Journal of the Neurological Sciences 174 (2000) 9299

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Neuromuscular disturbance outlasts other symptoms of exercise-induced
muscle damage
a , a b a c
*
Michael R. Deschenes , Rhonda E. Brewer , Jill A. Bush , Raymond W. McCoy , Jeff S. Volek ,
c
William J. Kraemer
a
Department of Kinesiology, The College of William & Mary, Williamsburg, VA 23187-8795, USA
b
Department of Kinesiology, The Pennsylvania State University, University Park, PA 16802, USA
c
The Human Performance Laboratory, Ball State University, Muncie, IN 47306, USA
Received 26 July 1999; received in revised form 13 October 1999; accepted 22 December 1999
Abstract
This study examined the biochemical, immunological, functional, and neuromuscular responses associated with exercise-induced
muscle damage in the quadriceps of untrained men. Muscle damage and soreness was elicited with maximal concentric/ eccentric muscle
21
actions at 0.53 rads s . Signicant (P,0.05) soreness was evident 1, 2, and 3 days following muscle insult, while plasma creatine
kinase, a marker of muscle damage, was elevated 3 and 5 days post-insult. Plasma interleukin-Ib was signicantly increased within 5 min,
and remained elevated 1, 2, 5, and 7 days post-insult. Maximal isometric quadriceps function was impaired (P,0.05) for 5 days following
21
muscle challenge. Maximal isokinetic performance at 1.09 rads s was diminished (P,0.05) for 2 days post-insult; no signicant
21
decrements at 3.14 rads s were noted. Average electrical activation (iEMG) of the quadriceps was unaltered, but iEMG activity of the
rectus femoris where soreness was focused was signicantly increased. Neuromuscular efciency (torque/ iEMG) was compromised
throughout the 10-day post-insult period investigated. While other symptoms of exercise-induced muscle damage dissipate within 7 days,
neuromuscular perturbation persists for at least 10 days. 2000 Elsevier Science B.V. All rights reserved.
Keywords: Creatine kinase; IL-Ib; EMG; Isometric; Isokinetic; Soreness
1. Introduction and is generally indicated by elevated concentrations of
circulating macrophages, monocytes, and interleukins
Participation in unaccustomed physical exertion, par- [11,12].
ticularly that involving eccentric, or muscle lengthening, Such muscle insult is also associated with a diminution
actions typically results in muscle damage [14]. This of maximal force production and localized discomfort
damage is evidenced by ultrastructural damage, i.e. Z-line [13,14], yet these manifestations of exercise-induced mus-
streaming [5,6], and due to sarcolemmal disruption, in- cle damage exhibit disparate temporal characteristics. For
creased blood-borne levels of intramuscular proteins such example, while impaired force production is most pro-
as creatine kinase (CK), lactate dehydrogenase (LDH), and nounced within 24 h [2,13,15], peak soreness is not
myoglobin [710]. Consequent to this damage, an in- reported until 23 days following eccentric work [13].
ammatory response is mounted by the immune system Accordingly, the discomfort accompanying unfamiliar
muscular exertion is referred to as delayed onset muscle
soreness or DOMS. The precise mechanism for the
postponement of discomfort following muscle insult is
*Corresponding author. Tel.: 11-757-221-2778; fax: 11-757-221-
unknown, but it has been suggested that it is related to a
2761.
E-mail address: mrdesc@facstaff.wm.edu (M.R. Deschenes) local inammatory stimulation [12,16]. That is, the release
0022-510X/ 00/ $ see front matter 2000 Elsevier Science B.V. All rights reserved.
PI I : S0022- 510X( 00) 00258- 6
M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299 93
of pain substances such as bradykinin and histamine is tube via venipuncture of an antecubital vein. Whole blood
delayed after the onset of muscle injury. was centrifuged at 30003g for 15 min at 48C; plasma was
Similar to delayed soreness, the exact cause(s) of stored at 2758C until analysis.
attenuated muscle strength following unaccustomed exer- The subject then performed a light (40 W), nonspecic
tion is not yet fully understood. It has been suggested that warm-up protocol on an electrically braked cycle ergome-
the decrement in force production may be due to damage ter (Excalibur Unit, Lode, Groningen, The Netherlands) for
to the contractile elements, impaired electrical activation, 5 min. One min into the warm-up, the subject was asked to
and/ or disruption of the excitationcontraction (EC) rate the level of soreness of the non-dominant thigh. If any
coupling process [17]. soreness was indicated, the subject was asked whether the
To date, the investigation of the effects of, and recovery discomfort was more pronounced in the inner, middle, or
from, exercise-induced muscle damage has mainly em- outer region of the quadriceps.
ployed an animal model [7,17,18], or in humans, a small, The subject was then situated on the isokinetic device
seldom used muscle, i.e. biceps brachii [1,2,14,19], or the with the knee aligned with its axis, to test muscle function
quadriceps following submaximal exertion [8,9,11,20]. The of the non-dominant leg. In preparation for concurrent
aim of the present study was twofold: (1) to examine the isometric testing and electromyographic (EMG) analysis,
effects of exercise-induced muscle trauma in a large, the involved knee was positioned at a 958 angle, and a one
commonly used muscle mass (quadriceps) in humans square inch area of skin over each of the three surface
following a maximal, voluntary effort, and (2) to assess the quadriceps muscles (vastus medialis, rectus femoris, vastus
impact of muscle insult on neuromuscular function over an lateralis) was shaved, abraded, and cleansed with an
extended period of time. alcohol wipe. Following the longitudinal contour of the
muscle, 2-mm-diameter electrodes lled with electrolyte
gel were secured on the skin with adhesive collars at an
interelectrode distance of 2 cm, and traced with ink. These
2. Materials and methods
tracings enabled electrode placement at the same sites for
each subsequent test. EMG signals were amplied by a
2.1. Subjects
factor of 1000 and passed through a bandwidth lter set at
30 and 500 Hz, along with a 60-Hz notch lter. Signals
Nine healthy, untrained men (20.961.0 years,
were digitized at a sampling frequency of 1000 Hz, and
174.862.0 cm, 72.366.1 kg; mean6S.E.) agreed to par-
recorded by an on-line computer system during the rst 5 s
ticipate in the study. After receiving a verbal description of
of a 15-s maximal isometric contraction of the knee
the investigation, its potential risks, and the experimental
extensors. The EMG signal was then full wave rectied,
procedures employed, the subjects provided written in-
and integrated (iEMG in mV/ s). Subjects were given a 1-s
formed consent. Each subject also completed a medical
ramp-up phase to achieve full muscle activation before
questionnaire that was examined by a physician to ensure
EMG and force measurements were begun.
that no contraindications to participation existed. All
Following completion of the 15-s isometric contraction,
experimental procedures were approved by the Committee
the electrodes were removed from the subjects leg. In
for the Protection of Human Subjects at The College of
preparation for isokinetic testing, the subjects limb weight
William & Mary.
was measured by the dynamometer so that performance
variables could be corrected for that resistance. A full
2.2. Experimental protocol range of motion was then selected by the subject that was
adhered to for isokinetic testing at each subsequent ses-
Subjects initially performed a familiarization trial on an sion.
isokinetic dynamometer (model 900-350, Biodex Co., Maximal isokinetic muscle function (peak torque, and
Shirley, NY). Since an extended prophylactic effect for total work) was then determined with ve repetition sets of
muscle damage occurs following a single exertional effort alternating, concentric actions of the knee extensors and
that is specic to the musculature involved [15,19,21], exors of the non-dominant leg at velocities of 1.05 and
21
only the subjects dominant leg was exercised during the 3.14 rads s . A rest interval of 5 min occurred between
familiarization trial; the nondominant leg would be used sets, and verbal encouragement was provided during all
during actual data collection sessions. To ensure that any isokinetic, as well as isometric, testing.
potential elevations in circulating markers of muscle After collection of baseline (day 0) data was completed,
damage or inammation were dissipated following this subjects rested for 5 min before carrying out the damage-
exercise, a 1014-day period interspersed the familiariza- inducing event. This muscle insult was presented as four
tion trial from the rst data collection session [22,23]. At sets of 25 repetitions, interspersed by 3-min rest periods, of
the rst test session (day 0) baseline data were collected. unilateral, alternating concentric/ eccentric muscle actions
21
The subjects height and weight were determined, and a of the non-dominant knee extensors at 0.53 rads s . Five
3-ml sample of blood was collected into a heparin-treated min following completion of the concentric/ eccentric work
94 M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299
protocol, another blood sample was drawn, centrifuged, contraction divided by average iEMG during that time,
and plasma was stored at 2758C until analysis. was determined and compared among test sessions.
With the exception of the muscle damaging protocol and
the blood sample taken 5 min after that muscle challenge, 2.4. Statistical analysis
data collection, i.e. blood draw, soreness assessment,
EMG, isometric and isokinetic testing, was repeated (61 Data are reported as means6S.E. To assess changes in
h), at 24 h (day 1), 48 h (day 2), 72 h (day 3), 120 h (day each variable of interest during the experimental period,
5), 168 h (day 7), and 240 h (day 10) after collection of repeated measures one-way ANOVAs were utilized. When
baseline data. For 10 days prior to, and throughout the signicant F-ratios were found, Fisher PLSD post-hoc
study period, subjects were instructed to avoid anti-in- analyses were conducted. In this report, only statistically
ammatory medications, maintain normal sleeping pat- signicant differences from baseline values will be iden-
terns, and to refrain from unaccustomed muscular exertion. tied since deviations from, and resumptions to, pre-insult
For 68 h prior to arriving at the laboratory for testing, conditions were the focus of the study. Statistical signi-
subjects consumed only water. cance was established at P#0.05.
2.3. Quantitation 3. Results
Subjective ratings of muscular soreness were determined The combination of concentric/ eccentric actions of the
with a 010 scale where 05no soreness, 55moderate knee extensors resulted in damage, and soreness of the
soreness, and 105extreme soreness. Plasma CK activities involved muscles. Muscle damage was indicated by in-
were measured in duplicate with a Vitros blood chemistry creased plasma CK activity at days 3 and 5. In addition,
analyzer (model DT 60 II, Johnson and Johnson Clinical signicant elevations in plasma IL-Ib were detected at 5
Diagnostics, Rochester, NY). Plasma concentrations of min post-insult, as well as on days 1, 2, 5, and 7.
interleukin Ib (IL-Ib) were assessed in duplicate with Responses of these blood-borne variables are presented in
ELISA kits (Cytimmune Sciences Inc., College Park, MD). Figs. 1 and 2, respectively.
During the 15-s maximal isometric contractions of knee There was signicant overall soreness of the quadriceps
extensors, peak and total torque, along with fatigue (% at 1, 2, and 3 days following muscle insult, with the
difference in torque from rst to last 5 s) were calculated highest ratings occurring at day 2. When subjects were
by the Advantage software accompanying the Biodex asked to localize where pain was most acute, 74% of the
dynamometer. During isokinetic maximal actions of knee time the central region of the thigh (rectus femoris) was
extensors and exors, peak torque and total work were identied. This was signicantly greater than the inner
ascertained with the Advantage software. In addition to (vastus medialis) or outer (vastus lateralis) regions at 26%
average iEMG values derived from the three surface and 12%, respectively. Soreness of the quadriceps over the
muscles of the quadriceps, iEMG data from each of those 10-day investigation is illustrated in Fig. 3.
muscles were examined individually over the experimental Performance of the quadriceps, during both isometric
period. In addition, neuromuscular efciency, dened as and isokinetic actions was signicantly diminished follow-
total torque generated during the rst 5 s of the isometric ing muscle insult. During the 15-s maximal isometric
Fig. 1. Plasma activities of creatine kinase (CK). Values are means6S.E., n59. *Indicates signicant difference (P,0.05) from day 0.
M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299 95
evidenced no performance decrements (data not shown).
Data regarding quadriceps function are found in Table 1.
Integrated EMG data, averaged among the measured
knee extensor muscles, exhibited no signicant differences
during the study. In addition to this overall iEMG value,
the electrical activation of the rectus femoris, vastus
lateralis, and vastus medialis was examined individually.
Results of these analyses revealed no signicant variation
in iEMG data for the vastus lateralis or vastus medialis
subsequent to the DOMS-inducing event. However, electri-
cal activation of the rectus femoris was signicantly
enhanced at days 2, 3, and 10. The results of iEMG
analyses are presented in Fig. 4.
Neuromuscular efciency (torque/ iEMG) was deter-
mined during the rst 5 s of the maximal isometric
contraction of the quadriceps. This parameter demonstrated
Fig. 2. Plasma concentrations of interleukin Ib (IL-Ib). Values are
a signicant diminution at each of the 10 days studied means6S.E., n59. *Indicates signicant difference (P,0.05) from day 0.
following the collection of baseline data at day 0. Changes
in neuromuscular efciency during the investigation can be
contraction, signicant decrements in peak and total torque
observed in Fig. 5.
produced were evident at days 1, 2, 3, and 5 compared to
baseline values at day 0. Torque generated during the rst
5 s of the isometric contraction demonstrated an identical
4. Discussion
` pattern of disturbance. Specically, vis-a-vis baseline data,
total force produced during the initial 5 s was less at days
The data presented here add to a large body of evidence
1, 2, 3, and 5. Interestingly, no changes in fatigue during
conrming that unaccustomed muscle exertion results in
15-s isometric contractions were demonstrated throughout
muscle damage, and a sequela of associated physiological
the experimental period.
responses. The plasma CK elevations consequent to mus-
Isokinetic data indicate that during ve repetition sets at
21
cle insult noted here are similar to previous studies despite
1.09 and 3.14 rads s , signicant impairments in quad-
the use of high intensity isokinetic resistance exercise to
riceps performance were displayed only at the slower
21
elicit damage and pain in the quadriceps. While other
velocity of movement. At 1.09 rads s , peak torque was
researchers have studied circulating levels of intramuscular
signicantly lower at day 1 compared to day 0, while total
proteins to indicate muscle damage in the knee extensors,
work was signicantly decreased at days 1 and 2 relative
21
they typically employed submaximal efforts such as down-
to baseline values. However, at 3.14 rads s , neither peak
hill running [7,8,20], cycle ergometry [5,6], or bench
torque nor total work produced by the quadriceps varied
stepping [4,9] as the unaccustomed exertion. Maximal
signicantly during the investigation. Knee exors, which
eccentric contractions have been used to investigate circu-
were not subjected to damage-inducing eccentric actions,
lating markers of sarcolemmal damage, however, those
studies focused on the elbow exors [1,14,19]. Despite
differences in muscles utilized, or intensity of muscle
exertion, the temporal responses of blood-borne CK levels
noted here are consistent with those earlier studies. Spe-
cically, elevations in CK activity are most marked 35
days following unaccustomed physical activity.
Muscle soreness manifested itself most acutely 23 days
following muscle insult. The intensity of the discomfort in
the quadriceps reported by our subjects was similar to that
experienced in the elbow exors following eccentric
contractions in previous studies [19,24]. Based upon these
and other investigations [1,4] comparing the effects of
different modes of muscle actions on soreness, it is
reasonable to assume that the eccentric component of our
exercise protocol was principally responsible for the
muscle damage and soreness evinced in our subjects.
It has been postulated [16] that the delay that occurs
Fig. 3. Soreness of quadriceps muscles. Values are means6S.E., n59.
*Indicates signicant difference (P,0.05) from day 0. between muscle insult and the onset of soreness is concor-
96 M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299
Table 1
a
Muscle function of knee extensors
Day 0 Day 1 Day 2 Day 3 Day 5 Day 7 Day 10
Peak torque (N)
21
0 rads s 219.0620.3* 179.7621.2 190.3621.2 188.1623.3 194.9622.8 203.6624.8 214.5625.0
Total torque (N)
21
0 rads s 2846.96262.9* 2338.46275.7 2473.86274.0 2445.86303.4 2531.26297.6 2647.56321.2 2787.76325.2
Torque, rst 5 s (N)
21
0 rads s 942.1685.8* 775.2687.9 823.9685.5 823.6698.5 824.0686.1 869.6695.6 917.06110.0
Peak torque (Nm)
21 D
1.09 rads s 203.3621.9 177.3614.0 191.8612.6 194.8615.0 197.6618.9 197.3618.0 200.7618.1
Total work (J)
21 [
1.09 rads s 754.7674.5 653.1651.0 686.8639.2 723.0651.4 741.5658.3 734.0664.7 754.1666.5
Peak torque (Nm)
21
3.14 rads s 150.8614.3 135.068.2 139.969.8 142.8612.4 148.5613.7 145.0614.1 148.0614.4
Total work (J)
21
3.14 rads s 504.8653.6 470.7624.6 478.0625.4 494.9634.3 524.3638.6 500.9640.0 512.3647.7
a D
Values are means6S.E.; n59. *Indicates signicant difference (P,0.05) from days 1, 2, 3, and 5. Indicates signicant difference (P,0.05) from day
[
1. Indicates signicant difference (P,0.05) from days 1 and 2.
dant with the time necessary for the activation of the cle function included testing during isometric and iso-
inammatory response resulting from muscle damage. Our kinetic contractions. During both modes of contraction,
plasma IL-Ib data indicate that the inammatory response signicant decreases in force production followed muscle
of the immune system was almost immediately stimulated injury. Isometric functional capacity of the quadriceps was
by muscle insult. After adjusting for plasma volume shifts measured during a 15-s maximal effort. A signicant
[25], a nearly threefold increase in this cytokine was decrement in peak torque, and total torque during the 15-s
observed as early as 5 min following the series of contraction was detected 24 h following the damage-
concentric/ eccentric muscle actions. This acute response inducing protocol, and persisted through the next 4 days.
supports the role of IL-Ib as a pro-inammatory cytokine By day 7, neither peak nor total torque produced were
that is responsible for the release of other soluble agents of signicantly different from baseline values.
the immune system that more directly mediate the in- The 15-s isometric test was also used to evaluate muscle
ammatory response [26], and suggests that it may serve fatigue. Our results indicate that while the capacity of the
as a particularly sensitive marker of muscle damage. affected muscle to produce force was modulated by
Unlike previous studies, the present evaluation of mus- damage, the ability to maintain force was not. Based upon
Fig. 4. Average iEMG among three surface quadriceps muscles, and iEMG for rectus femoris. Values are means6S.E., n59. *Indicates signicant
difference (P,0.05) from day 0.
M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299 97
21
0 to day 1 occurred at 0 rads s , while the least severe
21
was at 3.14 rads s . Moreover, knee extensor function
(peak and total torque) remained signicantly compro-
mised for 5 days post-insult during maximal contractions
21 21
at 0 rads s , for 2 days (total work) at 1.09 rads s ,
21
while it was never signicantly affected at 3.14 rads s .
In fact, it may be that these data do not conict with
evidence that fast-twitch bers suffer more damage than
slow-twitch muscle. Rather, these results reasonably imply
that a relationship exists between the degree of functional
disturbance, and the capacity to generate force prior to
muscle insult. In essence, the greater the ability to produce
force, the greater the degree and duration of the loss
suffered following injury. Alternatively, it is possible that
since muscle damage was elicited with slow muscle
Fig. 5. Neuromuscular efciency (torque/ average iEMG). Values are actions, the effects are most pronounced during move-
means6S.E., n59. *Indicates signicant difference (P,0.05) from day 0.
ments of similar velocities.
In an effort to determine potential causes of the di-
these performance data, it appears that the bioenergetics of minished muscle function observed post-insult, iEMG
the involved muscle, at least with respect to the phos- analyses of the three surface muscles of the quadriceps, i.e.
phagen pathway, remained unaffected. rectus femoris, vastus lateralis, vastus medialis, were
Isokinetic muscle performance was tested at two differ- performed. Komi and Viitasalo [30] had previously ex-
21
ent velocities. As with isometric results, at 1.09 rads s amined the electrical activation of the rectus femoris and
isokinetic performance of the quadriceps, measured as vastus lateralis following unaccustomed exertion, but those
peak torque and total work during the ve repetition set, authors collected iEMG data during submaximal contrac-
was signicantly impaired as a result of muscle damage. tions, and for only 2 days following muscle challenge. In
However, in contrast to isometric function of the knee the present study, iEMG data were collected on each of the
extensors which was reduced through day 5 of the study, three surface muscles during maximal isometric contrac-
21
peak torque at 1.09 rads s demonstrated a signicant tions for 10 days following muscle insult. In analyzing the
decrement only at day 1, while total work was hindered at effects of exercise-induced damage on muscle activation,
days 1 and 2. we examined each of the three muscles individually, as
Interestingly, when knee extensor muscle performance well as the average iEMG value among the three muscles.
21
was assessed at 3.14 rads s , no signicant decreases in This indicator of overall activation of the quadriceps
peak torque, or total work were noted. Although both remained steady over the period of investigation, sug-
slow- and fast-twitch muscle bers are activated at this gesting that decreased muscle performance cannot be
speed, fast-twitch bers primarily, if not exclusively, ascribed to impaired electrical excitation of the contractile
account for force production [27]. Our in vivo functional apparatus. When iEMG data for each of the three muscles
results appear to be at odds with ultrastructural evidence were considered individually, no impact of injury was
that exercise-induced muscle damage occurs mainly in apparent in either the vastus lateralis or the vastus
fast-twitch bers [5,28]. Presumably, if damage was medialis. However, electrical activation of the rectus
specic to fast-twitch bers in this study, functional femoris showed increases during the study period. These
diminution would be most pronounced at the velocity of results are interesting in that iEMG alterations were
21
3.14 rads s , where these bers are almost exclusively localized to the same quadriceps muscle in which soreness
responsible for force production. Yet, signicant functional was most acute. This localized soreness cannot be attribu-
disturbances are restricted to the slower movement velocity ted to normal conditions of greater recruitment, since data
21
of 1.09 rads s where, during maximal exertion, both collected at day 0 indicate that the rectus femoris is
slow- and fast-twitch muscle bers contribute to force activated no more than the vasti muscles during maximal
production [29]. exertion prior to muscle insult (data not shown). Unlike the
Upon closer examination, our results reveal that the vasti muscles, the rectus femoris crosses both the hip and
amount of force generated determines both the degree, and knee joints [31]. Perhaps this anatomical arrangement
the duration of the functional impairment of muscle leaves the rectus femoris more susceptible to stretch-
damaged during unaccustomed exertion. Consistent with related damage from eccentric actions, and consequently to
the forcevelocity curve, our baseline data illustrate that greater discomfort and electrical activation. The data
21
peak torque of the quadriceps was greatest at 0 rads s presented here do not allow rm conclusions to be drawn
21
(219.0 Nm) and lowest at 3.14 rads s (150.8 Nm). on these points.
Interestingly, the most severe drop in peak torque from day During the initial 5 s of the maximal isometric muscle
98 M.R. Deschenes et al. / Journal of the Neurological Sciences 174 (2000) 9299
action, measurements on muscle force production and signicant alterations in electrical activation. Yet overall
electrical activation were concurrently performed. This iEMG analyses indicate that the reduction in force pro-
enabled us to examine neuromuscular efciency, or the duction was unrelated to muscle activation which remained
capacity of the contractile elements to respond to electrical unchanged for 10 days following muscle insult. These data
stimulation of the sarcolemma. Komi and Viitasalo [30] contributed to the most striking nding of our inves-
had found that 2 days following unaccustomed exertion, a tigation: neuromuscular efciency was signicantly im-
disturbance in this relationship existed such that, relative to paired the rst day following insult, and remained compro-
original efforts, a greater degree of electrical activation mised for 10 days. This prolonged disturbance is probably
was necessary to perform the same submaximal muscle related to dysfunction within the EC coupling mecha-
task. Our data extend those ndings. During maximal nism; further research is needed to determine at what point
effort, signicant decrements in neuromuscular efciency subsequent to unaccustomed exertion this disorder is
were detected throughout the 10-day post-insult interval. resolved.
Initially, this nding may be explained by the signicant
reduction in torque produced during the 5-s interval
documented on days 1, 2, 3, and 5, without concomitant Acknowledgements
changes in overall iEMG values. This implies that early
neuromuscular dysfunction is mainly due to damage to The authors would like to express their gratitude to the
contractile elements [5,6]. Yet, torque returned to normal subjects who generously gave their time and effort to
values by day 7, while neuromuscular alterations persisted enable the successful completion of this study. This project
through day 10. The continuation of this disturbance may was supported, in part, by a grant from the Faculty
be related to the gradual trend (P50.07) toward an Research Committee of The College of William & Mary.
increase in average iEMG activity, mainly accounted for
by the rectus femoris, during the study period. Still, an
impaired capacity of the myolaments to respond to References
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