Infant Behavior & Development 23 (2000) 271–284

Optic flow sensitivity in neonates

François Jouena,*, Jean-Claude Lepecqb, Olivier Gapennec,
Bennett I. Bertenthald
a
Laboratoire de Psychobiologie du Développement, 41 rue Gay Lussac, EPHE/CNRS, F-75005
Paris, France
b
Neurosciences Cognitives et Imagerie Cérébrale, LENA, UPR CNRS 640, F-75651 Paris, France
c
COSTEC, Université Technologique de Compiègne F-60000 Compiègne, France
d
Department of Psychology, University of Chicago, Chicago, IL 60637, USA

Received 16 January 2001; received in revised form 15 February 2001; accepted 15 March 2001

Abstract
The present research investigates neonatal sensitivity to optic flow. Twenty five 3-day-old infants
were placed inside a dark room and observed while presented with a 10 s bilateral and backward
peripheral optic motion. Seven constant flow velocity conditions were used (2.5, 5.0, 10.0, 15.0, 20.0,
25.0 and 30.0 degrees per second) and were compared to a baseline motionless condition. Sagittal
deviation of the head was computer-sampled at 60 Hz frequency with pressure transducers. As
indicated by the mean head pressure during optic flow exposure, infants reacted with backward
leaning of the head whose magnitude was linearly related to the optic flow velocity. Additionally,
isolated head postural responses were identified. The magnitude of these responses was clearly related
to the optic flow velocity. © 2000 Elsevier Science Inc. All rights reserved.

Keywords: Neonate; Optic flow; Vestibular system

1. Introduction

When exposed to a linear backward optic flow, stationary standing adults evidence
backward postural reactions and report a sensation of forward motion. Such a sensation falls
under the category of visually-induced illusory body motion, or vection, and in this particular
case is referred to as a forward linear vection. Similarly, when exposed to a forward flow,

* Corresponding author. Tel.: ⫹33-1-44-10-78-83; fax: ⫹33-1-43-26-88-16.

E-mail address: fjouen@free.fr (F. Jouen).

0163-6383/00/$ – see front matter © 2000 Elsevier Science Inc. All rights reserved.
PII: S 0 1 6 3 - 6 3 8 3 ( 0 1 ) 0 0 0 4 4 - 3
272 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

adults exhibit forward postural reactions and report a feeling of backward motion (backward
linear vection). Generally, adults display postural reactions in the very direction of the flow
and report a feeling of translation in the opposite direction (see Andersen, 1986 for a review).
In infancy research, vection has not been investigated although a huge amount of work has
been devoted to the postural reactions shown by infants when exposed to optical flow. Such
reactions are now theoretically framed as a part of perception-action couplings in early
ontogeny (Bertenthal, 1996).
Lee and Aronson (1974) were the first to show that infants’ posture may be affected by
visual information. Babies who had recently learned to stand were tested on a motionless
floor, within a moveable room. The infants faced the interior end wall and the whole
structure, except the floor, was moved so that the end wall approached or receded. Subjects
compensated for a nonexistent loss of balance signaled by the optic flow pattern and fell in
the direction of the optic flow. If the end wall moved away from the baby, the infant fell
forward and if the wall moved toward the baby, the infant fell over backward. Peripheral
vision is particularly important for maintaining postural stability. Pope (1984) showed that
in 3-month-old infants, movement in the center of the visual field did not result in significant
postural adjustment, whereas a slight movement in the periphery is sufficient to induce a
complete loss of stability. These results have been extended by Bertenthal and co-workers in
infants aged between 5 and 9 months (Bertenthal & Bai, 1989, Bertenthal, 1990 for a
review). Using an enclosure that permitted independent movement of the front and side
walls, these authors showed that 7- and 9-month-olds compensated in a directionally
appropriate reaction to whole-room movement. Nine-month-olds, but not 7-month-olds
responded with systematic compensation to side wall movement suggesting a developmental
trend between 7 and 9 months in the ability to use peripheral optic flow for postural control.
Using a similar type of moving-room apparatus, Higgins, Campos and Kermoian (1996) also
demonstrated that 8.5-month-old infants without locomotor experience showed minimal
compensation to peripheral optic flow. In contrast, 8.5-month-old babies with crawling or
walker experience showed significantly higher degrees of postural compensation to periph-
eral flow (see also, Campos et al., 2000).
Although these results suggest increased sensitivity to peripheral optic flow following the
onset of locomotion, they do not imply that infants are insensitive to peripheral optic flow
prior the acquisition of locomotor skill. Butterworth and Hicks (1977) found that infants too
young to walk but nevertheless capable of unaided sitting compensate for visually specified
motion when seated in a moving room. This finding has been extended by Pope (1984) and
Jouen (1986) to younger infants. They investigated the effects of optical flow exposure on the
control of head posture in 2-month-old infants. They found that even infants unable to sit
without support perform directionally appropriate compensatory movements of the head
when stimulated with optic flow. Testing even younger infants, Jouen (1988) and Jouen and
Lepecq (1989) provided evidence for directionally appropriate head movements under rough
optical flow in 3-day-old newborn infants.
Although the existence of a coupling between vision and posture seems now well
established at birth and throughout early development, the quality of this coupling remains
 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284 273

to be investigated. In other words, to what extent do infants’ postural reactions covary with
the optical flow characteristics? A first and important answer to this question has been
recently provided by Bertenthal, Rose and Bai (1997). In a series of studies, 5, 7, 9 and 13
month-olds were tested while sitting in an moving room that oscillated at two different
frequencies (0.3 and 0.6 Hz). Postural sway was quantified as the center of pressure of the
seat mounted to a force plate. The postural sway responses of the infants covaried with the
oscillations of the moving room and the magnitude of this covariation increased with age.
Such investigations should be extended to younger infants, and especially to newborn infants
for whom the visual postural coupling is unaffected by previous postural or visual experi-
ence. How newborns regulate the timing and magnitude of their postural reactions according
to the optical flow properties remains an unanswered question. The present research repre-
sents an initial effort to address this issue. Specifically, the present research aims to test the
newborns’ sensitivity to kinematic properties of the optical flow.

2. Method

2.1. Subjects

Twenty-five (13 female and 12 male) 3-day-old (SD: 1.5) Caucasian, middle class,
newborn infants were studied in this experiment. All subjects had APGAR Scores of at least
8, had normal deliveries and were not on medication. These infants were directly recruited
through the maternity ward of Belvédère hospital in the Rouen community, which served a
wide variety of socioeconomic backgrounds.

2.2. Apparatus

The apparatus consisted of a visual stimulation and a response device. The visual
stimulation device was an adaptation of the one used by Bonnet (1987). The infants were
installed in a specially designed baby seat that allowed them to remain in a seated position
(reclined 25 degrees). They had a 14 inch monochrome monitor on each side of their head.
The monitors were placed parallel to each other and 31cm apart. The entire apparatus was
placed inside a dark isolated experimental chamber. The visual stimuli consisted of two
identical pseudorandom dot patterns (23 cm wide ⫻ 18 cm high) computed and displayed on
the monitors (see Fig. 1). Their angular size covered 56 degrees of each peripheral visual
field. The white/black ratio was 33:100. The two patterns moved horizontally across the
screens at the same constant velocity. In addition to a baseline condition with motionless
patterns, seven experimental velocities were used. The physical velocities were 0.01, 0.02,
0.04, 0.06, 0.08, 0.10 and 0.12 m/sec which correspond to angular perceived velocities of 2.5,
5.0, 10.0, 15.0, 20.0, 25.0 and 30.0 degrees per second (see appendix for the details).
The response device was composed of two air bags orthogonally positioned along the
occipital-temporal part of the skull. The pressure in the air bags was sensitive to any shift in
the center of the head mass. Head pressure was continuously monitored by two pressure
274 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

Fig. 1. Pseudo-random dot patterns used as bilateral optical stimulation.

transducers and digitally sampled by a 12-bit A/D converter at a rate of 60 Hz. The
integration of left and right signals reflected the head movements along the anterior-posterior
sagittal X-axis.
An infrared video camera was also used to record the infant’s behavioral state. The entire
experimental procedure was computer-driven. A special multitasking unit was used to
generate optic flow on the monitors, sample head mass center deviations and control the
order of the trials.

2.3. Procedure

All the infants were observed in the morning after their bath and less than one hour after
feeding. Each subject performed one baseline trial before each of the seven experimental
trials. The baseline trial consisted of a 10 s recording of the head pressure with motionless
visual patterns. Each experimental trial consisted of a pre-test, a test and a post-test. For each
pre- and post-test period, the infant was exposed to the motionless patterns for 2 s. Between
pre- and post-test the pattern was moved at a constant angular velocity for 8 s. Half the
infants (13 ss: 7 female and 6 male) were randomly assigned to an increasing order condition
from 2.5 to 30.0 deg/sec. The other half (12 ss: 6 female and 6 male) was assigned to a
decreasing order condition from 30.0 to 2.5 deg/sec.
Each baseline and experimental trial could begin only after the following two conditions
had been met: 1) the infant’s eyes must be open and, 2) the infant’s head must be stable. Head
stability was assessed by two criteria. First, the head pressure standard deviation had to
remain below 25% of the mean inside a permanently sampled mobile window of 60
consecutive measures (1 sec). Secondly, the head pressure slope had to remain within ⫹0.25
and ⫺0.25 inside the same mobile window. When these two criteria were simultaneously
reached the trial could begin. The length of intertrial interval depended on the newborn’s
head stability and lasted approximately 5 s.
 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284 275

Fig. 2. Evolution of individual mean head pressure in both increasing (left part of the graph) and decreasing (right
part of the graph) conditions in relation to the optical flow velocity. Squared line corresponds to the general mean
head pressure observed for each condition.

2.4. Measures

Due to the interindividual differences in terms of head mass and neuro-muscular tonicity,
all the 200 original recordings (200 ⫽ 25 subjects ⫻ 8 trials) were normalized by a
transformation into z values following the usual formula: zx⫽x-m/sd, where m represents the
mean and sd the standard deviation of individual sampled data among the 8 velocity
conditions. In order to limit the aliasing effects of high frequency components, the signals
were also submitted to a low pass filter involving data smoothing with a moving window of
15 samples (60 Hz/4), eliminating all frequencies equal to or above 4 Hz.

3. Results

The results are presented in two main sections. The first one considers optic flow
sensitivity in terms of mean head pressure throughout the stimulation period. In the second
section, optic flow sensitivity is analyzed in terms of head postural response. Thus, for each
experimental trial, a head postural response is defined and its characteristics in terms of
magnitude and latency are studied with respect to optic flow velocity.

3.1. Mean head pressure during the stimulation period

In order to test neonatal sensitivity to optic flow, we simply calculated the mean head
pressure throughout a trial for each velocity condition and for each subject. We then tested
the mean pressure difference in reference to the motionless baseline condition for each
subject in each experimental condition. Possible order effects due to the increasing or
decreasing sequence of stimulation were first tested. Fig. 2 summarizes, for all subjects in
both order conditions, the evolution of mean head pressure relative to the optic flow velocity.
Mean head pressures were analyzed by means of a 2 ⫻ 8 mixed-model ANOVA with
276 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

Fig. 3. Observed (square) and theoretical (diamond) gain, dispersion (1 SD) and probalities (0.10 to 0.01) as a
function of incremental optical flow velocity.

order condition (Increasing or Decreasing) as a between subjects factor and with velocity
(0 –30 deg/sec) as a within subject factor.
No significant differences between increasing and decreasing velocity order conditions
were found. For each velocity condition, the mean head pressure was comparable across both
orders. When we tested the effect of optic flow velocity, it was clear that the mean head
pressure was greater in all experimental conditions than in the motionless baseline one. All
velocities in Fig. 2, collapsed across order, were found to be significantly above baseline by
performing t tests. Moreover, the ANOVA, which measures the effect of angular velocity on
mean head pressure, reveals a highly significant effect F(7,161) ⫽ 4,52; p ⬍ .0001. Lastly
a linear trend analysis revealed that head pressure is linearly related to velocity (r ⫽ 0.94,
N ⫽ 8, p ⫽ .0005). It can be concluded from this analysis that 3 day-old infants exhibit a clear
response to a backward optic flow which related to the angular velocity of the optic flow.
In order to assess the relation between head pressure magnitude and optic flow velocity,
we reasoned in terms of the gain associated with each of these two variables. We calculated
first the incremental pressure of the head by dividing the mean head pressure recorded for
each experimental condition by the mean head pressure recorded in the slowest velocity
condition (2.5 deg/sec). The incremental velocity of optic flow was then obtained by dividing
each velocity value by the slowest velocity value (2.5 deg/sec) used in the experiment. The
optic flow incremental velocity so defined is equal to 2, 4, 6, 8, 10 and 12 (i.e. 5:2.5 ⫽ 2).
Fig. 3 plots the incremental pressure of the head versus the incremental velocity of optic
flow. The slope of this function represents the gain of the system. As shown, the incremental
pressure of the head is strictly and linearly related to the incremental velocity of optic flow
(F(6,138)⫽36,41; p ⬍ .00001). The increment of head pressure from one velocity condition
to the next is significant, as revealed by paired t test comparisons. Lastly, the observed gain
is highly correlated (r ⫽ 0.97, N ⫽ 6, p ⫽ 0.001) to the theoretical one (equal to 1) which
 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284 277

Fig. 4. Determination of the main dependent variables (magnitude and latency of the response). The vertical solid
lines correspond to the following temporal events: onset and end of the visual motion. The vertical dashed lines
correspond to the calculation of latency. The horizontal dashed lines illustrate the measure of the response
magnitude (see text).

would be expected if the head pressure increment strictly followed the arithmetical progres-
sion of optic flow velocity.

3.2. Head postural response

In most individual recordings it was possible to identify a postural response characterized

mainly by head deflection that was greater in magnitude than other head deflections during
the test phase. In order to isolate such responses (one response per trial), the original data
were submitted to the following analyses (Fig. 4). For each trial, we calculated with, an
automatic data scanning program, the elapsed time for head pressure to reach the maximum
and also the preceding minimum value. The software then calculated the magnitude of the
response as the difference between the maximum and the preceding minimum values of the
recorded data. The latency was defined as the time at which the excursion from the baseline
exceeds 10% of the maximum calculated from the onset of the visual motion.
The magnitude of the head postural response in each velocity condition (from 2.5 to 30.0
deg/sec) was compared to a baseline magnitude from the motionless trial. The baseline
magnitude is operationally defined as the arithmetic difference between the maximum and
minimum head pressure recorded during the baseline trial. A 2 ⫻ 8 mixed-model ANOVA
with order (Increasing or Decreasing) as a between subjects factor and with velocity (0 –30
deg/sec) as a within subjects factor revealed only (Fig. 5) a significant effect of the optic flow
velocity (F(7,161)⫽5,41; p ⬍ .0001).
Repeated One-way ANOVAs that compared the magnitude in each velocity condition to
the magnitude in the baseline motionless condition indicated that in velocity conditions (from
2.5 to 30.0 deg/sec) the magnitude was always significantly greater than in baseline (Table
278 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

Fig. 5. Mean, dispersion (SD) and linear regression of head postural response magnitude as a function of velocity
condition.

1). Lastly, posthoc paired t tests performed on these data reveal 4 clusters of velocities within
which no significant differences are found: velocities under 5 deg/sec, velocities between 5
and 10 deg/sec, velocities between 15 and 25 deg/sec and velocities over 25 deg/sec.
Between the 4 clusters of velocities, significant differences are observed and magnitude is
linearly correlated with the velocity of the optic flow display (r ⫽ 0,94, N ⫽ 8, p ⫽ .001).
The analysis of head postural response magnitude is largely convergent with the analysis
of head mean pressure magnitude throughout the stimulation period. Significant postural
response can be identified even in the slowest (2.5 deg/sec) velocity condition. Lestienne,
Soechting and Berthoz (1977) have demonstrated that for adults standing on a force platform,
the minimum velocity of peripheral stimulation required to produce a significant postural
change is less than 0.02 m/sec (5.0 deg/sec). This means that, expressed in linear velocity,
the same velocity induces a significant postural response in both neonates and adults.
Postural responses observed in neonates seem to be roughly similar to those observed in
adults and, sensitivity seems to be in the same order of magnitude.
The 2 ⫻ 8 mixed-model ANOVA with order (Increasing or Decreasing) as a between
subjects factor and with velocity (0 –30 deg/sec) as a within subjects factor failed to reveal

Table 1
Repeated one-way ANOVAs for each velocity condition compared to the baseline motionless condition
Velocity (deg/sec) F1-49 P
2.5 3,04 0.02
5.0 3,59 0.02
10.0 6,10 0.01
15.0 6,14 0.001
20.0 4,75 0.001
25.0 6,13 0.001
30.0 5,65 0.001
 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284 279

a significant effect of the optic flow velocity on latency to respond. Latencies of head postural
response obtained in neonate have to be discussed because they present some impressive
similarities with identical indices recorded in standing adults. First, latencies obtained in
human adults do not depend on the increase of optic flow velocity from 0.02 to 2.7 m/sec
(Lestienne et al., 1976; Lestienne, Soechting & Berthoz, 1977). These authors, and also
Flückiger and Baumberger (1988), observed that the latency, defined as the time at which the
excursion from the baseline exceeds 10% of the maximum of the postural reaction induced
by optical flow, is equal to 1.2 ⫾ 0.3 sec. The value we obtained in the neonate is of course
higher: 1.90 ⫾ 1.75. This means that the latency of the postural reaction is slightly shorter
in standing adult than in sitting neonate.

4. Discussion

The present study unambiguously confirms that infants are sensitive to optic flow within
3 days after birth (Jouen, 1988; Jouen & Lepecq, 1989, Jouen, 1990). When exposed to
backward flow, newborns react with a backward leaning of the head. More impressively,
mean head pressure is also tightly coupled to velocity. The magnitude of the head pressure
significantly and linearly increases with the velocity of the optic flow. Finally, the postural
head responses are velocity tuned. This visually induced modulation of head posture as soon
as birth raises three kinds of problems with respect to the ontogenetic determinants of such
sensitivity.
As revealed by experiments with adults (Berthoz, Pavard & Young, 1975; Lestienne,
Soechting & Berthoz, 1977), the thresholds of velocity for self-motion perception and
visually induced postural readjustments are within the same limits as those for detection of
the visual images themselves. Similarly, our results are consistent with data collected in
infants concerning the perception of visual motion and velocity thresholds. Using the visual
preference method and an infant-controlled habituation procedure, Slater, Morison, Town
and Rose (1985) have demonstrated that infants are sensitive to a rotating or translating
stimulus as soon as birth. Much of the additional research is directed toward somewhat older
infants (Kaufmann, Stucki & Kaufmann-Hayoz, 1985; von Hofsten, Kellman & Putaansu,
1992; Banton & Bertenthal, 1997). Volkmann and Dobson (1976) have studied 1- to
3-month-olds’ preferences for a moving versus an identical adjacent motionless display.
Infants demonstrated visual preference for motion that increased asymptotically with veloc-
ity up to a peak of 15.6 deg/sec. Aslin and Shea (1990) and Dannemiller and Freedland
(1989) both reported absolute thresholds for vertical motion in the 3-month-olds to be
approximately 5.0 deg/sec. Bertenthal and Bradbury (1992) tested infants with random dot
kinematograms and reported even lower thresholds ranging from 3.5 deg/sec at 3 months to
1.2 deg/sec at 5 months. Using linear motion of bars, Dannemiller and Freedland (1991)
found that 20-week-old infants discriminated 3.3 deg/sec from 5.0 deg/sec but failed to
discriminate 5.0 from 10.0 deg/sec. Lastly, research with OKN (Rosander & von Hofsten,
2000) or smooth pursuit (Phillips et al., 1997) showed sensitivity to differential velocity
before 2–3 months of age.
However the measures used in these experiments do not require scaling of visual infor-
280 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

mation to a postural response which probably involves different neural networks (Bertenthal,
1996). As a consequence, the rate-limiting factor for postural responding might not be the
detection of image motion, but rather applying the appropriate force in response to the
velocity of this motion. No anatomical study has demonstrated direct connections between
the axons of the retinal ganglionic cells and the motoneurons involved in the tonic activity
of the dorsal cervical extensors. Consequently, modulation of muscular activity by vision is
assumed to involve indirect neural tracts (Jouen, 1990). Various neurons have been described
in the central nervous system; they are related to peripheral vision and have appropriate
direction and velocity coding to be used by the oculomotor system or to modulate the activity
of the peripheral vestibular system (Stein & Meredith, 1993).
Apart from the visual projection to the vestibular nuclei, brain stem structures that relay
visual information are indirectly connected to the vestibular neurons. The accessory optical
system (AOS) receives input from the retina and relays visual signals to the vestibulo-
cerebellum (Simpson, Soodak & Hess, 1979). AOS neurons have large receptive fields and
respond optimally to textured patterns of 20 degrees. Most of the cells are sensitive to field
movement between 0.05 and 1 degree/sec. Single-cell recordings in the nucleus of the optic
tract (NOT) of the pretectum have demonstrated a particular sensitivity of these cells to large
and slowly horizontally moving patterns. Precht and Cazin (1979) and Precht and Strata
(1980) have shown that in rats and cats, lesions in the NOT eliminate or modify the responses
of central vestibular neurons to optokinetic stimuli. The distribution of preferred-direction of
AOS and pretectal neurons reveals an organization along spatial planes that coincides exactly
with the orthogonal organization of the vestibular semicircular canals. As information in the
vestibular system is also coded in vector components in these 3 planes, AOS and pretectal
neurons would be expected to be directly involved in visual-vestibular interactions. Neurons
of the superior colliculus are also known to respond greater to moving stimuli than to
stationary visual ones. Vestibular influences also have been demonstrated in the lateral
geniculate body (Papaioannou, 1973).
Another possible pathway consists of the cortical projections from primary visual areas to
the pontine nuclei, as demonstrated by Brodal (1978). Single neurons in the pontine nuclei
are movement- and direction-sensitive and have large receptive fields. Moreover, the fibers
in the pontine nuclei project to the cerebellum and subsequently to the vestibular nuclei. All
these brain stem visual structures involved in the coding of the direction and optical
information specifying self-motion are known to be mature at birth (Banks & Salapatek,
1984; Stein & Meredith, 1993).
The neonate’s head postural responses observed in the present study are remarkably
consistent with global postural reactions exhibited later in development. Indeed, the back-
ward leaning of the head in response to backward flow is similar to the backward leaning of
the whole body observed under analogous conditions in older infants (Bertenthal & Bai,
1989; Bertenthal, 1990; Butterworth, 1992; Butterworth & Hicks, 1977, Delorme, Fringon &
Lagacé, 1989; Jouen, 1988; Jouen & Lepecq, 1989; Lee & Aronson, 1974; Sveistrup, Foster
& Woollacott, 1992), in children (Baumberger, 1993) and in adults (Fluckiger & Baum-
berger, 1988; Dichgans & Brandt, 1978; Lestienne et al., 1976; Lestienne, Soechting &
Berthoz, 1977; Lishman & Lee, 1973). Similarly, the variation of neonates’ head response
magnitude with the flow velocity is identical to global postural reactions observed in adults,
 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284 281

in whom the magnitude of the body leaning has been shown to linearly increase with optic
flow velocity from 0.02 up to 1.1 m/sec (Lestienne et al., 1976; Lestienne, Soechting &
Berthoz, 1977). The major characteristics of global postural reactions in response to optic
flow observed throughout infancy, childhood and adulthood are thus present at a local
postural level within three days after birth. The present findings thus suggest that learning is
not necessary for the emergence of optic flow sensitivity, although experience, and partic-
ularly self-produced mobility, may play a role in the subsequent development of visual
postural coupling (Gibson, 1966; Bertenthal, 1990).
A considerable amount of psychophysiological and neurophysiological data supports the
fact that the control of head posture implies multimodal integrative and cooperative pro-
cesses in which visual, vestibular and proprioceptive captors are involved (Berthoz, 1989;
Guitton et al., 1986). Studies on eye-head coordination and gaze stabilization have demon-
strated that visual inputs could modulate the activity of the dorsal neck muscles and, by
extension, head motion. Visually induced electromyographic (EMG) responses can be
recorded under the influence of optokinetic stimulation. During upward vertical linear
optokinetic stimulation, a correlation between the direction and velocity of the slow phase
upward vertical component of optokinetic nystagmus (OKN) and the compensating EMG
activity of splenius and longus capitis muscles (Borel, Lacour & Xerri, 1988; Lacour, Vidal
& Xerri, 1981) has been observed. Other authors have shown that a modulation of the
vestibulocollic reflex (VCR), mainly involved in the postural head control, is induced by an
eye position signal in a fixed head as well as in a free head condition (Crommelinck,
Roucoux & Veraart, 1982; Vidal, Roucoux & Berthoz, 1982). These interactive processes
between the visual system and neck muscles are controlled by at least four types of
subcortical neurons: second order neurons from lateral and median vestibular nuclei, tecto-
reticulo-spinal neurons and reticulo-spinal neurons (for a review see Berthoz, 1989). The
visual inputs are conveyed by at least two important visual projection areas: the superior
colliculus (SC) and the optic tectum nuclei (OTN), or pretectum, which are both known to
be functionally mature at birth (Stein & Meredith, 1993). The present findings thus suggest
that the subcortical neural networks involved in visual proprioceptive control of posture are
functional as soon as birth.

Acknowledgments

We owe thanks to J.J. Lockman for his helpful comments about the first version of this
article, and to S. Margules and S. Harpner for their aid in translating.

Appendix

Relation between the linear velocity of the stimulation and the visual angle (from Berthoz,
Pavard & Young, 1975).
The angular velocity d␣/day␶ of a point belonging to the stimulation is a function of:
282 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

a) the linear velocity of visual stimulation (LV: from 0.01 to 0.12 m/s)
b) the distance between the eye and the screen (d’: 0.155 m)
c) the angle of incidence (␣:56°).
Trigonometric considerations show that d␣/day␶ (in deg/sec)⫽180/3.14*LV/day’*sin2␣.

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