Joen Optic Flow Sensitivity in Neonates Vison Infants

Optic flow sensitivity in neonates

Francois Jouena,*, Jean-Claude Lepecqb, Olivier Gapennec,Bennett I. Bertenthald

a Laboratoire de Psychobiologie du Developpement, 41 rue Gay Lussac, EPHE/CNRS, F-75005Paris, France

bNeurosciences Cognitives et Imagerie Cerebrale, LENA, UPR CNRS 640, F-75651 Paris, FrancecCOSTEC, Universite Technologique de Compiegne F-60000 Compiegne, France

dDepartment 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 infantswere placed inside a dark room and observed while presented with a 10 s bilateral and backwardperipheral 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. Sagittaldeviation of the head was computer-sampled at 60 Hz frequency with pressure transducers. Asindicated by the mean head pressure during optic flow exposure, infants reacted with backwardleaning 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 relatedto 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 evidencebackward postural reactions and report a sensation of forward motion. Such a sensation fallsunder the category of visually-induced illusory body motion, or vection, and in this particularcase 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: [email protected] (F. Jouen).

Infant Behavior & Development 23 (2000) 271–284

0163-6383/00/$ – see front matter © 2000 Elsevier Science Inc. All rights reserved.PII: S0163-6383(01)00044-3

adults exhibit forward postural reactions and report a feeling of backward motion (backwardlinear vection). Generally, adults display postural reactions in the very direction of the flowand 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 hasbeen devoted to the postural reactions shown by infants when exposed to optical flow. Suchreactions are now theoretically framed as a part of perception-action couplings in earlyontogeny (Bertenthal, 1996).

Lee and Aronson (1974) were the first to show that infants’ posture may be affected byvisual information. Babies who had recently learned to stand were tested on a motionlessfloor, within a moveable room. The infants faced the interior end wall and the wholestructure, except the floor, was moved so that the end wall approached or receded. Subjectscompensated for a nonexistent loss of balance signaled by the optic flow pattern and fell inthe direction of the optic flow. If the end wall moved away from the baby, the infant fellforward and if the wall moved toward the baby, the infant fell over backward. Peripheralvision is particularly important for maintaining postural stability. Pope (1984) showed thatin 3-month-old infants, movement in the center of the visual field did not result in significantpostural adjustment, whereas a slight movement in the periphery is sufficient to induce acomplete loss of stability. These results have been extended by Bertenthal and co-workers ininfants aged between 5 and 9 months (Bertenthal & Bai, 1989, Bertenthal, 1990 for areview). Using an enclosure that permitted independent movement of the front and sidewalls, these authors showed that 7- and 9-month-olds compensated in a directionallyappropriate reaction to whole-room movement. Nine-month-olds, but not 7-month-oldsresponded with systematic compensation to side wall movement suggesting a developmentaltrend 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) alsodemonstrated that 8.5-month-old infants without locomotor experience showed minimalcompensation to peripheral optic flow. In contrast, 8.5-month-old babies with crawling orwalker 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 theonset of locomotion, they do not imply that infants are insensitive to peripheral optic flowprior the acquisition of locomotor skill. Butterworth and Hicks (1977) found that infants tooyoung to walk but nevertheless capable of unaided sitting compensate for visually specifiedmotion when seated in a moving room. This finding has been extended by Pope (1984) andJouen (1986) to younger infants. They investigated the effects of optical flow exposure on thecontrol of head posture in 2-month-old infants. They found that even infants unable to sitwithout support perform directionally appropriate compensatory movements of the headwhen stimulated with optic flow. Testing even younger infants, Jouen (1988) and Jouen andLepecq (1989) provided evidence for directionally appropriate head movements under roughoptical flow in 3-day-old newborn infants.

Although the existence of a coupling between vision and posture seems now wellestablished at birth and throughout early development, the quality of this coupling remains

272 F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

to be investigated. In other words, to what extent do infants’ postural reactions covary withthe optical flow characteristics? A first and important answer to this question has beenrecently provided by Bertenthal, Rose and Bai (1997). In a series of studies, 5, 7, 9 and 13month-olds were tested while sitting in an moving room that oscillated at two differentfrequencies (0.3 and 0.6 Hz). Postural sway was quantified as the center of pressure of theseat mounted to a force plate. The postural sway responses of the infants covaried with theoscillations 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 infantsfor 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 accordingto 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 thenewborns’ 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 least8, had normal deliveries and were not on medication. These infants were directly recruitedthrough the maternity ward of Belvedere hospital in the Rouen community, which served awide variety of socioeconomic backgrounds.

2.2. Apparatus

The apparatus consisted of a visual stimulation and a response device. The visualstimulation device was an adaptation of the one used by Bonnet (1987). The infants wereinstalled 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 wasplaced inside a dark isolated experimental chamber. The visual stimuli consisted of twoidentical pseudorandom dot patterns (23 cm wide � 18 cm high) computed and displayed onthe monitors (see Fig. 1). Their angular size covered 56 degrees of each peripheral visualfield. The white/black ratio was 33:100. The two patterns moved horizontally across thescreens at the same constant velocity. In addition to a baseline condition with motionlesspatterns, 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 theoccipital-temporal part of the skull. The pressure in the air bags was sensitive to any shift inthe center of the head mass. Head pressure was continuously monitored by two pressure

273F. Jouen et al. / Infant Behavior & Development 23 (2000) 271–284

transducers and digitally sampled by a 12-bit A/D converter at a rate of 60 Hz. Theintegration of left and right signals reflected the head movements along the anterior-posteriorsagittal X-axis.

An infrared video camera was also used to record the infant’s behavioral state. The entireexperimental procedure was computer-driven. A special multitasking unit was used togenerate optic flow on the monitors, sample head mass center deviations and control theorder of the trials.

2.3. Procedure

All the infants were observed in the morning after their bath and less than one hour afterfeeding. Each subject performed one baseline trial before each of the seven experimentaltrials. The baseline trial consisted of a 10 s recording of the head pressure with motionlessvisual patterns. Each experimental trial consisted of a pre-test, a test and a post-test. For eachpre- and post-test period, the infant was exposed to the motionless patterns for 2 s. Betweenpre- and post-test the pattern was moved at a constant angular velocity for 8 s. Half theinfants (13 ss: 7 female and 6 male) were randomly assigned to an increasing order conditionfrom 2.5 to 30.0 deg/sec. The other half (12 ss: 6 female and 6 male) was assigned to adecreasing order condition from 30.0 to 2.5 deg/sec.

Each baseline and experimental trial could begin only after the following two conditionshad been met: 1) the infant’s eyes must be open and, 2) the infant’s head must be stable. Headstability was assessed by two criteria. First, the head pressure standard deviation had toremain below 25% of the mean inside a permanently sampled mobile window of 60consecutive measures (1 sec). Secondly, the head pressure slope had to remain within �0.25and �0.25 inside the same mobile window. When these two criteria were simultaneouslyreached the trial could begin. The length of intertrial interval depended on the newborn’shead stability and lasted approximately 5 s.

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


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 atransformation into z values following the usual formula: zx�x-m/sd, where m represents themean and sd the standard deviation of individual sampled data among the 8 velocityconditions. In order to limit the aliasing effects of high frequency components, the signalswere also submitted to a low pass filter involving data smoothing with a moving window of15 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 flowsensitivity in terms of mean head pressure throughout the stimulation period. In the secondsection, optic flow sensitivity is analyzed in terms of head postural response. Thus, for eachexperimental trial, a head postural response is defined and its characteristics in terms ofmagnitude 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 headpressure throughout a trial for each velocity condition and for each subject. We then testedthe mean pressure difference in reference to the motionless baseline condition for eachsubject in each experimental condition. Possible order effects due to the increasing ordecreasing sequence of stimulation were first tested. Fig. 2 summarizes, for all subjects inboth 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

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


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 conditionswere found. For each velocity condition, the mean head pressure was comparable across bothorders. When we tested the effect of optic flow velocity, it was clear that the mean headpressure was greater in all experimental conditions than in the motionless baseline one. Allvelocities in Fig. 2, collapsed across order, were found to be significantly above baseline byperforming t tests. Moreover, the ANOVA, which measures the effect of angular velocity onmean head pressure, reveals a highly significant effect F(7,161) � 4,52; p � .0001. Lastlya 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 clearresponse 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 calculatedfirst the incremental pressure of the head by dividing the mean head pressure recorded foreach experimental condition by the mean head pressure recorded in the slowest velocitycondition (2.5 deg/sec). The incremental velocity of optic flow was then obtained by dividingeach velocity value by the slowest velocity value (2.5 deg/sec) used in the experiment. Theoptic 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 opticflow. The slope of this function represents the gain of the system. As shown, the incrementalpressure 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 conditionto the next is significant, as revealed by paired t test comparisons. Lastly, the observed gainis highly correlated (r � 0.97, N � 6, p � 0.001) to the theoretical one (equal to 1) which

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


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 characterizedmainly by head deflection that was greater in magnitude than other head deflections duringthe test phase. In order to isolate such responses (one response per trial), the original datawere submitted to the following analyses (Fig. 4). For each trial, we calculated with, anautomatic data scanning program, the elapsed time for head pressure to reach the maximumand also the preceding minimum value. The software then calculated the magnitude of theresponse as the difference between the maximum and the preceding minimum values of therecorded data. The latency was defined as the time at which the excursion from the baselineexceeds 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 30.0deg/sec) was compared to a baseline magnitude from the motionless trial. The baselinemagnitude is operationally defined as the arithmetic difference between the maximum andminimum head pressure recorded during the baseline trial. A 2 � 8 mixed-model ANOVAwith order (Increasing or Decreasing) as a between subjects factor and with velocity (0–30deg/sec) as a within subjects factor revealed only (Fig. 5) a significant effect of the optic flowvelocity (F(7,161)�5,41; p � .0001).

Repeated One-way ANOVAs that compared the magnitude in each velocity condition tothe magnitude in the baseline motionless condition indicated that in velocity conditions (from2.5 to 30.0 deg/sec) the magnitude was always significantly greater than in baseline (Table

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


1). Lastly, posthoc paired t tests performed on these data reveal 4 clusters of velocities withinwhich no significant differences are found: velocities under 5 deg/sec, velocities between 5and 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 islinearly 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 analysisof head mean pressure magnitude throughout the stimulation period. Significant posturalresponse 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 posturalchange 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 inadults and, sensitivity seems to be in the same order of magnitude.

The 2 � 8 mixed-model ANOVA with order (Increasing or Decreasing) as a betweensubjects factor and with velocity (0–30 deg/sec) as a within subjects factor failed to reveal

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

Table 1Repeated one-way ANOVAs for each velocity condition compared to the baseline motionless condition

Velocity (deg/sec) F1-49 P

2.5 3,04 0.025.0 3,59 0.0210.0 6,10 0.0115.0 6,14 0.00120.0 4,75 0.00125.0 6,13 0.00130.0 5,65 0.001


a significant effect of the optic flow velocity on latency to respond. Latencies of head posturalresponse obtained in neonate have to be discussed because they present some impressivesimilarities with identical indices recorded in standing adults. First, latencies obtained inhuman 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 alsoFluckiger and Baumberger (1988), observed that the latency, defined as the time at which theexcursion from the baseline exceeds 10% of the maximum of the postural reaction inducedby optical flow, is equal to 1.2 � 0.3 sec. The value we obtained in the neonate is of coursehigher: 1.90 � 1.75. This means that the latency of the postural reaction is slightly shorterin standing adult than in sitting neonate.

4. Discussion

The present study unambiguously confirms that infants are sensitive to optic flow within3 days after birth (Jouen, 1988; Jouen & Lepecq, 1989, Jouen, 1990). When exposed tobackward 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 pressuresignificantly and linearly increases with the velocity of the optic flow. Finally, the posturalhead responses are velocity tuned. This visually induced modulation of head posture as soonas birth raises three kinds of problems with respect to the ontogenetic determinants of suchsensitivity.

As revealed by experiments with adults (Berthoz, Pavard & Young, 1975; Lestienne,Soechting & Berthoz, 1977), the thresholds of velocity for self-motion perception andvisually induced postural readjustments are within the same limits as those for detection ofthe visual images themselves. Similarly, our results are consistent with data collected ininfants concerning the perception of visual motion and velocity thresholds. Using the visualpreference method and an infant-controlled habituation procedure, Slater, Morison, Townand Rose (1985) have demonstrated that infants are sensitive to a rotating or translatingstimulus as soon as birth. Much of the additional research is directed toward somewhat olderinfants (Kaufmann, Stucki & Kaufmann-Hayoz, 1985; von Hofsten, Kellman & Putaansu,1992; Banton & Bertenthal, 1997). Volkmann and Dobson (1976) have studied 1- to3-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 beapproximately 5.0 deg/sec. Bertenthal and Bradbury (1992) tested infants with random dotkinematograms and reported even lower thresholds ranging from 3.5 deg/sec at 3 months to1.2 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 todiscriminate 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 velocitybefore 2–3 months of age.

However the measures used in these experiments do not require scaling of visual infor-


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 thedetection of image motion, but rather applying the appropriate force in response to thevelocity of this motion. No anatomical study has demonstrated direct connections betweenthe axons of the retinal ganglionic cells and the motoneurons involved in the tonic activityof the dorsal cervical extensors. Consequently, modulation of muscular activity by vision isassumed to involve indirect neural tracts (Jouen, 1990). Various neurons have been describedin the central nervous system; they are related to peripheral vision and have appropriatedirection and velocity coding to be used by the oculomotor system or to modulate the activityof the peripheral vestibular system (Stein & Meredith, 1993).

Apart from the visual projection to the vestibular nuclei, brain stem structures that relayvisual information are indirectly connected to the vestibular neurons. The accessory opticalsystem (AOS) receives input from the retina and relays visual signals to the vestibulo-cerebellum (Simpson, Soodak & Hess, 1979). AOS neurons have large receptive fields andrespond optimally to textured patterns of 20 degrees. Most of the cells are sensitive to fieldmovement between 0.05 and 1 degree/sec. Single-cell recordings in the nucleus of the optictract (NOT) of the pretectum have demonstrated a particular sensitivity of these cells to largeand 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 responsesof central vestibular neurons to optokinetic stimuli. The distribution of preferred-direction ofAOS and pretectal neurons reveals an organization along spatial planes that coincides exactlywith the orthogonal organization of the vestibular semicircular canals. As information in thevestibular system is also coded in vector components in these 3 planes, AOS and pretectalneurons would be expected to be directly involved in visual-vestibular interactions. Neuronsof the superior colliculus are also known to respond greater to moving stimuli than tostationary visual ones. Vestibular influences also have been demonstrated in the lateralgeniculate body (Papaioannou, 1973).

Another possible pathway consists of the cortical projections from primary visual areas tothe pontine nuclei, as demonstrated by Brodal (1978). Single neurons in the pontine nucleiare movement- and direction-sensitive and have large receptive fields. Moreover, the fibersin the pontine nuclei project to the cerebellum and subsequently to the vestibular nuclei. Allthese brain stem visual structures involved in the coding of the direction and opticalinformation 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 remarkablyconsistent 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 ofthe whole body observed under analogous conditions in older infants (Bertenthal & Bai,1989; Bertenthal, 1990; Butterworth, 1992; Butterworth & Hicks, 1977, Delorme, Fringon &Lagace, 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 responsemagnitude with the flow velocity is identical to global postural reactions observed in adults,


in whom the magnitude of the body leaning has been shown to linearly increase with opticflow 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 opticflow observed throughout infancy, childhood and adulthood are thus present at a localpostural level within three days after birth. The present findings thus suggest that learning isnot 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 visualpostural coupling (Gibson, 1966; Bertenthal, 1990).

A considerable amount of psychophysiological and neurophysiological data supports thefact 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, byextension, head motion. Visually induced electromyographic (EMG) responses can berecorded under the influence of optokinetic stimulation. During upward vertical linearoptokinetic stimulation, a correlation between the direction and velocity of the slow phaseupward vertical component of optokinetic nystagmus (OKN) and the compensating EMGactivity 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 thevestibulocollic reflex (VCR), mainly involved in the postural head control, is induced by aneye position signal in a fixed head as well as in a free head condition (Crommelinck,Roucoux & Veraart, 1982; Vidal, Roucoux & Berthoz, 1982). These interactive processesbetween the visual system and neck muscles are controlled by at least four types ofsubcortical neurons: second order neurons from lateral and median vestibular nuclei, tecto-reticulo-spinal neurons and reticulo-spinal neurons (for a review see Berthoz, 1989). Thevisual inputs are conveyed by at least two important visual projection areas: the superiorcolliculus (SC) and the optic tectum nuclei (OTN), or pretectum, which are both known tobe functionally mature at birth (Stein & Meredith, 1993). The present findings thus suggestthat the subcortical neural networks involved in visual proprioceptive control of posture arefunctional as soon as birth.

Acknowledgments

We owe thanks to J.J. Lockman for his helpful comments about the first version of thisarticle, 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:


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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Joen Optic Flow Sensitivity in Neonates Vison Infants

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