Oscillatory Patterns in Sympathetic Neural Discharge and

Cardiovascular Variables During Orthostatic Stimulus

Raffaello Furlan, MD; Alberto Porta, MS, PhD; Fernando Costa, MD; Jens Tank, MD;
Lemont Baker, MS; Richard Schiavi, MS; David Robertson, MD;
Alberto Malliani, MD; Rogelio Mosqueda-Garcia, MD, PhD

Background—We tested the hypothesis that a common oscillatory pattern might characterize the rhythmic discharge of
muscle sympathetic nerve activity (MSNA) and the spontaneous variability of heart rate and systolic arterial pressure
(SAP) during a physiological increase of sympathetic activity induced by the head-up tilt maneuver.
Methods and Results—Ten healthy subjects underwent continuous recordings of ECG, intra-arterial pressure, respiratory
activity, central venous pressure, and MSNA, both in the recumbent position and during 75° head-up tilt. Venous
samplings for catecholamine assessment were obtained at rest and during the fifth minute of tilt. Spectrum and
cross-spectrum analyses of R-R interval, SAP, and MSNA variabilities and of respiratory activity provided the low (LF,
0.1 Hz) and high frequency (HF, 0.27 Hz) rhythmic components of each signal and assessed their linear relationships.
Compared with the recumbent position, tilt reduced central venous pressure, but blood pressure was unchanged. Heart
rate, MSNA, and plasma epinephrine and norepinephrine levels increased, suggesting a marked enhancement of overall
sympathetic activity. During tilt, LFMSNA increased compared with the level in the supine position; this mirrored similar
changes observed in the LF components of R-R interval and SAP variabilities. The increase of LFMSNA was proportional
to the amount of the sympathetic discharge. The coupling between LF components of MSNA and R-R interval and SAP
variabilities was enhanced during tilt compared with rest.
Conclusions—During the sympathetic activation induced by tilt, a similar oscillatory pattern based on an increased LF
rhythmicity characterized the spontaneous variability of neural sympathetic discharge, R-R interval, and arterial
pressure. (Circulation. 2000;101:886-892.)
Key Words: nervous system, autonomic 䡲 catecholamines 䡲 spectrum analysis 䡲 tilt-table test

D uring the last decade, a lively discussion has addressed

the physiological interpretation of the rhythmic fluctu-
ations that characterize the variability of cardiovascular
signals such as heart period or systolic arterial pressure
(SAP). Spectral methodology has been essential in assessing
the amplitude and frequency of these oscillations, which in
numerous and various experimental approaches, seemed to
reflect some patterns of autonomic regulation.1
One possible view holds that low and high frequency
(LFR-R and HFR-R) spectral components of heart rate variabil-
ity, despite their mixed origin,2 when measured in normalized
units or as LF/HF ratio (ie, in their reciprocal relationship),
furnish quantitative markers of, respectively, cardiac sympa-
thetic and vagal modulation.2–5 However, when LFR-R is
measured in absolute units, it does not furnish an index
of sympathetic modulation.6,7 Instead, the LF component
(LFSAP) of SAP variability is a widely accepted marker of
sympathetic vasomotor control.2,8
Similar rhythmic LF and HF periodicities characterize the
discharge activity of medullary and sympathetic pregangli-
onic neurons of anesthetized animals9,10 and postganglionic
sympathetic fibers of humans.11–13 Recently, direct recordings
of muscle sympathetic nerve activity (MSNA) furnished
evidence that the LF and HF spectral components of neural
discharge variability undergo changes correlated to those
undergone by the spectral components of the variability of
cardiovascular signals during graded changes in arterial
pressure obtained with the infusion of vasoactive drugs.12
However, it is not clear whether a more physiological
stimulus that would induce an increase in sympathetic activ-
ity (such as upright tilt) would result in similar oscillatory
patterns in sympathetic neural discharge and in the sponta-
neous rhythmicity of the R-R interval and SAP. Furthermore,
no data are available on the modifications of the rhythmic
components of MSNA or other indices of sympathetic func-

Received June 8, 1999; revision received September 7, 1999; accepted September 23, 1999.
From Centro Ricerche Cardiovascolari, CNR, Medicina Interna II, Ospedale L. Sacco, and Centro LITA di Vialba, Università degli Studi di Milano,
Milan, Italy (R.F., A.P., A.M.); the Division of Clinical Pharmacology, Department of Medicine (F.C., D.R.), and the Department of Biomedical
Engeneering (L.B., R.S.), Vanderbilt University, Nashville, Tenn; Clinic Bavaria Kreischa, Kreischa, Germany (J.T.); and the Division of Clinical
Pharmacology, Dupont Pharmaceuticals Co, Wilmington, Del (R.M.-G.).
Correspondence to Dr Raffaello Furlan, Unità Sincopi e Disturbi della Postura, Medicina Interna II, Ospedale L. Sacco, Università di Milano, Via G.B.
Grassi 74, 20157 Milano, Italy. E-mail raffaellof@fisiopat.sacco.unimi.it
© 2000 American Heart Association, Inc.
Circulation is available at http://www.circulationaha.org

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 Furlan et al Neural Sympathetic Oscillations During Tilt 887

tion, such as plasma catecholamine level, during orthostatic TABLE 1. Hemodynamic and Respiratory Parameters, MSNA
stress. These aspects are of paramount importance to under- Measurements, and Catecholamine Plasma Levels at Rest and
stand better the possible abnormalities in the central integra- During Upright Tilt
tion of autonomic control of the cardiovascular system that Rest Tilt
characterize orthostatic intolerance syndromes such as neu- HR, bpm 57⫾2 84⫾4*
rally mediated syncope 14 –16 or chronic orthostatic R-R interval, ms 1054⫾32 727⫾31*
intolerance.17
SAP, mm Hg 131⫾3 122⫾3
In the present experiments, recordings of MSNA were
DAP, mm Hg 73⫾3 72⫾4
maintained in control supine conditions and during passive
CVP, mm Hg 4.4⫾1.1 ⫺4.7⫾1.1*
head-up tilt in healthy volunteers. It was thus possible to
correlate the rhythmic fluctuations occurring in sympathetic RESP, cycles/min 16⫾4 15⫾2
outflow with similar oscillations detected from cardiovascu- MSNA, bursts/min 16⫾4 35⫾4*
lar variables during the most natural stimulus leading to MSNA, bursts/100 beats 29⫾7 44⫾6*
sympathetic excitation and vagal withdrawal, that is, the NE, pg/mL 227⫾26 430⫾32*
orthostatic position. E, pg/mL 47.3⫾13.9 117⫾18*
Data are mean⫾SEM. HR indicates heart rate; DAP, diastolic arterial
Methods pressure; RESP, respiratory frequency; NE, norepinephrine; and E, epinephrine.
Ten healthy, normotensive volunteers (4 women and 6 men aged Tilt data refer to the period from the fifth to the tenth minute of passive
27⫾1 years) with no evidence of organic disease (on the basis of the orthostatism.
interview and physical examination) were admitted to the study. *P⬍0.05 vs rest.
In every subject, we recorded the ECG with an AC amplifier and
direct arterial pressure with an intra-arterial catheter placed into the
radial artery of the nondominant arm. The catheter was connected to
Data Analysis
Microneurography recordings were considered to reflect MSNA in
a pressure transducer positioned at the midchest level to counterbal-
accordance with criteria previously described.20 CVP values are
ance hydrostatic modifications during the tilt procedure. Central
referenced to atmospheric pressure; no atrial transmural pressure
venous pressure (CVP) was recorded with a miniature (diameter, 3F)
values are provided because intrathoracic esophageal pressure was
catheter tip pressure transducer (Millar) introduced into a vein of the not evaluated concomitantly.
antecubital fossa and advanced near the right atrium. An intravenous
line was positioned in the opposite arm for blood sampling. Place-
ment of intra-arterial, CVP, and venous catheters was done under
aseptic conditions and local anesthesia. Respiratory activity was
evaluated by means of a thoracic bellows positioned at the midchest
level and connected to a pressure transducer.
MSNA was obtained by the microneurography technique, as
reported in detail elsewhere.18 Briefly, a unipolar tungsten electrode
was placed on the right peroneal nerve near the fibular head for
multiunit postganglionic sympathetic nerve recording. A reference
electrode was inserted subcutaneously, close to the recording needle.
The raw neural signal was amplified (1000-fold amplification), fed
to a band pass filter (bandwidth between 700 and 2000 Hz), rectified,
and integrated (time constant, 0.1 s) by a nerve traffic analyzer
system (Bioengineering Dept, University of Iowa). Neural sympa-
thetic activity, ECG, arterial blood pressure, respiratory activity, and
CVP signals were recorded on the paper of an optic chart recorder,
concomitantly digitized at 300 samples/s by an analogical to digital
board (AT-MIO 16E2, National Instrument), and stored on the hard
disk of a personal computer.
High-performance liquid chromatography with electrochemical
detection19 was used to measure plasma epinephrine and norepineph-
rine levels on venous blood samples.

Protocol
After overnight fasting, subjects were placed on an electrically
driven tilt table provided with a footrest. They then underwent the
instrumentation procedure as described above. Thirty minutes after
instrumentation, continuous signal recordings were initiated for
baseline resting condition (15 minutes), and a blood sample was
withdrawn for plasma catecholamine evaluation. Thereafter, the tilt
table angle was advanced at 15° intervals every 3 minutes until the
75° head-up position was reached. That position was maintained for
30 minutes. A second blood sample was obtained after 5 minutes of
75° tilt.
The experimental protocol was approved by the Vanderbilt Uni-
versity Institutional Review Board in Human Research, and written
informed consent was provided by all subjects.
Figure 1. Spontaneous fluctuations involving postganglionic
neural sympathetic activity (MSNA) and hemodynamic parame-
ters in a healthy human at rest and during 75° head-up tilt. Note
during tilt that neural discharge activity is grouped after a LF
pattern with a period of 10 s, which is coupled with analogous
rhythmic fluctuations in blood pressure (BP; ie, Mayer waves)
and heart rate (HR; measured in beats/min). A different oscilla-
tion at HF (period of ⬇3 s), synchronous with modifications of
respiratory activity (RESP), characterized CVP signal major fluc-
tuation. EKG indicates electrocardiogram.

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888 Circulation February 29, 2000
Analog data were analyzed offline after analog-to-digital conver-
sion at 300 samples per second per channel. The principles of the
software for data acquisition and spectrum and cross-spectrum
analysis of R-R interval and SAP variability and respiratory activity
have been described in detail elsewhere.3,21 The methodology we
used for signal processing and autoregressive and bivariate spectrum
analysis of MSNA variability were previously used to assess the
rhythmic content of thoracic efferent preganglionic sympathetic
activity in anesthetized cats.9 Briefly, the integrated sympathetic
nerve signal was low-pass filtered by a 2400 coefficients finite
impulse response filter with a cut frequency at 0.5 Hz. Thereafter, the
neural signal was sampled once per cardiac cycle synchronously with
the R-wave peak of the ECG from which the neurogram was
obtained. This neurogram, in turn, underwent autoregressive spec-
trum and cross-spectrum analyses.
The power of the LF and HF oscillatory components of R-R
interval variability is provided in both absolute (ms2) and normalized
units, in accordance with the suggestions of the Task Force of the
European Society of Cardiology and the North American Society of
Pacing and Electrophysiology.1 A similar approach was used to
express results of MSNA variability spectrum analysis. Absolute
values of each component were computed as the integral of the
oscillatory components LF (mean, 0.1⫾0.01 Hz; frequency bands
were between 0.04 and 0.12 Hz) and HF (mean, 0.27⫾0.02 Hz;
frequency bands were between 0.16 and 0.38 Hz). Normalization
was obtained by dividing the absolute power of each component by
total variance (minus the power of the very-low-frequency compo-
nent) and then multiplying by 100.3 The use of normalized units for
the LF and HF components of R-R interval and MSNA variability
was necessary because of the marked changes in these variances
when comparing different subjects and experimental conditions. The
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Figure 2. Examples of spectrum and cross-
spectrum analyses of sympathetic nerve dis-
charge rhythmicity and respiratory activity. Note
2 oscillatory components in spectrum of MSNA
variability. A coherence (K2) value ⬎0.5 is
obtained in HF region of cross-spectrum, sug-
gesting breathing periodicity plays an important
role in modulating sympathetic neural traffic.
PSD indicates power spectrum density; RESP,
respiratory activity; and a.u., arbitrary units.
LFR-R/HFR-R ratio, which is nondimensional, may furnish a frame of
reference for studies using different methodologies, and it represents
a further global index to explore the sympathovagal modulation of
sinoatrial node spontaneous activity.3
The squared coherence function (K2) was computed as the square
of the cross-spectrum normalized by the product of the spectra of the
2 signals.21 K2 quantifies the amount of linear link between oscilla-
tions with the same frequency contained in 2 different signals.
Values of coherence higher than 0.5 were considered significant.
Cross-spectrum analyses between respiratory activity and R-R
interval, SAP, and MSNA variabilities were also used to identify the
respiratory-linked oscillatory component of those signals, thus ex-
cluding a possible breathing entrainment of the LF oscillations.
Data are expressed as mean⫾SEM. Student’s t test for paired
observations was used to evaluate the changes produced by the tilt
maneuver. Differences were considered significant at P⬍0.05.
Results
Table 1 shows the effects produced by the tilt maneuver on
hemodynamics and on the biochemical and neurophysiolog-
ical estimates of sympathetic activity. During tilt, no major
changes could be observed in systemic blood pressure and
respiratory rate, although R-R interval and CVP values were
reduced. MSNA and plasma norepinephrine and epinephrine
levels increased, as expected.
Figure 1 illustrates the spontaneous rhythmic oscillations
of sympathetic neural discharge and the hemodynamic sig-
nals recorded in a subject at rest and during upright tilt. The
 Furlan et al Neural Sympathetic Oscillations During Tilt 889

TABLE 2. Spectral Measurements of MSNA, R-R Interval, and during tilt, K2 decreased in the HF band compared with values
SAP Variability at Rest and During Upright Tilt in the recumbent position. Finally, peak coherence values in
Rest Tilt the HF band between respiratory activity and MSNA were
unaffected by orthostatic stimulus.
MSNA
LF, nu 27⫾4 47⫾3*
Discussion
HF, nu 67⫾4 50⫾3*
In the present study, we addressed the hypothesis that a
LF/HF 0.5⫾0.1 1.0⫾0.2* common oscillatory pattern might characterize the variability
R-R interval of neural sympathetic discharge, heart rate, and blood pres-
Variance, ms2 7305⫾1263 3942⫾1029* sure in healthy humans during the sympathetic activation that
LF, ms2 1827⫾377 1762⫾845 attends orthostatic stress. Our findings suggest that an in-
LF, nu 43⫾5 82⫾4* crease in the LF component of all these variability signals is
HF, ms 2
2316⫾609 250⫾86* the hallmark characterizing the enhanced sympathetic mod-
HF, nu 48⫾5 15⫾4*
ulation that occurs during upright tilt. They also indicate that
the physiological increase of sympathetic activity produced
LF/HF 1.0⫾0.2 8.7⫾1.6*
by tilt enhanced the coupling between the LF component of
SAP
sympathetic discharge and the LF components of R-R inter-
Variance, mm Hg2 9⫾1 34⫾7* val and SAP variability, leaving unchanged the relationship
LF, mm Hg2 2⫾0.5 24⫾5* between respiratory activity and MSNA in the HF
HF, mm Hg2 2⫾0.4 6⫾0.9* component.
Data are mean⫾SEM. nu indicates normalized units.
*P⬍0.05 vs rest. Effects of Tilt on LF Component of
MSNA Variability
neural activity was clustered in series of bursts that tended to The results of the present study indicate that, at rest, the
occur with a periodicity of 10 s, which mirrored spontaneous pattern of sympathetic neural discharge was characterized by
LF fluctuations of blood pressure (Mayer waves). 2 major oscillatory components at ⬇0.1 and 0.27 Hz, which
A second spontaneous oscillation at the respiratory fre- were synchronous with analogous fluctuations in R-R interval
quency characterizes CVP variability. The crucial role played and SAP spontaneous variability. This observation agrees
by respiratory activity in modulating sympathetic neural with previous findings obtained from both animal and clinical
traffic is depicted in Figure 2. A large oscillatory component studies suggesting that the sympathetic preganglionic9 and
at the same frequency of breathing is evident in the spectrum postganglionic11–13 discharge activity also display LF and HF
of MSNA while the subject is recumbent. The linear relation- oscillatory patterns.
ship between the spontaneous periodicity of respiration and In our group of healthy subjects, upright tilt was charac-
MSNA is presented in the coherence diagram (bottom) terized by the expected reduction of CVP and a marked
showing a K2 value that is ⬎0.5 only in the HF respiratory increase in heart rate, MSNA, and plasma epinephrine and
band. norepinephrine levels, which is suggestive of a remarkable
Results of the frequency domain analysis of spontaneous enhancement in overall sympathetic activity. However, we
R-R interval, SAP, and MSNA variability are summarized in did not assess the contribution of norepinephrine spillover
Table 2. During tilt, the normalized power of the LF oscilla- and clearance during tilt22 in sustaining such high plasma
tory component of MSNA variability (LFMSNA) increased, levels of norepinephrine.
whereas HFMSNA decreased, thus paralleling similar changes Similar to what was observed during hypoglycemia,23 our
in the R-R interval and systolic blood pressure spectral findings indicate that a generalized increase in neural sym-
profiles (Figure 3). In addition, the amount of increase in pathetic traffic develops as a consequence of a gravitational
sympathetic neural discharge in response to tilt was corre- stimulus. In addition, they also highlight the flexibility of the
lated to the increase of the normalized power of LFMSNA sympathetic response, which in other peculiar conditions,
(Figure 4). such as during mental stress24 and hyperinsulinemia25 in
The linear relationships between R-R interval, SAP, and humans or hemorrhage26 in animals, may selectively activate
MSNA spontaneous variability, as assessed in the LF and HF only some compartments.
bands by the peak coherence values, are summarized in In the present study, the tilt maneuver was attended by a
Figure 5. At rest in the LF region, 7 of 10 subjects showed K2 clear modification in the pattern of sympathetic neural dis-
values ⬎0.5 between MSNA and R-R interval variability charge; this consisted of an increased number of bursts
(mean, 0.58⫾0.06), whereas the K2 between MSNA and SAP clustered with a periodicity of ⬇10 s, which were detected by
variability was ⬎0.5 in 9 of the 10 subjects (mean, power spectral analysis as a marked increase of LFMSNA.
0.64⫾0.03). During the tilt maneuver, K2 between MSNA Therefore, during tilt, LFMSNA became predominant in the
and R-R interval and SAP variability increased (P⬍0.05) in autospectrum, mimicking analogous changes in R-R interval
the LF band in all subjects (0.81⫾0.04 and 0.81⫾0.05, and SAP variability. Importantly, the increase in the sympa-
respectively), suggesting enhanced coupling between the LF thetic neural discharge induced by tilt was accompanied by a
spontaneous fluctuations of each pair of signals. Conversely, proportional enhancement of the relative power of LFMSNA,

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890 Circulation February 29, 2000
indicating a link between this oscillatory component and the
amount of sympathetic neural firing.
These observations agree with the results of previous
studies in humans in which baroreflex unloading induced by
nitroprusside administration was associated with an increase
in MSNA and in the LF component of its variability.11,12
A 0.1 Hz discharge rhythmicity linked with LF spontane-
ous oscillations of R-R interval9,10 and blood pressure27–29 has
been recorded in anesthetized animals from sympathetic
neurons belonging to different areas of the nervous system
involved in cardiovascular regulation. Results compatible
with ours were obtained in several studies in which stimuli
eliciting an enhancement of sympathetic efferent activity,
such as inferior cava vein,9 common carotid bilateral occlu-
sions,29 or an increase of cerebrospinal fluid pressure,28 were
characterized by the onset of a typical LF oscillatory pattern
in sympathetic neural discharge, coupled with pronounced LF
fluctuations of systemic arterial pressure. Interestingly, those
oscillations persisted after baroreflex afferent inflow was
abolished (by a stabilizer device connected to the arterial
system of the animal29 or in dogs with spinal section after
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Figure 3. Effects produced by tilt on power
spectra of MSNA, R-R interval (RR), SAP vari-
ability, and respiratory activity (RESP). At rest,
2 oscillatory components characterize spectra
of MSNA, R-R interval, and SAP variability.
During sympathetic activation induced by tilt,
LF component of MSNA increases, thus resem-
bling changes obtained in same oscillatory
component of R-R interval and SAP variability.
Although HFMSNA amplitude is unaffected by
passive orthostatism, its relative power is
reduced during tilt. Differences in y-axis scales
between rest and tilt in power spectrum density
(PSD) of R-R interval variability reflect marked
reduction of total R-R interval variance induced
by orthostatic stress. a.u. indicates arbitrary
units.
bilateral vagal denervation28). These observations suggest
that baroreflex activity is not necessary to the genesis of LF
fluctuations of both preganglionic sympathetic efferent dis-
charge and blood pressure and that these fluctuations may be
the result of the activity of a central oscillator. In keeping
with this, the results of a recent study by Cooley et al30
challenged the necessary role of arterial baroreceptors in
generating LF oscillations of R-R interval variability. Indeed,
in 2 patients with intractable heart failure, the abolition of
spontaneous fluctuations of systemic blood pressure by a left
ventricular assist device was characterized by persistent LF
components in the power spectrum of R-R interval
variability.30
However, in humans, the importance of baroreflex inhibi-
tory activity on tonic MSNA is well established. In addition,
theoretical31 and experimental32,33 models have hypothesized
a role of baroreceptor mechanisms in the genesis of the LF
oscillatory components of both R-R interval and SAP vari-
ability.33 Thus, a limitation of the present study is that we
could not differentiate the relative contribution exerted by an
intrinsic LF rhythmicity inherent in any sympathetic excita-
 Furlan et al
Figure 4. Relationship between modifications in MSNA pro-
duced by tilt maneuver and concomitant increase in power of
LF oscillations of sympathetic discharge activity. Note that
increase in sympathetic neural discharge is associated with
concomitant increase of power of its LF rhythmicity. n.u. indi-
cates normalized units.
tion2 from the likely concomitant reflex mechanisms occur-
ring during baroreflex unloading.
Effects of Tilt on HF Component of
MSNA Variability
The results of the present study also indicated that the
normalized power of HFMSNA decreased during tilt. This
pattern suggests that recordings of sympathetic activity might
also offer a window to the likely link between HFMSNA
Figure 5. Relationship between R-R interval (RR), SAP, and
MSNA rhythmic fluctuations, as assessed by coherence function
(K2) in LF and HF regions, both at rest (white bars) and during
tilt (dashed bars). Tilt increases coupling between signals in LF
band, suggesting a preferential pattern of spontaneous fluctua-
tions of signals during sympathetic activation on basis of fre-
quency at 0.1 Hz. HF oscillatory component shows opposite
pattern. Tilt does not affect relationship between respiration and
MSNA variability. *P⬍0.05 vs tilt.
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Neural Sympathetic Oscillations During Tilt 891
rhythmicity and parasympathetic central regulation. In keep-
ing with this hypothesis is the previous finding of an increase
in HFMSNA during the baroreflex-mediated enhancement of
parasympathetic activity obtained by increasing systemic
blood pressure with phenylephrine administration.12 In addi-
tion, a reduction of neural sympathetic discharge and a
concomitant enhancement of the normalized power of HFMSNA
were evident after low-dose atropine infusion; this directly
increases central parasympathetic activity by activating cen-
tral muscarinic receptors.13 However, the finding in the
present study of a high peak coherence between MSNA and
the respiratory signal, which was unaffected by modifications
in the autonomic profile induced by tilt, suggests a strong link
between respiration and sympathetic efferent discharge,
which might be mediated by vagal afferents.34
Regarding this aspect, our results agree with previous
observations emphasizing the role of respiratory activity in
generating rhythmic oscillations in heart rate and MSNA.35
Clinical Implications
The definition of a pattern of MSNA variability during the
gravitational stimulus in healthy subjects may have important
clinical implications by furnishing a frame of reference in
those conditions characterized by orthostatic intolerance. In
fact, manipulation of sympathetic activity by centrally acting
drugs such as clonidine and yohimbine worsened or, con-
versely, ameliorated orthostatic tolerance in patients with
neurally mediated syncope.16 In a group of subjects affected
by chronic orthostatic intolerance syndrome, the exaggerated
tachycardia in the absence of changes in mean blood pressure
during standing have been attributed to a central abnormality
in the distribution of sympathetic activity to the vessels and to
the heart.17 It is, therefore, conceivable that in these and
related diseases,36 abnormalities in the sympathetic drive to
different cardiovascular target organs may coexist with con-
comitant impairments in sympathetic discharge rhythmicity.
Conclusions
The results of the present study indicate that the overall
sympathetic excitation induced by upright tilt is characterized
by an increase of the discharge activity of the sympathetic
peroneal fibers and by a proportional modification of their LF
rhythmic pattern of firing. These changes were associated
with concomitant relative reductions of HFMSNA and were
mirrored by analogous modifications in R-R interval and SAP
variability. Thus, a common oscillatory pattern can be de-
tected from the variability of the sympathetic outflow and the
cardiovascular target functions. These findings are consistent
with the hypothesis of a central-pattern push-pull organiza-
tion, according to which sympathetic excitation and vagal
withdrawal are linked to an enhancement of LF oscillation
and, conversely, vagal excitation and sympathetic inhibition
are linked to an increase in the HF component.2
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 Oscillatory Patterns in Sympathetic Neural Discharge and Cardiovascular Variables
During Orthostatic Stimulus
Raffaello Furlan, Alberto Porta, Fernando Costa, Jens Tank, Lemont Baker, Richard Schiavi,
David Robertson, Alberto Malliani and Rogelio Mosqueda-Garcia

Circulation. 2000;101:886-892
doi: 10.1161/01.CIR.101.8.886
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