Prog. Nwo-PsycJxophcmnacwL & BioL Psychic&. 1999, Vol. 23, pp.

447463
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FROIUTAL REACTMTY AND SENSATION SEEKING AN ERP STUDY

IN SKYDIVERS

ANNICK PIERSON, JACQUES LE HOUEZEC, ARNAUD FOSSAERT, STEPHANIE DUBAL

and ROLAND JOUVENT

CNRS UMR 7593, H6pital de la Salp&i&e,

Paris, France

(Final form, March 1999)

Abstract

Pierson Annick, Jacques Le Houezec, Amaud Fossaert, StP;phanie Dubal and Roland Jouvent: Frontal
Reactivity and Sensation Seeking: An ERP Study in Skydivers. Prog. Neuro-Psychopharmacol. & Biol.
Psychiat. 1999,2& pp. 447463.81999 ElsevierScienazInc.

1. In the line of Zuckerman’s studies on sensation seeking and optimal level of arousal, the authors
hypothesized that high sensation seeking might be used to compensate for anhedonia due to basal
arousal deficit. A population of interest was found with parachutists practicing skydiving,
generally described aa very high sensation seekers.
2. After clinical assessment of emotional and affective components, amplitudes of the frontal P3 of
the ERP were used as indices of arousal.
3. Skydivers presented more negative symptoms (anhedonia and blunted-affect) than controls. This
was observed in isolation from any depressive episode, which would suggest the presence of
emotional deficit as a trait. As expected, skydivers presented more sensation seeking than controls.
These two results taken together could indicate that sensation seeking is an adaptive reaction to
anhedonia.
4. ERP results showed that l?ontal P3 amplitudes were larger in skydivers than in controls, whereas
in a previous study we showed the opposite in depressed patients with a similar emotional deficit.
This could indicate that the frontal P3 amplitude does not reflect the emotional deficit per se. We
suggest that it rather reflects the capacity to use some behaviors which improve automatic
attentional processes in order to obtain arousing stimulation that could counterbalance the
emotional deficit. Depressions with emotional deficit might be due to the lack of such a capacity.

Kev_words: emotional deficit, ERP, frontal reactivity, negative symptoms, orienting reflex, personality,
sensation seeking.

Abbreviations: Abram+Taylor blunted-affect scale (AT), Cchelle d’humeur depressive - depressive

mood scale (EHD), 6chelle de ralentissement dkpressif - depressive retardation scale (ERD), event
related potentials (E&P), Hamilton rating scale for depression (I-IRSD), Montgomery and Asberg
depressive rating scale (MADRS), orienting reflex (OR), reaction time (RT), t&personality
questionnaire (TPQ).
448 A. Pierson et al.

Introduction

Sensation seeking was described by Zuckerman (1979) as a “trait defined by the need for varied,
novel and complex sensations and experiences and the willingness to take physical and social risks for

the sake of experience”. Zuckerman et al (1972) asserted that the optimal level of arousal is the

physiological basis for the sensation seeking trait. Such an approach is in keeping with a biological

perspective, being based upon Berlyne’s (1950) and Hebb’s (1955) concept of an optimal level of
arousal and upon Eysenck and Eysenck’s later theory incorporating this same notion. Eysenck’s
hypothesis was that certain personality traits (extraversion for the case in question) may be explained by
chronic under-arousal which makes these individuals seek sources of stimulation so as to increase their

arousal to an hedonic point (Eysenck and Eysenck 1978).

Following along the same lines, our group conducted several studies investigating the relationships
between sensation-seeking and emotional deficiency, i.e. blunted-affect and anhedonia. From this
research, the authors concluded that high sensation seeking can be considered as intended to provide
sensations and to compensate for anhedonia due to a basal arousal deficit (Carton et al 1992a). It was
also shown that frontal P3 amplitudes were abnormally low in depressed patients with psychomotor
retardation and blunted-affect, and a correlation was observed between frontal amplitude and negative
symptoms (Pierson et al 1991; Partiot et al 1993).

The present study was designed to investigate indices of frontal dysfunction in subjects presenting a
sensation seeking trait. The task selected was designed to analyze cortical indices of arousal related to
certain cognitive processes by manipulating two experimental variables:
1) the probability of occurrence of the stimulus, which mainly required automatic orienting

processes,
2) the classification of the stimulus as being a target or a non-target for a motor response, that mainly
required selective attentional (and therefore controlled) processes.
The task consisted in differentiating two types of stimulus presented in different proportions, with

either frequent or rare stimuli being the target for the motor response (oddball task).

Working on the assumption that sensation seeking is an adaptive behavior intended to compensate for

basal anhedonia or blunted-affect, a clinical emotional and affective evaluation of the subjects was
made. A population of interest for such a study appeared to be people practicing high-risk sports,
generally described as very high sensation seekers. Several studies have reported relationships between
sensation seeking and the practice of sport (Hymbaugh and Garrett 1974; Zuckerman 1979), with a
higher relationship observed in men (Straub 1982). The present study examined parachutists of both
sexes practicing skydiving as a sport.
 Frontal reactivity and sensation seeking 449

Inclusion Criteria

Skydivers were recruited from a French center for sportive parachuting; they had been jumping and
skydiving regularly for at least two years. Controls were recruited by advertising in public buildings and
were matched for age, sex and social-cultural background. All subjects were volunteers and were paid

(4OUS$) for taking part in the study. To be included they had to give a written consent. Each group
consisted of twenty-two subjects (16M, 6F).

Exclusion Criteria

Interviews were conducted by a psychiatrist to ensure that no subjects were suffering from
psychiatric or neurological conditions or any visual problems that could interfere with the task.

Psvchooatholonical Evaluations

To evaluate depressive and anxious symptomatology as well as emotional disturbances, a psychiatrist

conducted a semi-structured interview with each subject. The following scales were used to rate the
subjects: Montgomery and Asberg Depressive Rating Scale (MADRS, Montgomery and Asberg 1979),
Hamilton Rating Scale for Depression, 17 item version (HRSD, Hamilton 1967), Covi Brief Anxiety
Scale (Lipman et al 1981; Covi and Lipman 1984) Tyrer Anxiety Scale (Tyrer et al 1984; French
version validated by Lemperiere et al 1989) and Widlbcher Depressive Retardation Scale (ERD -
Echelle de Ralentissement Depressif, Widlocher 1983). Emotional disturbances were assessed with the
Abrams-Taylor Scale for Blunted AITect (AT, Abrams and Taylor 1978; French version by Ammar et al
1983) and the Depressive Mood Scale (EHD - Echelle d’Humeur Depressive, Jouvent et al 1988; Carton
et al 1992b; Bungener et al 1996). The EHD scale, consisting of 20 items, has a coherent factorial

structure based on five factors: anhedonia, hyper-emotionality, emotional hyper-or hypo-expressiveness,

sadness, hyper-attentiveness. Two factors focus on emotional deficits: anhedonia and emotional hypo-
expressiveness and the conjugate use of these two clinical constructs provides a more accurate overall
assessment of blunted-affect/emotional deficit. Two other clinical constructs (hyper-emotionality and
emotional hyper-expressiveness) assess lack of control.

Subjects also completed the following questionnaires: Chapman’s Physical Anhedonia Scale (PAS,

Chapman et al 1976, Chapman and Chapman 1978; French version validated by Loas 1993),
Zuckerman’s Sensation Seeking Scale (SSS-form V, including four dimensions: Thrill and Adventure
Seeking (TAS), Experience Seeking (ES), Disinhibition (Dis) and Boredom Susceptibility (BS)
(Zuckerman et al 1964; French version validated by Carton et aI 1992b), and the Cloninger

Tridimensional-Personality-Questionnaire (TPQ, Cloninger 1987; French version validated by Lepine

450 A. Pierson et aI.
1994) indexing three independent dimensions, the dimension of interest in this study being Novelty
Seeking (NS)

ERP Procedure

After clinical assessment, the subject was placed in a dimly-lit, sound-attenuated chamber for ERP

recording. The subject was seated comfortably, 150 cm from the computer screen where visual stimuli

were presented for the oddball task. The subject was asked to stare at a fixation point (0.5 cm x 0.5 cm
white square) and to press a response button as fast as possible after the target stimulus (3cm x 3cm
yellow square), but to ignore the non-target stimulus (red square).

Two task conditions were used with varying probability of the target occurrence (20% target/llO%
non-target or 80% target/20% non-target). Under each experimental condition, the subject was presented
with stimuli until 30 responses had been obtained for the rare stimulus. The Inter-Stimulus-Interval (ISI)
varied at random between 1 set and 1.5 sec. The order of presentation of the two task conditions was
counterbalanced across subjects in each group. Test sessions lasted 30 minutes. Training trials were
conducted before commencing the ERP recording. Subjects were asked to relax and to blink as little as
possible during the test.

EEG was recorded from 9 electrodes (Fpz, Fz, Cz, Pz, Oz, T3, T4, C3, C4) according to the
international lo-20 system (Jasper 1958) with respect to a linked-earlobe reference. The EEG electrodes
were embedded in a cap (Electrocap, Dallas, USA). The earth electrode was attached to the right cheek.
Two electrodes for bipolar recording of vertical ocular movements were placed above and under the left

eye in order to reject trials contaminated by electro-oculogram (EOG 2 f 50 uV). Electrode impedance

was controlled and maintained as being less than 2 kilo-ohms. Signals were low-pass filtered at 25 Hz
and high-pass filtered at 0.16 Hz by analogue filters in the amplifiers. Data were digitized on-line at a
sampling rate of 250 Hz using the InstEP system (Ottawa, Canada). The sampling period was 1024 msec
(200 msec pre-stimulus to 824 msec post-stimulus).

Assessment

After elimination of trials with excessive eye movements, ERP were averaged for each subject
according to task condition (target or non-target) and response (correct, omission, false alarm). Because
of the small number of errors observed in the task, only ERP corresponding to correct responses were
analyzed; corresponding Reaction Times (RT) were obtained from the average procedure. The ERP
recordings were then low-pass filtered (off-line procedure of the InstEP system) using a digital filter set
at 12 Hz (-3 dB/octave). Computer-assisted scoring of ERP components was realized with InstEP

system using time-windows determined from grand-average (overall subjects) ERP. Peak amplitudes
were measured for the N2 and P3 components. N2 was measured at Fz between 200 msec and 350 msec
 Frontal reactivity and sensation seeking 451

P3 was measured at Fz and at Pz between 280 msec and 500 msec. Peak amplitudes were calculated

with respect to the 200 msec pre-stimulus baseline. The frontal N2-P3 complex was assessed by

measuring peak-to-peak amplitudes of these components as well as their separate values The P3b was
measured at the parietal site (Pz) The FzTz P3 amplitude ratio and RT were also analyzed. All scorings
were performed automatically by computer. In case of aberrant measures, individual recordings were

examined independently by two judges to reach consensus

Statistical Analvsis

Comparisons between demographic and clinical parameters for each group were tested using the
Mann and Whitney U-test. Analysis of Variance (ANOVA) was used to compare RT and amplitudes of
the ERP components in the two groups. Significance levels were adjusted using the Greenhouse-Geisser
correction when appropriate. Student t-test was then used for 2 x 2 planned comparisons with
Satterthwaite’s approximation for the degrees of freedom when variances were unequal. Correlations (r

of Bravais-Pearson) were calculated between the different measures of interest (anhedonia, blunted-
affect, sensation seeking, novelty seeking). The level of significance was set at p < 05

Results

One control subject was excluded because of noisy EEG recordings The high risk-taking group
therefore consisted of 22 skydivers and the control group of 21 subjects. Pairing of the two groups

according to demographic parameters (age, sex, social-professional background) was satisfactory and no
significant statistical differences were found. The mean age was 24.8 for both groups (Table 1)

Table 1
Demographic Parameters in the Two Groups of Subjects

Skydivers Controls
16 Men 15 Men Mann-Whitney
6 Women 6 Women
(n=22) (n=21)
m SD m SD U P
Age 24.77 3.85 24.76 3.31 220.5 0.80
Cultural level* 5.63 1.18 6.00 1.10 190.0 0.31
* the cultural level was scored from 1 = illiterate to 6 = long higher education

Clinical Results

The medical history, specifically psychiatric disorders and addictions, were similar in the two groups.
A history of minor depressive episodes was mentioned by 4 skydivers and 3 controls. Drugs had been
used, but rarely (less than three times) and only by a minority of subjects in each group (hashish: 8
452 A. Pierson et al.

skydivers and 7 controls; heroin: 1 skydiver); 7 skydivers and 4 controls were smokers. No subjects
suffered from any psychiatric disorder at the time of the experiment. This was confirmed by the low
scores observed on the depression and anxiety scales in both groups. No difference between the two
groups was observed for these scores. The data used for clinical and psychological assessments and the
results of the statistical comparisons of the two groups are presented in Table 2.

For Sensation Seeking, skydivers showed significantly more Thrill and Adventure Seeking and more
Boredom Susceptibility than controls. There was a tendency for Experience Seeking to be greater in
skydivers than in controls (p = ,064). Novelty Seeking as measured on the TPQ also tended to be greater
in skydivers, but again the difference did not reach significance (p =.077; Table 2).

Table 2
Scores for Clinical and Personality Evaluations in the Two
Groups of Subjects

Skydivers Controls Mann-

n=22 n=2 1 Whitney
m SD m SD U p+
MADRS (Montgomery and Asberg 1979) 3.95 3.84 2.18 2.44 174.5 .I62
HRSD (Hamilton 1967) 3.05 2.68 1.73 I .64 171.5 .I41
Covi (Covi and Lipman 1984) 1.00 1.02 0.86 I .08 214.5 ,670
Tyrer (Tyrer et al 1984) 4.63 5.04 2.32 2.03 180.0 ,210
Widlijcher ERD (WidlCicher 1983) 0.59 0.73 0.68 1.67 187.0 .I97
Abrams-Taylor (Abrams and Taylor 1.91 2.39 0.41 1.18 128.0 .004 **
1978)
EHD F1:anhedonia (Jouvent et al 1988) 1.41 1.99 0.50 1.37 159.5 ,041 *
EHD F2:hyperemotivity 3.68 2.98 3.45 2.36 223.5 .854
EHD F3:emotional expressiveness -0.86 0.99 0.59 2.58 221.0 ,793
EHD F4:sadness 0.77 1.34 0.45 0.86 221.5 .770
EHD F5:anxious hyperattentivity 0.86 0.99 0.77 0.87 221.0 ,793
EHD:blunted-affect (Fl + F3 hypo) 3.50 3.78 1.14 2.92 132.5 .008 **
EHD:lack of control (F2 + F3 hyper) 4.91 3.90 4.68 3.29 223.5 .854
Chapman PAS (Chapman and Chapman 14.95 5.41 13.77 6.87 208.0 .575
1978)
Zuckerman:Thrill and Adventure 9.23 0.61 7.36 2.42 137.5 .018 *
(Zuckerman et al 1964)
Zuckerman:Experience Seeking 6.64 2.08 5.36 2.24 155.5 .064
Zuckerman:Disinhibition 4.91 2.05 4.59 2.87 200.0 .455
Zuckerman:Boredom Susceptibility 5.18 1.71 3.82 2.15 143.0 ,030 *
TPQ Novelty Seeking (Cloninger 1987) 19.77 3.90 17.36 6.18 158.5 ,077
+ * p1.05 ** p1.01

Another important observation concerns emotional blunting in skydivers. Compared to the controls,
the skydivers had significantly higher level of both hetero- and self-rated emotional deficits. Higher
 Frontal reactivity and sensation seeking 453
scores were observed on both the Abram+Taylor blunted-affect scale and the Depressive Mood Scale
(EHD - anhedonia and hypo-expressiveness). A positive correlation was also found for these self-rated
(AT) and hetero-evaluated (EHD) deficits: Abrams-Taylor/EHD anhedonia [r(22) = .63, p = ,001 in
skydivers, r(21) = 57, p < .Ol in controls], and Abrams-Taylor/EHD emotional hypo-expressiveness
[r(22) = .75, p < .OOOlin skydivers, r(21) = .62, p < .Ol in controls].

No correlation was found between these indices of emotional deficits and sensation or novelty
seeking within each group. However, another dimension of the EHD (lack of control) showed the two
groups of subjects to be different, both for the personality trait (experience seeking in skydivers, novelty
seeking in controls) and the direction of the correlation. A negative correlation was found in skydivers
between experience seeking and both hyper-emotionality [r(21) = -.53, p < .Ol] and lack of control
[r(21) = -.55, p < .Ol], whereas a positive correlation was found in controls between novelty seeking and
both hyper- emotionality [r(22) = .61, p < ,011 and lack of control [r(22) = .51, p < ,021.

ERP and Behavioral Results

ANOVA conducted for each parameter used two inter-group factors (Group: skydivers vs controls
and Sex: males vs females) and two intra-group factors with two levels each (Target: targets vs non-
targets and Proportion: 20/80% vs 80/20%). No main effect of sex was found; the results are therefore
presented for males and females together.

No group effect was observed on RT (348 f 52 ms in skydivers vs 342 f 63 ms in controls)

Grand-average ERP traces are shown in Fig 1. ERP amplitudes are given in Table 3 ERP amplitudes
revealed two main group effects. The first was that the frontal N2P3 peak-to-peak amplitude was greater
for skydivers compared to controls in all experimental conditions (no interactions) (F(1,41) = 5.85, p <
.02). This result seems to be mainly attributable to frontal P3 amplitudes, which were significantly
greater in skydivers than in controls (p = -05) rather than to frontal N2 amplitudes which did not show
any significant inter-group difference.

A further difference between the two groups was observed for the frontal/parietal P3 ratio (Fig 2)
which was, on average, significantly higher in skydivers than in controls (F(1,41) = 3.96, p < .05).
Again this result seems to be due to differences in frontal P3 amplitudes (Fz), since the parietal P3
component (Pz) did not show any significant difference in amplitude between the two groups. A Group
x Target interaction did, however, indicate that the difference between the two groups was significant
for non-targets only (t(84) = 2.47, p < .02). In skydivers, the frontal/par&al P3 ratio was in fact
significantly greater for non-targets (0.76) than for targets (0.43) (t(76) = 3.77, p < ,001).
454 A. Pierson et al.

-00
x
 Table 3
ERP Amplitudes in the Two Groups of Subjects for the Different Experimental Conditions

Infrequent Frequent Infrequent Frequent

Targets Targets Non-Targets Non-Targets
Skydivers Controls Skydivers Controls Skydivers Controls Skydivers Controls
7
m SD m SD m SD m SD m SD m SD m SD m SD
Amp* N2b -1.12 2.31 -0.46 3.55 0.22 2.18 0.58 2.71 -2.83 3.37 -2.08 4.22 0.67 1.30 0.30 1.48 i
Amp P3a 7.14 4.71 6.75 4.24 4.67 3.64 3.66 2.67 11.71 5.70 10.58 4.78 3.10 2.37 1.55 1.44 2
Amp N2bP3a 8.26 5.06 7.21 3.80 4.45 3.39 3.08 2.60 14.54 5.49 12.66 5.71 3.03 2.73 1.25 2.02 R
Amp P3b 16.93 5.59 16.63 4.30 10.34 3.80 8.14 3.67 16.13 5.95 16.87 5.76 5.60 3.11 4.49 2.10 4g

Amp P3a/P3b 0.45 0.31 0.39 0.18 0.41 0.33 0.47 0.37 0.76 0.31 0.65 0.30 0.75 0.59 0.40 0.41 !z
Lat* P3b 363.4 30.2 362.7 34.9 332.7 29.0 322.5 42.4 377.5 49.4 352.6 40.6 345.8 44.6 364.4 75.3
* Amp. = Amplitude Lat. = Latency B
I
!z

i
456 A. Pierson et al.

FzfPz

1 .oo
skydivers
controls @J
0.80

0.60

targets nontargets

Fig 2 The ordinate gives the ratio of P3 amplitudes measured at Fz and Pz sites respectively (Fflz).
This ratio was significantly larger in skydivers compared to controls only for non-targets. This was
essentially due to a larger frontal (Fz) P3 in skydivers.

The other effects of the experimental conditions were similar in both groups. The classic effect of the
probability of occurrence of the stimulus on P3 amplitudes was found, with a greater parietal P3 for
infrequent compared to frequent stimulations, irrespective of target/non-target classification. Moreover,
this typical effect was also observed for indices related to automatic processes, frontal N2 and P3

amplitudes and N2P3 peak-to-peak amplitude. A target/non-target effect on these same ERP parameters

was also apparent in both groups of subjects, but depended on the probability of stimulation, with
amplitudes for targets greater than for non-targets, but only when stimuli were frequent (80%).

An Emotional Deficit in Skvdivers

The main finding of the present study was that skydivers presented a larger emotional deficit
(anhedonia and blunted-affect) than controls. These negative symptoms were observed independently of
any depressive episode. This point supports the argument that the presence of an emotional deficit
constitutes a trait.
 Frontal reactivity and sensation seeking 457

As was expected, sensation seeking (Thrill and Adventure Seeking, Boredom Susceptibility) was
greater in skydivers than in controls. The existence of negative symptoms in these subjects indicates that
sensation seeking may be an adaptive reaction to emotional deficits. It could be assumed that with
repetition, the sensation seeking value of jumps may be diminished. Indeed, it has been established that
anxiety or fear are reported by both experienced and inexperienced subjects (according to a different
time pattern), which would suggest that a jump always has a certain character of novelty (Schedlowski
and Tewes 1992; Bockheler 1995). A number of psychological studies have pointed out how unique this
threatening situation is (Galor and Tenenbaum 1986).

Frontal P3 Amnlitude as an Index of Larger Orienting Reactions in Skvdivers Comnared to Controls

From the cognitive point of view, the two groups of subjects did not show any significant difference
in the effects of the two intra group variables on ERP (targets vs non-targets and frequent vs rare
stimuli). This indicates, in particular, that controlled processes which are necessary to classify stimuli as
being targets or non-targets were similar in skydivers and in controls. In contrast, there was an effect on
ERP of the inter-group variable (skydivers vs controls): the frontal P3 component was larger in
skydivers than in controls whatever could be the stimulus but this difference between the two groups
was only significant for non-targets. As the non-target stimuli required more automatic processing,
results are in favor of an increase of automatic processes in skydivers, compared to controls. Moreover,
as frontal P3 amplitude (P3a) is known to index, with the preceding negative component (N2b),
orienting response (Renault 1983, Niiatanen and Gaillard 1983, Wijers et al 1989) it could be suggested
that skydivers have developed an ability to react automatically to all stimuli, including non-relevant
stimuli.

Why an Orienting Resnonse to Non-Relevant Stimuli in Skydivers?

Sokolov’s theory of orienting (1963) postulates that orienting reflexes occur when recognizing
novelty but also when recognizing any discriminable change in the stimulus The basic finding of an
anteriorly distributed positivity (P3a), with a relatively shorter latency than the classical pa&al P3
(P3b) in response to rare non-targets has been observed in a number of studies (Graham and Hackley,
1991). Sokolov (1960) proposed that the development of a neuronal model for repetitive stimuli should
block inputs in the reticular formation. This accounts for habituation provoked by repetition and
explains that the most frequent stimuli in an oddball task do not provoke as large orienting responses
than rare stimuli. Before complete habituation, residual indices of orienting might however be observed
for frequent stimuli and it is plausible that larger indices of orienting (here, NZb-P3a amplitude) in
skydivers by comparison to controls may be related to a more important residual orienting response not
only to rare but also to frequent non-targets (because, in comparison with controls, OR should be larger
and/or because of a less rapid habituation).
458 A. Pierson et al.

What is the Orienting in an Oddball Task?

It is common to consider after Lindsley (1970) that the orienting reflex results from a sudden increase
in reticular or thalamic non-specific activation. However Naatanen (1986) proposed that the orienting

response should have two components: an arousal component, resulting from a transient increase in non-

specific activation but also an attentional component consisting in a reflex-like attention switch to the

eliciting stimulus. For his part, Sanders (1983) in his cognitive-energetical model of arousal, stress and
performance related linear stages of information processing to energetical resources and proposed that
perceptive stages be dependent on arousal (phasic activation) in contrast to motor stages which he
proposed be dependent on tonic activation. This model is based upon the Pribram and McGuiness
(1975) neurophysiological scheme describing three different systems in the control of attention and the
major neuroanatomical structures involved in these processes. In the same perspective, Tucker and
Williamson (1984) consider the phasic arousal system as regulating the phasic response of the brain to
environmental events and as being the primary mechanism of the orienting reflex. This mechanism

should be dependent on norepinephrine pathways.

The position of Pribram and MC Guiness (1975) is that the phasic arousal system is integral to

emotional processes (the tonic activation system being important to motivational processes). Emotions
are considered as the processes by which the brain evaluates the subjective significance of
environmental events. In the same view, Tucker and Williamson (1984) suggest that the emotional
characteristic of the phasic arousal system and of its norepinephrine substrate is depression-elation. That

is coherent with catecholaminergic models of affective disorders that several authors relate to the only
form of depression with emotional deficit, in contrast with agitated and impulsive depression that could
be more dependent on serotoninergic pathways (Jouvent et al 1991, Hansenne et al 1995, 1998).

An Impairment in Frontal Structures of the Brain?

Although ERPs recorded at the frontal sites do not indicate that they originate from frontal sources,
the question of such frontal generators can be raised since some neuropsychological data describe a
syndrome caused by, or involving, deficiencies of the dorsomedial frontal cortex with a pseudo-
depressive state and lack of interest in the outside world (George et al 1994).

Several authors (George et al. 1994) consider that frontal regions may in some way regulate or
govern the primitive unmodulated behaviors (drives or natural tendencies) which are ingrained in sub-
cortical structures and the limbic system.

Other studies, principally those by Knight (1984), have observed a reduction in the amplitude of the

P3 wave in reaction to novel stimuli in frontally lesioned patients and interpreted this as the expression
 Frontal reactivity and sensation seeking 459

of a deficit in prefrontal control of the external/sensory-intemablimbic system. Knight also

demonstrated that in normal subjects the frontal P3 was greater in amplitude, and that the frontahparietal
P3 ratio was greater than 1, for novel stimuli. In the present study, while the fiontal/parietal P3 ratio is
not greater than 1, the higher ratio for non-targets in skydivers compared to controls (0.76 vs 0.36)
seems to suggest that skydivers show an abnormally strong orienting reaction towards non-targets. This
could be due to an involvement of frontal structures in processing not only of novel or rare relevant
stimuli but also of irrelevant stimuli in these individuals.

The Development of Orienting as a Mean of Adaptation

The increase of the P3a component amplitude in skydivers in comparison with controls is in
opposition to what we found in depressed patients with blunted-atTect (Pierson et al 1991, Partiot et al
1993). The authors have shown that blunted-affect in depression was primarily related to abnormally
low P3a amplitudes that they have interpreted as an impairment of orienting automatic processes, This
deficit extended to cover controlled processes, but only for high effort-demanding tasks (Pierson et al,
1996).
On the basis of Pribram and MC Guiness’s and Tucker and Williamson’s approaches, the larger OR
observed in skydivers than in controls for non-target stimuli seem to indicate that skydivers give an
emotional significance not only to relevant but also to irrelevant stimuli. The abnormal reactivity to non-
targets of skydivers could be interpreted as an adaptive reinforcement procedure which consists in using
the context as arousing stimuli. It may be suggested that skydiving is an adaptive form of behavior that
contributes to the development of orienting reactions.

Indeed, when individuals display a primary emotional deficit, two different attitudes are possible:
either they let the emotional deficit take hold and (with time or because of specific events) reach a stage
of clinical depression, or they react by developing certain adaptive forms of behavior aimed at
compensating for the deficit. High-risk taking activity is one of these possible forms of adaptive
behavior. This interpretation of skydiving as an active adaptation procedure is reinforced by the
existence of correlations showing that while skydivers register the highest scores for experience seeking,
they record the lowest scores for hyper-emotionality and lack of control. Their experience seeking is not
an impulsive behavior resulting from lack of control. This last result is coherent with an improvement in
the orienting attentional component.

Conclusion: The Frontal P3 Amplitude as an Index of Adaptive Capacities

Examining two populations with emotional deficits, we observed opposite results concerning some
indices of orienting (P3a amplitude). The former population (depressed patients), examined in a
previous work (Pierson et al 1991, Partiot et al 1993), showed a deficit in these indices whereas the
460 A. Pierson et al.

other population (skydivers), examined in the present study, showed, on the contrary, a larger
development of these indices than control subjects. This opposition suggests that, in individuals with

emotional deficits, the frontal P3 amplitude measured in reaction time tasks does not reflect the

emotional deficit per se. We suggest that it rather reflects the capacity to use some behaviors which
improve automatic attentional processes in order to obtain arousing stimulation that could

counterbalance the emotional deficit. Depressions with emotional deficit might be due to the lack of
such a capacity.

Acknowledgement

The authors thank B. Renault and R Ragot for their valuable comments in revision qf manuscript

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