Download Autonomic Nervous System Responses to

Autonomic
Nervous System Responses
Relation to Ventilatory
Threshold*
Yoshiharu
Yoshio
Yamanwto,
PhD; Richard
Nakamura,
L. Hughson,
Ph.D.;
our recent
approach
to study
control
of heart
rate during
means
rate
ofheart
reference
(Tvent).
The
nervous
rationale
with
nomic
(HRV)
for the
(cardiac)
system
gether
variability
study
was
that
parasympathetic
(PNS
the
and
SNS,
underlying
analysis
to ventilatory
markedly
reduced
HRV
respectively)
sympathoadrenal
and epinephrine
enhanced
above
rine
was
and
SNS
the
low-dimensional
less
complex
Tvent,
that
indicators
concentrations
Tvent,
and that
indicators
were
autonomic
considered
activity.
respect
with
with
dynamics
DF)
to-
These
appearance
that
might
findings
to implication
in
uring
incremental
exercise,
(VE) begins
oxygen
uptake
to
the
increase
(o)
or
minute
nonlinearly
noninvasive
using
onset
on
bicarbonate
abrupt
blood
Since
these
on the
specific
work
lactacidosis
buffering
early
rate
(WR)
acidosis,
evidence
has
stimulus.12
was
However,
both
such
is,
dioxide
an increase
not
always
in
accom-
compromise
methodologically
speaking,
parameters
it
of
relating
by what
question
to
is called
arises;
authorsl2
should
only
may
and
they
be
said
have
physio-
response
to
“V-slope”
method.’
exercise,
not
Then
all ofthe
reinterpret
most
performed
VE
these
were not
to this question
regarded
that
been
the
we
responded
that
can
AT
that AT determined
for the subjects
was
at
rose
by
pattern
of
counterevidences
Dempsey
were
et
provided.
develop
lactacidemia
during
The
clinically
examined
given
by the
by these
by VE responses
whose
ventilatory
could be
control
“appropriately.”
However,
consid-
the
responses
as such
appropriate
Tvent,3’4
that
responses
during
subse-
an
alteration
in
concomitant
could change
included
recently
these
factors
in heavy
forced
concept
the
2 more
was
the
because
original
to modify
despite
of
of
authors
the
Dempsey
could
not
marked
acidemia.
of the
anaerobic
significance
the
Department
of
Kinesiology,
University
Ontario,
Canada
(Drs.
Yamamoto
and
of Sports
Science,
Waseda
University,
Japan
(Dr. Nakamura).
of Tvent
of
shown
to
intensity.
catecholamines,”
of potassium.’5
recent
review
and
All of
of Cunningham
we introduce
our recent
system
control
during
rate
variability
spectral
(HRV;
respiratory
As the
controller
with
was
directly,
the
delineate
the detailed
for Tvent.
However,
implication
has
been
of the
discussed
last
they
ty.’7
not
for
results
for
on beat-
reference
to its
investigating
ofthese
physiologic
there
was
results
in the
intervals
special
method
and
approachesl321
to
exercise
by means
heartbeat
analysis
to Tvent.
HRV
Waterloo,
Hughson),
and
Tokorozawa,
PARAMETERS
TO
studies
mechanisms
at least
one
clinical
cardiology.
section.
EVALUATE
CONTROL
The
analysis.
2 or more
at frequency
of
are
coherent
with
nervous
that
might
system
Nervous
System
with time series
spectral
human
HRV spectra
have
harmonic
components.”
solely
system
usually
blood
AUTONOMIC
HEART
mediated
nervous
components
demonstrated
<0. 15 Hz
Autonomic
major
>0. 15 Hz,
parasympathetic
other
OF
study
of HRV has evolved
In the frequency
domain,
displayed
are
Waterloo,
Department
Saitama,
in the
were
to high
of
in
to
a lack
of hyperventilation
than
lactacidosis
at a moderate
temperature,
concentration
listed
basis)
could
not
responsible
This
a Tvent
inability
On the contrary,
trained
athletes
body
arterial
article,
nervous
of heart
the
direct
report
demonstrated
despite
their
exercise
the
al,
other
exercise
are
relationship
in
LA
onset
et al18 to modulate
feedback
control
of ventilation,
are thought
to be related
to the sympathoadrenal
to-beat
blood
change
both
These
more
the
for
is necessary.
Several
factors
affect
YE during
In this
autonomic
lactate
failed
exercise
the enzyme,
myophosphorylase.
et al’
reported
that
highly
(AT)
to
exercise
One
et al’#{176}
in which
they
with
McArdle’s
disease
evidence
blood
before
that
Hagberg
patients
This
leading
responsible
per se at Tvent
potential
Tvent.8
the
review
threshold
doubt
cast
strenuous
status could not induce
the
and that epinephrine
infusion
hyperventilate
of
ofTvent.
that
significantly
exercise,56
moderate
dysoxia,
occurrence
preliminary
to
activation
have
observations
of ventilatory
incremental
acid-base
Tvent,7
a
resultant
cellular
in the
mechanisms
respect
threshold
(Tvent).
to be due to the
studies
that
the
(LA)
the
many
resulted
induced
to alter
quent
reports,
included
concentration
lactacidosis
and the
system.’2
hypothesisl2
metabolic
206S
in carbon
with
Tvent
determinations
necessarily
ering
ventilation
with
exercise
intensity,
namely
the ventilatory
This al)rupt
increase
in VE was considered
*From
That
increase
commensurate
while
problems
mechanisms
been
cardiol-
an
that YE tracked
the
increase
in Vco2,
this
is circular
logic.
To answer
this question,
therefore,
a study on the physiologic
D
In
by this
validated
ogy.
and
always
presented
results
because
latter
method?
The answer
of
have
was
exchange
that
to lactacidosis
definite
the
reflect
for clinical
(Vco,)
due
to gas
Methodologically
(DF) of HRV
that PNS was
the
more
emphasized
logically.
auto-
of change
norepineph-
incline
panied
and SNS indicator)
these changes
in PNS
associated
(low
recently
raise
activity,
the rate
(plasma
to
they
H
has been
of cerebral
and
output
and sympathetic
complexity
below
activity
with
threshold
system
in terms
of fractal
dimension
series.
The experimental
results
showed
time
and
autonomic
exercise
by
spectral
to its relationship
to reflect
shown
in
Ph.D.
We introduce
nervous
system
special
to Exercise
seen
pressure
(PNS)
at
or
and
Response
PNS
to Exercise
Downloaded From: http://journal.publications.chestnet.org/pdfaccess.ashx?url=/data/journals/chest/21646/ on 05/07/2017
at
both
The
0. 1 Hz
It
is
levels
activity.””
below
variability.”
the latter
components
be associated
with
(SNS)
One
by changing
has
and
been
frequencies
sympathetic
activities.un
(Yamamoto,
Hug/non,
Nakamura)
There
are
taken
in the
components
to indicate
PNS
of a nonharmonic
and
HRV
SNS
component
band
spectrom.’
This
to be
“fractal”
in nature
signal
and
can
developed
graining
that
monic
permits
simultaneous
components
well
related
as computation
The
Yamamoto
the
to
and
fact
that
power
after
whereas,
the
was
used
the
original
and
shown
the
components,
nonharmonic
(fractal)
subtracted
Sxx’ form
(Ph) and
ratio
HRV
power
with
magnitude
retains
its
of
Eight
plus
tests
the harmonic
consideration
nonharmonic
has
been
is
13
component
plane
with
of this
repre-
component.
spectral
was
plotted
plot.
for the
effect
(j) vs log
frequency
slope
of the
From
the value
of 3,
for 1 <33.
For 3>3
1 and infinity
(oo), respectively.
value
of DF or 3 can have
the
taken
DF
was
and
The
in
spec-
meanings.
The
brain
oscillators
complex
and the
dynamics.”
reflection
ofthe
DF
number
dimensionality
correlation
consists
interactions
vs log
power
work
regression
was
is
thought
ofthese
of human
dimension
to
cerebral
(the
be
lower
encephalographic
be 8 to -0o in the
(EEG)
awake,
recording,
eye-opened,
high-dimensional
system
is also
physical
evaluated
of DF)
by
“information
epileptic
As to the
implication
the
producing
agent
activity
that
toward
the
correlation
had
of this
finding,
the
a stable
seizure
periodic
test
to
A
rich:’
stimuli
that
cen-
dimension
a low mean
they
tended
motion.
value
drive
In
of
of 2.05.
speculated
to
the
such
PNS
to decrease
subjects
intensity
dra-
exercised.
reached
was
it increased
autonomic
detail,”
more
by observing
that
male
on
an
60%
statistically
significantly
nervous
system
by adopting
catecholamines
volunteers
a ramp
and
performed
electrically
of a 5-mm
braked
warm-up
in a ramp
analyzed
for
DF
(3)
fashion
incremental
cycle
ergometer,
50 W, followed
exhaustion.
The
period
for
was
HRV
epinephrine
Overall
in Figures
et al,”
rate
was
Arterialized
from
an indwelling
heated
plasma
of change
so small
dorsum
in LA during
that
Tco,
of the
intensity.
Such
expected
for
1) pressures
states,
“isocapnic
a reduction
these
buffering”
CHEST
the
very
1). Thus,
is
I 101
showed
clear
at
in
PE’rCO2
an
ramps
anomaly
I 5 I MAY,
Downloaded From: http://journal.publications.chestnet.org/pdfaccess.ashx?url=/data/journals/chest/21646/ on 05/07/2017
1992
for
and
very
slow
ramp
linearly
in
the determination
(Tvent)
slow
hand
are shown
Wasserman
almost
method’s
was impossible
profile
of end-tidal
0,
hyperventilation
brain
venous
catheter
(NE),
increased
(Fig
The
particularly
norepinephrine
intensity
ofY-slope
the
Fig
of
of data.
periods
response
analysis.
5 mm
on the
LA,
to exercise
( PE’ICO,,
rate
slow ramp
responses
(E) concentrations.
responses
of the physiologic
variables
1 through
3. As was suggested
by
was
Nevertheless,
onset
of
the
relation
every
vein
analyses
10-mm
heart
spectral
sampled
in a superficial
the
of
by
rate
at
until
successive
nature
important
blood
in the first report
for the
dynamics
in the field of human
seizure
data,
of increase
in WR was 2.0 Wmin’.
This very
protocol
allowed
us to observe
quasi-steady-state
ofTvent
demonstrated
and
increment
Babloyantz
and
Destexhe,
existence
oflow
dimensional
during
these
for
6 healthy
Briefly,
exercise
EEG
and
exchange,
HRV
contrary,
while
studied
exercise
protocol
ters,”
it is not surprising
to see high DF (3 close
to unity)
in
the resting
human
HRV#{176}” with high level ofPNS
activity.’#{176}
physiology,
gas
the
the
the
on the
initial
WRs of 20
predetermined
indicator
when
Tvent,
for the
ofthe
From
until
indicator,
recently
steady-state
reported
state.’
and this makes
redundant
processing
of external
possible.
Considering
the fact that there are projections
could
transmit
the cerebral
activity
to the autonomic
to decrease
to
of HRV
of an electro-
has been
but quiet
called
very
For example,
activity,
limit
the
110%
PNS
exercise
position.
constant
alveolar
rest
to
to Tvent.
sitting
increased
continuously.
were calculated.
up to 100%
exercise
WR
physiologic
oscillators.”’
was
and
the
the
it reached
profile,
responses
relative
submaximal
in
WR
with
HRV
set
17-mm
until
showed
SNS
the
Tvent.
consisting
of (nonlinear)
VENTILATORY
intensity
100%,
compared
We have
as
DF
generate
of the
responses.
calculated
them
aging,3’
instability
observed
tests,
90%,
results
110%
responses
among
various
susceptibil-
challenges,’#{176}’
AND
six
ventilatory
The
plots
of a number
centers.
in
including
measurement
fashion
60%,
unchanged
at
we
completed
It continued
fractal
following
one
way, we have
SNS responses
autonomic
electrical
with
resting
while
matically
as
The
Tvent
the
031,
study,’
exercise
in a ramp
power
linear
and
submaximal
mm
total
HRV
by orthostatic
HRV were measured
and SNS indicators
characteristics,
Therefore,
in a log
as the
in the
reflex
albeit
observed
situations,
induced
one
W, or 30%,
in
of change
power-law
frequency
3
(P/Ph),
by
been
stress,’8
subjects
and
Tvent.
component
exponent.a
3 estimated
power
normalized
to have
in log
ft
frequency
was
also
RESPONSE
load
During
we
arrhythmia.
(fractal)
the
DF=2J(3-1)
the
sinus
reported
by
expressed
where
to high
of the
state
has
pathologic
In an earlier
constant
time
Thus,
attractor,
THRESHOLD
grained)
Sxx’
and
DF
AUTONOMIC
spectra
18
recovery
process
In an analogous
increase
ofdefensive
the
in
syncope
heart.”'”
harmonic
whereas,
indicator
of respiratory
The
trum
as
of low
PNS
respectively.182
to
and
to
on a chaotic
of a functional
physiologic
ity
remains
that
impossible
However,
challenges#{176} and exercise”
was associated
with
close
to 2.0 (13-- 1 .0), suggesting
that DF of HRV
reduction
on
From the harmonic
component,
the integrated
power
0.0 to 0. 15 Hz (P,) and 0. 15 to 1.0 Hz (Ph) was calculated.
PNS
and SNS activities
were
evaluated
by high frequency
power
indicative
The
based
spectra
represent
Sxx to obtain
of HRV
psychologic
cross-power
(coarse
to
DF
was
by
was
brain
shown
its rescaled
version,
does
not. A standard
and
rescaled
as
detailed
auto-power
(Sxx)
Sxx was
(fractal)
(Sxx’).
nonharmonic
been
component
with
to estimate
series
activities,’8
the method
Briefly,
har-
components.2#{176}
has
nonharmonic
time
SNS
nonharmonic
method
“
fractal
HRV
series
sented
this
of the
recently
be
diffIcult.
low
to orthostatic
analysis
would
extremely
the
of very
by
spectral
processing
be
activities,
sense is indicative
We have recently
and
it is cross-correlated
the harmonic
component
original
between
use
Hughson.
method
FFT
the
of DF from
rationale
broad-
suggested
evaluation
to PNS
information
would
be characterized
in some
series.”
coarse
those
consists
its
has been
the fractal
dimension
(DF) which
of the complexity
of a given time
(CGSA),
also
by
noise
, ‘‘#{176}
than
HRV
characterized
broad-band
a technique,
other
activities.
for all subjects.
and CO,
(PE’rO,)
evidence
moderate
at Tvent
because
of the
I Supplement
of the
exercise
is to
be
so-called
specific
15
2075
(mmol’
(L- mm)
80
L’)
6i
C
0
a)
60
0
C
C)
(L’ mm’)
2.0
40
>
(ngmL’)
a
2
-I
20
0
c%J
0
--1’
-
0
C
0
I..
a
0
-
0
1.0
1C
a
(Torr)
C,l
z0
(ng-mL1)
0.2
0
.40
w
-40
0
-20
delta
Work
FIGURE
1. VE,
VCo2,
Wmin’)
ramp
exercise.
20
Rate from
T
and PErCO,
responses
All variables
shown
-
40
a.
(W)
during
w
very
in Figures
slow
(2.0
1 through
0
3
delta
exercise.
protocol.
In Figures
been
shown
to the Tvent
as ensemble
determined
The statistical
above Tvent
comparison
showed
that
1 through
3, all variables
averages
for 6 subjects
by the PETO,
and PEiCO,
of regression
only PEi’CO,
systematically
at Tvent.
Figure
2 shows the responses
as intensity
and
Tvent.
and
E. Each
of
increase
increase
in the recent
study
of Mazzeo
and Marshall.”
the recent
study
of Savard
et al” suggested
that
creases
progressively
The
3.
study,”
low
gradually
values
over
it increased
the
slopes.
reached
2.04
higher
was
of
Tvent,
below
with
(lower
for
the
orthostatic
component
of
heart
SNS
indicator
greater
had
our
recent
C/)
z
in-
0
Tvent,
to
the
graded
the
fractal
less
21-
3
with
finding
dimensional,
w
for
at Tvent
that
a
change
A similar
responses
3r
C
C)
C
0
increased
value
of 1.92)
C)
a
Cl)
-40
delta
nervous
2085
summary,
system
responses
results
during
concerning
exercise
autonomic
in
0
-20
20
40
complex
dynamics.
In
0
40.
reaching
than
mean
indicating
low
I
80
earlier
(p<0.05)
thereafter.
stress,”
HRV
the
rate
ramp
i!1
(0I
z
summarized
At WR
to DF
DF)
slow
Cl)
of our
a significant
(corresponding
values
are
results
increased,
interestingly,
very
was
muscles
as WR
The
Tvent.
steeply
More
obtained
levels
at
more
the
during
intensity.
variables
with
decreased
WR
leg
exercise
of HRV-related
indicator
concentrations
exercising
greater
In agreement
PNS
very
the
with
responses
in Figure
in
from
responses
See text for abbreviations.
.
suggested
of catecholamines
E
(W)
A threshold-like
of the
However,
and
40
T
in
in
response
effiux
catecholamine
NE,
20
Rate from
0.1
1 changed
with an
greater
LA,
0
Work
profiles.
below
Figure
NE,
gradually
apparently
exceeded
blood
in
have
in relation
slopes
of LA,
LA, NE,
and E increased
exercise
intensity,
with
an
concentration
2.
FIGURE
text for abbreviations.
Wmin’
-20
-40
are ensemble
averages
for 6 subjects
in relation
to ventilatory
threshold
(Tvent)
determined
by the PETO,
and PEi’CO,
profiles,
and plotted
against
work
rate (WR)
relative
to WR at Tvent.
See
relation
Work
Rate
from
T
to
(W)
3. PNS indicator,
SNS indicator,
and the spectral
(13)responses during very slow ramp exercise.
FIGURE
Autonomic
Nervous
System
Response
to Exercise
Downloaded From: http://journal.publications.chestnet.org/pdfaccess.ashx?url=/data/journals/chest/21646/ on 05/07/2017
(Yamamoto,
exponent
Hughson, Nakamura)
Tvent
showed
Tvent,
that
indicators
in
that
the
PNS
rate
withdrew
was enhanced
PNS
and
appearance
less
Tvent,
indicators
the
and
were
below
autonomic
increases
anesthetized
activity
that
these
changes
associated
low-dimensional
complex
completely
in sympathoadrenal
above
SNS
of
reflect
almost
of changes
with
dynamics
the
the
that
might
ered
the
to
be
mechanisms
important
methodologically.
results
for
ventilatory
context,
Proper
of the
the
system
these
and
the
peripheral
beyond
the
could
have
by showing
that
cardiac
performance”
has been
via
demia
due
et
several
the
by their
mechanisms
impaired
originally
to decreased
oxygen
that
well-trained
could
as
matched
trained
runners
despite
the
maximal
cardiac
output
of the
patients.
show
oxidative
indicators
lactacidemia
might
Neither
such
if one
and
merely
Tvent,
reflect
affects
of the
reduced
skeletal
examined
the
lower
to
indicate
the
relationship
Tvent
physical
increase
anterior
susceptible
during
the
vagotomy
arrvthmia
using
kins,’
HRV
or
Electrical
to increase
atropine
formation.42
spectral
exercise
has
Recently,
analysis
had
the
of HRV
dogs
and
with
the
other
hand,
sympathetic
neural
could
threshold,
indicators
to
than
activity
also
and
provided
that
that
a
protec-
DF (3) of HRV
above
the
to increased
between
system
moderate
Tvent
exercise
risk
of cardiac
the change
in cardiac
status
and a possible
cardiac
exercise
remains
to be inves-
experimentally
Whipp
study
BJ,
respiratory
in future
was
Koyal
blood
gas
lactate,
and
J Appl Physiol
research.
supported
in part
by
SN,
Beaver
RL.
muscle
Methodologies
WL.
1984;
GC.
during
Anaerobic
exercise.
metabolites
in progressive
for measurement
Physiologist
Gass
exchange
1983; 54:1032-38
5 Davis
HA,
before
and
during
The
SW,
Ivy JL. Lactate
anaerobic
lactic
of the
anaerobic
27:304-11
acidosis.
threshold
as determined
J Appl Physiol
1981;
Eur
47: 141-49
6 Farrell
during
exercise.
acidosis
J Appl
Heigenhauser
GJF,
incremental
7 Kowalchuk
per
the
dogs
occlusion
approximating
complexity,
lead
and
a Holter
correlation
to 1 .0 (to 3.0)
of
their
J Appl
Physiol 1973; 35:236-43
2 Wasserman
K, Whipp
BJ, Koyal SN, Cleary
MG. Effect of
carotid
body resection
on ventilatory
and acid-base
control
during
exercise,
J AppI Physiol 1975; 39:354-58
3 Green
HJ, Hughson
RL, Orr GW, Ranney
DA. Anaerobic
exercise.
Hos-
dimension
generator
This
K,
and
8 Scheen
healed
to
of HRV
patient
wearing
They
stated
the
loss
as wellas
1 Wasserman
JM,
and
metabolic
shown
published
arrhythmogenesis
relationship
clinically
threshold.
resistant.
On
nervous
during
4 Hughson
of spontaischemia,’
been
PNS
autonomic
vulnerability
exercise.
stimulation
ventricular
reported
that
by a coronary
lower
as
innervattone
Biliman
in
nor
to
recently
Considering
Tvent
The
patients
cardiac
function.
the spectral
analysis
myocardial
infarctions,
to ventricular
fibrillation
submaximal
a
are
and
due
susceptibility
dimensionality
gradually
indicating
above
between
in cardiac
to
REFERENCES
intracellular
inactivity
ofthe
heart.
been
shown
heartbeat
(increased)
threshold
ratio,
the activity
of cardiac
autonomic
nerves
sinoatrial
node.”
The cardiac
parasympathetic
bilateral
as
efilux,
pyruvate
can
in the
and
muscle
in lactate
lactate
et al,”
their
electrical
stability
vagus nerves
has
age-
redox
potential
concentrations
changes
as
of lethal
observation
fibrillation
threshold
and decrease
the incidence
neous
ventricular
fibrillation
during
myocardial
while
graded
to
Vo2max
This
muscle
enzyme
disease
per se, not the impaired
It should
be emphasized
that
evaluated
ing the
during
capacity
of the
effiux
and
of muscle
Wasserman
Thus,
dysoxia.#{176}
responses
disease
LA response
during
exercise.”
by Connett
et aF’#{176}
has convincingly
review
availability.
by
endurance
of
in
suggested
LA
similar
capacity
shown
that lactate
interactive
functions
redox
similar
as
shown
In our results,
of SNS with
grants
from the Ministry
ofEducation,
Japan
(02951196),
from Meiji
Life Foundation
of Health
and Welfare,
Japan,
and from the Natural
Sciences
and Engineering
Research
Council
ofCanada.
The authors
are grateful
to Isao Muraoka,
Professor
of Waseda
University,
and
Minort,
Shinohara,
M.Sc. , at University
ofTokyo
for their
help in
conducting
part
of the experiments
in this study.
Coyle
heart
been
increase
to an integer
arrythmogenesis.
study,
has
The
extrasystole
PNS
almost
completely.
that changes
in cardiac
al”
onset
ACKNOWLEDGMENT:
lactaci-
Indeed,
ischemic
et
on the
in a cardiac
time ofhis death.
vulnerability.
tigated
performie,
transport.’”
exercise
substrate
lower
clas-
effect
might
Skinner
process
our
60%.
for
In
instability.
decreased
intensity
concept
cardiac
proposed,
patients
determinant
the recent
in
about
PNS activity.e
the enhancement
at Tvent
to the
from
this
by
observations
decreased
had the lower Tvent.
However,
this
mean
that the lower
Tvent
in these
caused
well
this
patients
with
the
and the higher
NYHA
reported
suggested
fractal
potential
that
by
observed
withdrawal
of
these data suggested
profile
dimension
also
without
electrical
1.0 prior
studies.
one
supported
the
al”
possible
Indeed,
of these
for
tion
investigators
offunction”
necessarily
ances
a delinescheme
at least
accompanied
was
arrhythmogenesis
monitor
at the
cardiology.
Later,
patients
scope
Tvent
Furthermore,
auto-
although
especially
preliminary
ofAT was originally
developed
by Wasserman
for evaluating
cardiac
performance
of cardiac
indirectly
sifications
does
not
between
was
It is of note
be evident
to fibrillation”
stimulation
decreased
threshold
of response
a cardiac
noninvasive
Within
this
vulnerability
ganglion
fibrillation
type
autonomic
also
accumulated
controller
was
patients.
maximal
link
Tvent,
results
for clinical
The concept
and McIlroy
of the
a possible
status
and
hyperventilation
implication
but
ventricular
same
the simultaneous
Taken together,
is consid-
physiologically,
interpretation
showed
central
In addition,
only
for Tvent
of the so-called
“AT” by
to exercise
is required.
results
nervous
ation
not
estimation
responses
our
nomic
responsible
ventricular
dogs,
stellate
threshold”
activity.
DIsCussIoN
Seeking
the
cardiorespiratory
J Appl
A, Lemaire
perfusion
triangulaire.
9 Dempsey
systems
Jones
NL.
responses
Arch
Int
Vidruk
in exercise:
d’adrenaline
Physiol
EH,
Mitchell
Fed
Proc
1983;
ofpH
on
progressive
d’hyperventilation
lors
Biochem
update.
Effect
during
Physiol 1984; 57:1558-63
P Abaissement
du seuil
intraveineuse
JA,
and the increase
of 1s/o,
Physiol 1987; 62:1551-55
d’un
exercise
91:187-96
GS.
Pulmonary
1985;
44:2260-70
control
10 Hagberg
JM, Coyle EF, Carroll JE, Miller JM, Martin WH,
Brooke
MH. Exercise
hyperventilation
in patients
with McArdle’s disease.
J Appl Physiol 1982; 52:991-94
11 Dempsey
JA, Hanson P. Henderson
K. Exercise-induced
arterial
hypoxemia
in healthy
humans
at sea-level,
J Physiol
1984;
355:161-74
CHEST
I
101
I
5
Downloaded From: http://journal.publications.chestnet.org/pdfaccess.ashx?url=/data/journals/chest/21646/ on 05/07/2017
I
MAY,
1992
I
Supplement
2095
12 Wasserman
the lactic
K, Beaver WL, Whipp BJ. Gas exchange
theory and
acidosis
(anaerobic)
threshold.
Circulation
1990;
challenges.
31
81:1114-30
13 Beaver
WL,
detecting
Wasserman
the
K,
anaerobic
Whipp
threshold
BJ. A new
by
method
for
exchange.
J AppI
gas
Physiol 1986; 60:2020-27
14 Whipp BJ. Ventilatory
control during exercise
in humans.
Ann
Rev Physiol 1983; 48:393-413
15 Paterson
DJ, Friedland
JS, Bascom DA, Clement
ID, Cunningham DA, Painter R, et al. Changes
in arterial K + and ventilation
during exercise
in normal
subjects
and subjects
with
McArdle’s
syndrome.
J Physiol 1990; 429:339-48
16 Cunningham
DJC, Robbins PA, Wolff CB. Integration
of respiratory response
to changes in alveolar
partial
pressures
of CO,
and 0, and in arterial
pH. In: Handbook
of physiology:
the
respiratory
system:
control
of breathing,
section
3, chap 15.
Bethesda,Md:
American
Physiological
Society, 1986; 2:475-528
17
Christensen
exercise.
18
NJ,
1991;
H.
Sympathetic
Ann Rev Physiol
Yamamoto
new
Ga1bO
Y, Hughson
method
nervous
activity
for studying
Coarse
heart
rate
spectral
variability.
23
Pagani
M,
Lombardi
F,
Guzzetto
5,
Rimoldi
0,
25
NY Acad
Mayer-Kress
Sci 1990; 591:402-09
G, Yates FE, Benton
M, T,rsch
W,
Poppl
SJ, et al. Dimensional
analysis of nonlinear
oscillations
heart, and muscle. Math Biosci 1988; 90:155-82
26 Mayer-Kress
G, Layne
SR Dimensionality
ofhuman
cephalogram.
Ann NY Acad Sci 1987; 504:62-87
in
brain,
27
Babloyantz
ofepilepsy.
28
Skinner
in the
1989;
A, Destexhe
L, Keidel
A. Low-dimensional
chaos
heart:
implications
for
clinical
cardiology
34
Wasserman
obic
35
of syncope.
Spectral
during
postural
Circulation
1990;
Strange
spillover
5, Kiens B, Christensen
NJ,
from skeletal
muscle during
of muscle
mass.
Am J Physiol
1989;
role
K, Mcllroy
metabolism
37
38
39
40
41
42
1964;
Weber
ICT, Kinasewitz
Detecting
the threshold
patients
during
of anaer-
exercise.
J
Am
GT,
Janicki
ventilation
JS,
during
Fishman
exercise
AP
in
Oxygen
patients
with
chronic cardiac failure. Circulation
1982; 65: 1213-23
Matsumura
N, Nishijima
H, Kojima
5, Hashimoto
F, Minami
M , Yasuda H . Determination
of anaerobic
threshold
for assessment of functional
state in patients
with chronic
heart failure.
Circulation
1983; 68:360-67
Lipkin DP, PerrinsJ,
Poole-Wilson
PA. Respiratory
gas exchange
in the assessment
ofpatients
with impaired
ventricular
function.
Br Heart J 1985; 54:321-28
Coyle EF, Martin WH, Ehsani AA, Hagberg
JM, Bboomfield
SA, Sinacore
DR. et al. Blood lactate threshold
in some welltrained
ischemic
heart disease
patients.
J Appl Physiol
1983;
54:18-23
Ivy JL, Withers
LIT, Van Handel
PJ, Elger DH, Costill
DL.
Muscle respiratory
capacity
and fiber type as determinants
of
the lactate threshold.
J AppI Physiol 1980; 48:523-27
Connet
RJ, Honig
CR, Gayeski
TEJ, Brooks
GA. Defining
hypoxia:
a systems
view of Vo,, glycolysis,
energetics,
and
intracellar
Po,. J Appl Physiol 1990; 68:833-42
Kolman BS , Verrier
RL, Lown B. The effect of the vagus nerve
stimulation
upon vulnerability
of canine ventricle:
role of the
sympathetic
parasympathetic
interactions.
Circulation
1976;
52:578-85
Hess
DS,
Hanlon
1982;
Billman
Levy
T, Scheinman
of ventricular
GE,
vagal
MN.
M,
tachycardia
Budge
R,
J. Termimessage.
Desal
by carotid
sinus
65:627-33
Hoskins
during
cardiac
44
MB.
cardiac
14:844-52
and
nation
43
in
Cardiol
utilization
tone
RS. Time
series
submaximal
exercise:
in animals
susceptible
1989;
analysis
of heart
evidence
for
rate
reduced
to ventricular
fibrilla-
80:146-57
Autonomic
interactions
in cardiac
45
46
Sci 1990; 601:209-21
Lown
B, Verrier
RL. Neural activity
N EngI J Med 1976; 294:1165-70
control.
Ann
NY
Acad
Biotechnology
1990; 8:1018-24
Saul JP, Albrecht
P. Berger
RD, Cohen RJ. Analysis oflong term
heart
rate variability:
methods,
1ff scaling
and implications.
Comp Cardiol 1988; 14:419-22
30 Butler GC, Yamamoto
Y, Xing HC, Northey
DR, Hughson
RL.
Heart rate variability
and fractal dimension
during orthostatic
EA,
in humans:
tion. Circulation
Chaos
AL.
and
P. Influence
of plasma catecholamines
on
during graded
exercise.
J Appl Physiol
Richter
variability
RE.
risk
before
257:H1812-18
29
2105
GK,
exercise
in an instance
Proc Natl Acad Sci USA 1986; 83:3513-17
JE, Goldberg
AL, Mayer-Kress
G, Ideker
and
B. Norepinephrine
Circulation
electroen-
variability
67:1319-22
Saltin
R,
Pizzinelli
R, et al. Power
spectral
analysis
of heart rate and
arterial
pressure
variabilities
as a marker
of sympatho-vagal
interaction
in man and conscious
dog. Circ Res 1986; 59:178-93
24 Goldberger
AL. Fractal electrodynamics
of the heartbeat.
Ann
to aging
Savard
Physiol
Furlan
tilt:
relations
rate
33
71:1143-50
19 Yamamoto
Y, Hughson
EL, Peterson
JC. Autonomic
control
of
heart
rate during
exercise
studied
by heart
rate variability
spectral analysis.
J Appl Physiol 1991; 71:1136-42
20 Yamamoto
Y, Nakamura
Y, Bufler GC, Hughson
RL. Fractal
dimension
of heart rate variability
and physiological
stress. Am
J Physiol, submitted
for publication
21 Nakamura
Y, Yamamoto
Y. Autonomic
control
of heart rate
during
physical
exercise
and fractal dimension
of heart
rate
variability.
J Appl Physiol, submitted
for publication
22 Saul JR Beat-to-beat
variations
of heart rate reflect modulation
ofcardiac
autonomic
outflow. NIPS 1990; 5:32-7
ofheart
Mazzeo RS, Marshall
the lactate
threshold
analysis:
J Appl
characteristics
32
36
graining
Am J Physiol, submitted
for publication
Mietus
J, Moody
GB, Goldberger
LA,
81:1803-10
during
1983; 45:139-45
RL.
Lipsitz
Furey
SA
autonomic
extrasystole
III,
Levy
activity,
MN.
and
threshold
The
interactions
arterial
in the
and ventricular
dog.
among
fibrillation.
heart
rate,
pressure
Am
upon the multiple
Heart
J 1983; 106:1112-
20
AUtOnOmiC
Nervous
System
Response
to Exercise
Downloaded From: http://journal.publications.chestnet.org/pdfaccess.ashx?url=/data/journals/chest/21646/ on 05/07/2017
(Yamamoto,
Hughson,
Nakamura)