFl4Cf3VC4/OAVpDDa8K2+Ya6H515kE= on 03/23/2023
that testing RE at higher velocities would reveal greater
differences between an individual’s hamstring muscle
strength and the H:Q ratios. This would perhaps result in
a higher correlation between H:Q ratios and RE, as well as
other variables related to RE.
Another possible explanation for the lack of a significant
correlation between H:Q ratio strength and submaximal V_ O2
at all the tested velocities in the present study could be the
difference in technique for treadmill running vs. overground
running. Overground running may place greater demands on
the hamstring muscles to produce propulsive forces than
treadmill running (39). Wank et al. (68) found greater elec-
tromyographic activity of the biceps femoris muscle during
treadmill running compared with overground running. Thus,
whether treadmill results are transferable to overground run-
ning remains unclear. In the present study, the relatively low
treadmill speed may not have reached the threshold level at
which hip extensors, which include the hamstring and glu-
tes, become more important for propulsion forces. The sig-
nificant differences between the 2 groups at the medium and
fast velocities may reflect the HT runners’ tendency to per-
form more race-specific workouts at higher intensities than
REC runners; for example, high-intensity interval training on
a track. This type of activity would target the hamstring
muscles specifically at higher velocities. Moreover, Koller
et al. (39) found a significant decrease in hamstring, but
not quadriceps, eccentric strength after overground running
(running a marathon), implying that the 2 muscle groups
may be exposed to a different level of fatigue during distance
running. These large muscle groups’ association with RE
may be explained by their high activation rate during run-
ning for propulsion and support (28), which makes them
consume a substantial amount of oxygen during locomotion.
There are 2 main ways of changing the H:Q ratios: (1)
by increasing hamstring torque and (2) by decreasing
quadriceps torque. The reasons for the higher H:Q ratios
among the HT runners in the present study were a combi-
nation of stronger hamstrings and weaker quadriceps
muscles in the HT vs. REC runners. The hamstring
muscles among the HT had higher peak torque at both
concentric and eccentric hamstring muscle actions. The
opposite was true for the quadriceps muscles; the REC
runners had higher values than the HT runners at all angu-
lar velocities. There were no differences in the amount of
self-reported strength training sessions between groups in
terms of frequency or hours per week. Nevertheless,
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Journal of Strength and Conditioning Research | www.nsca.com
among those HT and REC runners who reported that they
did strength training, the HT performed more specific
hamstring exercises per week.
In conclusion, this is the first study to investigate the
relationship between relative muscle strength in the thigh
muscles and RE. Highly trained female runners had signif-
icantly higher f-H:Q ratios at velocities approaching those
that occur during running. The f-H:Q ratios observed for all
the HT female runners indicated a significant functional
capacity of the hamstring muscles for providing muscular
stability at the knee joint in fast knee extension compared
with the REC runners. Taken together, the results from our
study suggest that relative hamstring to quadriceps muscle
strength may be a contributing factor for RE. As a final
point, the HT leg strength was lower compared with the
REC runners, despite higher f-H:Q ratios. Therefore, run-
ning performance in long-distance events, at least to a certain
level, may be related to greater hamstring muscle strength
relative to quadriceps muscle strength and not to absolute
muscle strength per se.
PRACTICAL APPLICATIONS
From a practical viewpoint, our results indicate that thigh
muscle strength distribution (relative concentric to eccentric
hamstring muscle strength), and not muscle strength per se,
may have a regulatory role in determining RE. In fact, no
significant correlation was found between RE and isolated
quadriceps or hamstring peak torque were found. Because of
the significantly higher f-H:Q and the significant relationship
between f-H:Q and RE, strength training to improve this
ratio should be considered for middle-distance and long-
distance running events to further lower the oxygen cost of
running (i.e., improve RE) and thereby, improve perfor-
mance. As running is basically a series of horizontal jumps
requiring a strong and highly efficient extensor apparatus,
and improved plyometrics has shown to be related to greater
RE, we suggest that runners should aim to include hamstring
muscle strengthening exercises that imply horizontal
motions.
Our results suggest that coaches and athletes in middle-
distance and long-distance running should focus on increas-
ing the functional hamstring:quadriceps muscle strength
ratio by implementing additional conditioning exercises or
exercises specifically targeting the hamstring muscles. These
exercises should primarily focus on eccentric hamstring
muscle actions and may include fast downhill running,
over-speed running, hill bounding, drills, as well as resistance
or plyometric.
ACKNOWLEDGMENTS
The authors are grateful to the 21 runners who took part in
this study. For their assistance and cooperation with this
research, the authors thank Gonzalo Villablanca, Morgan
Tapper, Alicia B. Olsen, and Annie Peck. The authors thank
Annie Bersagel and Håkon Brox for their valuable comments
VOLUME 28 | NUMBER 8 | AUGUST 2014 | 2225
 Functional Hamstring:Quadriceps Ratios and Running Economy
on drafts of this article. The authors have no funding or
conflicts of interest to disclose.
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