European Journal of Neuroscience, pp. 1–7, 2014 doi:10.1111/ejn.

12777

Evidence for a role of corrective eye movements during

gaze fixation in saccade planning
rez Zapata,1 Marıa Sole
Laura Pe  Puig,1 Jose Antonio Aznar-Casanova1,2 and Hans Supe
r1,2,3
1
Department of Basic Psychology, University of Barcelona, Pg. Vall d0 Hebron 171, 08035 Barcelona, Spain
2
Institute for Brain, Cognition and Behavior, Barcelona, Spain
3
Catalan Institution for Research and Advanced Studies, Barcelona, Spain

Keywords: corollary discharge, corrective eye movement, efferent copy, human, oculomotor, saccadic eye movement

Abstract
In a three-dimensional (3D) world most saccades are made towards visual targets that are located at different distances. We previ-
ously demonstrated that gaze shifts within 3D space consist of two stages: a target saccade followed by a corrective saccade during
gaze fixation that directs the eyes to the physical target location. We proposed that, by accurately positioning the eyes on the visual
object, the visual system maintains an orderly representation of the visual world. In this study we used a double saccade experiment
to assess the function of corrective saccades in humans. We found that, when a corrective eye movement occurred during fixation
on the first target point, the direction of the second saccade towards the next target point was accurate. When a corrective saccade
was absent, a directional error of the second target saccade was observed. This finding, which cannot be explained by current mod-
els of eye movement control, supports the idea of a two-step model in saccade programming. We suggest that the motor system
sends a corollary discharge when programming a corrective saccade for maintaining an orderly representation of the visual world. In
conclusion, our results indicate that corrective saccades have a role in programming target saccades within 3D space.

Introduction
In a three-dimensional (3D) world most saccades are made towards
visual targets that are located at different distances from the obser-
ver. Misperceptions, such as underestimations of the distance
between the observer and the object, are affected by the depth at
which the object is located (Cutting & Vishton, 1996; Foley et al.,
2004; Wagner, 2006; Aznar-Casanova et al., 2011). Consequently,
correctly gazing at targets in a 3D environment should involve fast
and robust mechanisms to prevent perceptual error accumulation.
To correctly guide saccadic eye movements to visual goals, the
oculomotor system makes use of bottom-up saliency (Masciocchi
et al., 2009), perceptual interpretations of the stimuli (e.g. Melcher
& Kowler, 1999; Super et al., 2003; Super et al., 2004; Vishwanath
& Kowler, 2004; de Grave et al., 2006; Super & Lamme, 2007) and
corollary discharge (Sommer & Wurtz, 2004, 2008; Joiner et al.,
2010) that signal information about eye position relative to space
(Ghasia et al., 2008). However, this corollary discharge may not
carry all of the information about the 3D visual environment
because eye position representations to higher cortical centers of the
superior colliculus, where corollary discharge originates, operate
in two-dimensional visual rather than 3D motor coordinates (Van
Opstal et al., 1991; Hepp et al., 1993; Klier et al., 2003).
In a recent study (Perez Zapata et al., 2013), we demonstrated
that gaze shifts to far targets in a 3D setting consist of two stages: a
Correspondence: Hans Super, 1Department of Basic Psychology, as above.
E-mail: hans.super@icrea.cat
Received 23 April 2014, revised 29 September 2014, accepted 7 October 2014
large eye movement toward the perceived target location followed
by a corrective saccade that directs the eyes to the physical target
location. In a two-dimensional setting we did not find evidence for
corrective saccades (Perez Zapata et al., 2013), suggesting that these
corrective saccades during gaze fixation have a specific role in cor-
recting depth information.
By accurately positioning the eyes on the visual object before
producing the next saccade the visual system maintains an orderly
representation of the 3D visual world. In general, corrective sac-
cades are thought to reduce the distance between the saccade land-
ing site and target location (Becker & Fuchs, 1969). However, other
studies suggest that corrective saccades can increase the distance
between the gaze position and target positions (Lemij & Collewjin,
1989), and also that the latency, amplitude, and orientation of cor-
rective saccades are modulated by target eccentricity (Ohl et al.,
2011; Wang et al., 2011). Thus, the precise function of corrective
saccades is not clear. In particular, the role of corrective saccades
after a target saccade in 3D space is unknown.
To test a possible role of corrective saccades in updating the change
in visual location during gaze fixation we used a double saccade
experiment in a 3D setting. As these corrective saccades seem, at least
in part, to be visually driven (Perez Zapata et al., 2013) the rate of
occurrence of corrective eye movements was manipulated by varying
the latency of onset/offset of the first target point. We then analysed
the direction of the second target saccade in order to assess the role of
corrective eye movements in saccade planning (Joiner et al., 2010).
The direction of the target saccade towards the second target was
accurate when a corrective saccade occurred during the eye fixation

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
2 L. Perez Zapata et al.

Fig. 1. Illustration of the two-step saccade experiment in a 3D setting. The
double-step paradigm was presented on a 30° inclined screen.
on the first target. When no corrective saccade was observed, the
direction of the second target saccade was inaccurate. Our data indi-
cate that corrective saccades may play a role in the programming of
target saccades within 3D space.
Materials and methods
Observers
We recorded the eye positions of six volunteers (three males and three
females, mean age 25 years, range 21–28 years; two of them were
authors of this study) participating in the experiments. They all had
normal visual acuity and stereo vision. Detailed instructions regarding
the experiments were given to the six observers. They all provided
written informed consent before taking part in the study, which was
conducted in accordance with the Helsinki Declaration and approved
by the Ethics Committee of the University of Barcelona.
Eye movement recording
Eye positions were recorded at a sampling rate of 500 Hz with a
head-mounted video-based tracking system (EyeLink II, SR
Research, ON, Canada). Head movements were prevented using a
dental imprint bite-bar (Bite-Buddy, UHCOTECH Head Spot, Uni-
versity of Houston). Each recording session lasted for about 4 min.
Before each session the gain and linearity of the eye tracker were
calibrated (nine-point calibration) for both horizontal and vertical
eye positions. For all participants, the mean of the errors reported
after the calibration procedure was of the order of 0.15  0.05°.
Task
The observers were seated in a fully-lit room (40 cd) that provided
access to all visual cues (surface slant and rectangular frame of pro-
jection). Each observer faced a swinging screen (49 9 39 cm)
slanted at 30° at a viewing distance to the base of 89 cm. The trial
began with a retro-projected (Pico projector; PK102, Optoma) black
screen with a white point of 0.02° (starting point) in the lower part
of the screen.
The observers were required to fixate on this point. After
removal of the starting point an upper point (first target; size,
0.02°) was presented at a vertical distance of 10.13° from the start-
ing point (Fig. 1). The observers had to move their gaze to this tar-
get point by making a saccadic eye movement. When the eyes
started to fixate on the target, i.e. the eyes entered a virtual region
of 3° surrounding the target point, the first target point was
removed and a second target (size, 0.02°) at a distance of 9.16° to
either the right or left side of the first target point was briefly
(50 ms) presented. The observers had to shift their gaze to this sec-
ond target point.
The two main reasons for this choice of experimental set-up were
as follows: (i) fixation mislocalisation for reactive saccades seems
not to be modulated by initial eye position (Zimmermann & Lappe,
2011), and (ii) previous studies indicate that corrective saccades
occur more frequently when saccades coincide with the direction of
the depth gradient (near to far) (Perez Zapata et al., 2013).
The interval between the arrival of the eyes at the first target and
the removal/presentation of the first/second target [response stimulus
interval (RSI)] was randomly set to 10, 20, 50, 60, 100, 150, 250,
and 300 ms (Panouilleres et al., 2011). In a separate block, an RSI
of 500 ms was used as a control. Each observer performed 10
blocks of 32 trials in two sessions. Each block lasted about 3 min.
A few trials were performed as training before the experimental ses-
sions began. Visual stimulation software was developed and pre-
sented using EventIDE (OkazoLab Ltd, UK).
Calculation of saccadic eye movements
To calculate the saccade direction we transformed the HRef record-
ings (X and Y coordinates of the left and right eye), provided by
the EyeLink II software, into angular units using algorithms
designed to calculate the 3D components (Sx, Sy, Sz) of both eye
gaze vectors (Grossberg et al., 1992). Thus, the 3D setting was
taken into account when analysing the collected data. These angu-
lar measures implied a transformation from the screen-collected
gaze positions to a 3D space where eye movement amplitudes
showed a misjudgment of the physical depth of the presentation
screen.
Sx,y,z are the three coordinates that represent the intersection point
of the lines describing the line of sight of each eye. When these
lines do not intersect this represents the midpoint of the smallest dis-
tance between them. The transformation was performed taking into
account the real distance between the screen and the observer [dis-
play–subject distance (DSD)] and the actual interpupil distance
(IPD), and converting them, taking into account a factor of 15 000
HRef/cm (which is a transformation given by the EyeLink II com-
mercial software) (section 4.4.2.2 ‘HREF’ in the EyeLink II User
Manual). Sx,y,z are calculated as follows:

IPDðIPD2 ðY1  Yr ÞðY1 þ Yr Þ þ 4IPDðXr Y12  X1 Yr2  ðXr þ X1 ÞDSD2 Þ þ 4ðXr2 ðY12 þ DSD2 Þ  X12 ðYr2 þ DSD2 ÞÞÞ
Sx ¼
2ð2Xr Y1 þ 2X1 Yr þ IPDðY1 þ Yr ÞÞ2 þ 8ððIPD þ X1  Xr Þ2 þ ðY1  Yr Þ2 ÞDSD2
2IPDðY1 Yr ð2X1 Yr þ 2X1 Yr þ IPDðY1 þ Yr ÞÞ þ ðIPD þ X1  Xr ÞðY1 þ Yr ÞDSD2 Þ
Sy ¼
ð2Xr Y1 þ 2X1 Yr þ IPDðY1 þ Yr ÞÞ2 þ 4ððIPD þ X1  Xr Þ2 þ ðY1  Yr Þ2 ÞDSD2
IPDðY1 þ Yr Þð2Xr Y1 þ 2X1 Yr þ IPDðY1 þ Yr ÞÞDSD þ 4IPDðIPD þ X1  Xr ÞDSD3
Sz ¼
ð2Xr Y1 þ 2X1 Yr þ IPDðY1 þ Yr ÞÞ2 þ 4ððIPD þ X1  Xr Þ2 þ ðY1  Yr Þ2 ÞDSD2

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
European Journal of Neuroscience, 1–7
 Corrective eye movements during gaze fixation 3

From these components we calculated the saccade amplitude by

(i) azimuth (horizontal component of the eye movement) and (ii) lat-
itude (vertical component of the eye movement):


Sx
AzimuthðaÞ ¼ arctan ð1Þ
Sz
!
Sy
LatitudeðaÞ ¼ arctan pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi ð2Þ
Sx þ S2z
2

Data analysis
For each condition (RSI) the onset, amplitude, and velocity of the
first and second target saccade were calculated. Saccades were iden-
tified based on a velocity criterion. An eye movement velocity of
over 60°/s followed by a decrease to below 0.05°/s within a window
of 50 ms was used to detect saccades. These saccades were then
categorised as first target saccade, corrective saccade (i.e. they
occurred in a time interval from the start of fixation on the first
target to the onset of the saccade to the second target), and second
target saccade.
The location of eye fixation was computed for two different peri- Fig. 2. A random sample of 30 individual trials (500 ms RSI) showing the
ods. Early fixation location was the average gaze position between traces of eye positions. We observed saccades that were followed by a cor-
90 and 130 ms after saccade onset towards the first target. This time rective eye movement (black lines) and saccades without a subsequent cor-
period occurs during target fixation and well before the onset of a rective eye movement (gray lines). Saccade amplitude is represented as
version changes in visual degrees. Dotted lines show the average eye posi-
corrective saccade (Perez Zapata et al., 2010, 2013). Late fixation tion of all trials. Arrows point to some corrective eye movements during fix-
location was the average gaze position between 380 and 420 ms ation. Inset shows the distribution of the onset latencies of all corrective eye
after saccade onset, which is after the corrective saccade onset. For movements.
the fixation location at the second target we used the average eye
position in a window of 150 ms starting at the moment that the sac-
cade velocity fell below 0.05°/s. To detect the latency and onset of
the corrective saccade we carried out a similar procedure to that
used in Super et al. (2004). Trials containing eye blinks and those
in which observers moved their eyes in whatever direction (right or
left) before fixating on the first target (anticipated eye movements)
were filtered out.

Results
We first classified the data from the RSI-500 condition according to
the presence or absence of a corrective eye movement. This revealed
that, in 73% of all trials, saccades towards the first target were fol-
lowed by a corrective eye movement (Fig. 2, arrows). The mean
onset latency of these eye movements was 213  4.7 ms
(mean  SEM) (Fig. 2, inset), calculated from the start of the first
saccade, and is in agreement with our previous report on corrective
eye movements (Perez Zapata et al., 2013). For the other conditions,
the results showed that the rate of corrective eye movements gradu-
ally decreased from RSI-300 (42%) to RSI-10 (11%) (Fig. 3). The
mean onset of the corrective eye movement across all conditions
was 217.7  4.20 ms (mean  SEM). No significant differences in
the onset latencies of corrective eye movements were found between
conditions (t312 = 0.12; P > 0.05). The corrective eye movements
Fig. 3. The frequency of trials with and without a corrective eye movement
had a mean velocity of 84.3  3.8°/s, with a mean amplitude of for the different conditions (RSI).
0.9  0.03°.

Amplitudes and velocities of the first target saccade
We assessed the landing points of the eyes at the first target, before
and after the corrective eye movement (Fig. 4a and b). For the trials
from all RSI conditions with a corrective eye movement, the landing
points during the early fixation period were different from the gaze
positions during the late fixation period, i.e. after the corrective eye
movement (Fig. 4c) (t312 = 2.7; P < 0.01). The mean amplitude
of the target saccade was 9.40  0.09° (mean  SEM) and, follow-
ing the correction of eye position, gaze was located near the physi-
cal location of the first target (mean  SEM: 10.11  0.08°), which
was located at a distance of 10.13° from the fixation point (Fig. 4c,

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
European Journal of Neuroscience, 1–7
4 L. Perez Zapata et al.

A
B

Fig. 4. Eye movement traces during the task (A and B) and average saccade amplitude to the first target (C). (A) Scan path of the eyes for trials with (black
traces) and without (gray traces) corrective saccades (just a sample of trials is shown for clarity). White squares (not at scale) represent the positions of the tar-
gets. (B) Average eye positions over time for trials with (black trace) and without (gray trace) corrective saccades. Shaded vertical bars represent the time win-
dows for the calculations of the landing positions. (C) Mean saccade amplitude to the first target for early (white) and late (black) fixation location. Horizontal
dashed line represents physical target distance. *Statistical (P < 0.01) difference. Error bars represent SEM.

horizontal dashed line). Analysis of amplitudes for different RSIs

revealed no differences for trials with either short or long RSI.
The landing points did not differ between the early and late fixa-
tion positions for trials without a corrective eye movement
(t443 = 1.76, P = 0.08). This observation was supported by the
finding that the mean amplitude of the first target saccade,
9.73  0.07° (mean  SEM), was similar to the saccade amplitude
when taking the gaze position at the late fixation period
(mean  SEM: 9.75  0.07°) (Fig. 4b and c) as the landing point.
The comparison between trials with and without corrective eye
movements showed no difference in the mean amplitude of the first
target saccade when taking as the landing point the gaze position
during the early fixation location (t312 = 1.70; P = 0.09). How-
ever, when the gaze position during the late fixation location was
taken as the landing point, the saccade amplitude was higher for the
trials with a corrective eye movement (t312 = 2.5; P < 0.01).

Onset of the second target saccade

We then calculated the onset latency of the second target saccade
(calculated from the onset of fixation on the first target). For trials
with a corrective eye movement, the mean onset latency of the sec-
ond target saccades was longer than the mean onset latency of the
target saccades for trials without corrective eye movements
(mean  SD: 692.4  241 ms vs. 505.8  197 ms; t340 = 11.24;
Fig. 5. Onset latencies of the second target saccade for each RSI condition
separately. Error bars represent SEM.
P < 0.0001). This was particularly the case for the intermediate RSI
values, whereas for the short and long RSIs no difference between
trials with and without corrective saccade was observed (Fig. 5).

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
European Journal of Neuroscience, 1–7
 Corrective eye movements during gaze fixation 5

Direction of the second target saccade agreement with previous findings (Perez Zapata et al., 2013), we
observed that ~75% (in the RSI-500 condition) of saccades to a dis-
To test whether the absence of a corrective eye movement has conse-
tant target were followed by a corrective eye movement. This per-
quences for saccade programming, we studied the accuracy of the
centage dropped (in the RSI-10 to RSI-300) when the first target
direction of the target saccade towards the second target in trials with
point was removed directly after fixation on it (Becker & Fuchs,
and without a corrective eye movement, and separately for long
1969; Bonnetblanc & Baraduc, 2007; Ohl et al., 2013).
(500 ms) and short (10–300 ms) RSIs. In order to assess the accuracy
of the direction of the target saccade we applied the method used by
Joiner et al. (2010). Hence we calculated two angles: the observed Control of corrective eye movements
(a0 ) and the expected (a) angle. The observed angle was defined as the
Corrective saccades may position the eyes at the visual object in
angle formed by the vector connecting the gaze position at the starting
depth if the landing point of the target saccade is inaccurate (Perez
point to the gaze position at the first target point during the late fixa-
Zapata et al., 2013). Thus, a saccade error of the first target saccade
tion period (i.e. after the occurrence of corrective eye movement) and
may explain the occurrence of corrective saccades. A saccade error
the vector connecting this latter gaze position to the gaze position at
may derive from the non-linear dynamics of saccades (Collewijn
the second target. The expected angle (a) was defined as the angle
et al., 1988), incorrect target mapping (Semmlow et al., 1993,
between the vector connecting the gaze position at the starting point to
1994), or conflicting sensory information (Perez Zapata et al.,
the gaze position at the first target during the late fixation period and
2013). Alternatively, insufficient 3D information or an inaccurate
the vector connecting the latter gaze position to the physical location
corollary discharge signal accompanying a target saccade may result
of the second target (Joiner et al., 2010) (Fig. 6).
in saccade error. Here we found correction of eye gaze after saccad-
The expected and observed angles in trials with corrective eye
ing to a distant target. Other authors have observed a corrective
movements were significantly different from those found in trials
saccade when the target is a simple, isolated visual object in a two-
without corrective eye movements (expected: t312 = 12.12;
dimensional setting (e.g. Becker & Fuchs, 1969; Ross et al., 2001;
P < 105; observed: t312 = 9.62; P < 105) (Fig. 7). For saccades
Bonnetblanc & Baraduc, 2007; Ohl et al., 2013), thereby suggesting
that were followed by a corrective eye movement, the mean value
that corrective eye movements are a common phenomenon.
(mean  SEM: 90.61  0.98°) of the observed angles was similar
Our observed corrective eye movements occurred at ~200 ms
(t312 = 0.67, P = 0.5) to the mean value (mean  SEM:
after the onset of fixation, which is indicative of visual control. This
90.21  0.79°) of the expected angles, and neither angle type dif-
is supported by our findings showing that removal of the target
fered significantly (P > 0.05) from 90°. By contrast, for saccades
information from the fovea suppresses the occurrence of corrective
that were not followed by a corrective eye movement, the mean
eye movements. It is also in line with recent findings showing that
value (mean  SEM: 96.61  0.44°) of the observed angles was
fixational eye movements are visually driven (Ko et al., 2010;
different from the mean (mean  SEM: 99.95  0.38°) of the
Poletti et al., 2013). However, corrective eye movements are also
expected angles (t312 = 5.83; P < 105) (Fig. 7).
observed after a saccade made in complete darkness (Leigh & Zee,
We also analysed the error in the angle of the second saccade,
2006) or without visual feedback (Ohl et al., 2013), which indicates
using the difference between the expected and the observed angle as
that the correction does not solely depend on visual information.
a dependent variable in an ANOVA and taking the presence/absence of
The simultaneous disappearance/appearance of the first/second target
the corrective saccade and each RSI condition as factors. The pres-
may suppress the occurrence of corrective eye movements, although
ence or absence of a corrective saccade was shown to be a significant
this does not explain the suppression of corrective saccades in the
factor (F1,871 = 7.37; P = 0.006), whereas neither the RSI condition
RSI-300 and RSI-500 conditions, in which the removal and presen-
nor the interaction were significant (F8,871 = 1.26, P = 0.263 and
tation of the target points occur after the time when corrective sac-
F8,871 = 1.16, P = 0.318, respectively). Taken together, these obser-
cades occur.
vations indicate that goal-directed saccades are more accurate once
It may be argued that, on trials with high oculomotor readiness,
the eye position has been corrected during fixation.
parallel programming takes place and no corrective saccades occur.
These trials would then show less accurate second saccades and this
Discussion is more likely to happen in trials with short RSIs. However, an expla-
nation of parallel programming of both first and second saccade is
In this study, we used a double saccade task to test whether correc-
doubtful. Firstly, the task was designed so that the second saccade
tive eye movements may play a role in oculomotor control. In
appeared only when the eyes landed at the first target location. There-
fore, the subjects could not know the location of the second target at
the moment when the first saccade was planned or executed (McPeek
et al., 2000). Secondly, the different conditions were randomly inter-
leaved and the observers were unaware of the RSIs.

Role of corrective eye movement in oculomotor control

Fig. 6. Schematic explanation of the observed and expected angle. Thick
trace represents eye positions over time and thin lines represent the vectors
for the calculation of the observed (gray line) and expected (dashed line)
angles. Positions of the visual targets (not at scale) are represented by the
black squares. Gray dashed circle denote the landing points at the second
target.
We tested the role of corrective eye movements by analysing the
direction of the second target saccade (Joiner et al., 2010). In trials
with a corrective eye movement the second saccade was accurate.
When no corrective saccade was observed the saccade towards the
second target was inaccurate, i.e. the empirically measured saccade
direction did not correspond to the expected saccade direction.
One possible explanation for the directional error of the second
target saccade would be an inaccurate saccade planning, due, for

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
European Journal of Neuroscience, 1–7
6 L. Perez Zapata et al.

A B

Fig. 7. Angle between the first and second target saccade. (A and B) Comparison of the distributions of the observed and expected angles from saccades with
and without a corrective eye movement. (C) Mean values of observed and expected angles. Error bars represent SEM. *Significant differences (P < 10).

instance, to a too short fixation period. However, we believe that
this is unlikely for three reasons. Firstly, we observed differences
between the expected and observed angles in all RSI conditions,
including the RSI-500 condition, in which saccade planning is not
compromised. Secondly, the minimum time (i.e. 394 ms in the
RSI-10 condition) between the onset of the corrective saccade and
the onset of the second saccade is sufficient to program a visually
guided saccade. Thirdly, in all conditions, including RSI-10, sac-
cades that followed a corrective eye movement were accurate, indi-
cating that saccade planning was correct in these cases. Interference
with the planning of the saccade to the first target is also an unlikely
explanation for these results (Ohl et al., 2013). The first target point
was removed after the execution of the first saccade, at the moment
that the eyes foveated the target location, and this means that
saccade planning was completed. Moreover, we found inaccurate
saccades in the RSI-500 condition when target information remained
present. Another possible explanation for the obtained results is that
the second target saccade showed a strong bias to move purely later-
ally; in other words, it went to the side at an angle of 90°, as if the
gaze were in precisely the same location as the target. Contrary to
this explanation, we found that the observed angles differed between
trials with and without corrective saccades. In trials with a corrective
eye movement the observed angles of the second saccade were
indeed around 90°, whereas in trials without a corrective saccade
the observed angles were significantly larger than 90°. However,
they did not reach the expected value for angular size. Thus, the
visual system was not able to completely correct for the saccade
error. We therefore suggest that when a corrective saccade, to
correct eye gaze in 3D space, is absent during fixation, the direction
of the next saccade is inaccurate.
Possible implications in updating visual location
Eye movements are internally monitored by sending the sensory cor-
tex an efferent copy (corollary discharge) of the planned saccade
(Sommer & Wurtz, 2008). The visual system uses this signal to
update the current location of the eyes (Joiner et al., 2010). We
show that the occurrence of a corrective saccade probably depends
on the presence of the fixation point at the time of eye fixation. This
means that visual information is needed to produce a corrective eye
movement, and that the corrective saccade is not pre-programmed
together with the target saccade. Accordingly, the motor system
needs to send a corollary discharge when programming a corrective
saccade so that the visual system is able to update for the change in
the visual location and to maintain an orderly representation of the
visual world.
Acknowledgements
This work was supported by grants to H.S. from the Spanish Ministry of
Education and Science (MICINN; PSI2010-18139 and SAF2009-10367) and
the Catalan government (AGAUR; 2009-SGR-308). We would like to thank
Juan Manuel Garcia for his technical support.
Abbreviations
3D, three-dimensional; RSI, response stimulus interval.

© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd
European Journal of Neuroscience, 1–7

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