Multisensory Research 31 (2018) 455–480 brill.

com/msr

Amending Ongoing Upper-Limb Reaches:

Visual and Proprioceptive Contributions?

Rachel Goodman, Valentin A. Crainic, Stephen R. Bested,

Darrin O. Wijeyaratnam, John de Grosbois and Luc Tremblay ∗
Perceptual Motor Behaviour Laboratory, Centre of Motor Control, Faculty of Kinesiology and
Physical Education, University of Toronto, Toronto, ON, M5S 2W6, Canada

Received 14 October 2016; accepted 2 November 2017

Abstract
In order to maximize the precise completion of voluntary actions, humans can theoretically utilize
both visual and proprioceptive information to plan and amend ongoing limb trajectories. Although
vision has been thought to be a more dominant sensory modality, research has shown that sensory
feedback may be processed as a function of its relevance and reliability. As well, theoretical models
of voluntary action have suggested that both vision and proprioception can be used to prepare online
trajectory amendments. However, empirical evidence regarding the use of proprioception for online
control has come from indirect manipulations from the sensory feedback (i.e., without directly per-
turbing the afferent information; e.g., visual–proprioceptive mismatch). In order to directly assess the
relative contributions of visual and proprioceptive feedback to the online control of voluntary actions,
direct perturbations to both vision (i.e., liquid crystal goggles) and proprioception (i.e., tendon vibra-
tion) were implemented in two experiments. The first experiment employed the manipulations while
participants simply performed a rapid goal-directed movement (30 cm amplitude). Results from this
first experiment yielded no significant evidence that proprioceptive feedback contributed to online
control processes. The second experiment employed an imperceptible target jump to elicit online
trajectory amendments. Without or with tendon vibration, participants still corrected for the target
jumps. The current study provided more evidence of the importance of vision for online control but
little support for the importance of proprioception for online limb–target regulation mechanisms.

Keywords
Motor control, proprioception, tendon vibration, online control, vision

* To whom correspondence should be addressed. E-mail: luc.tremblay@utoronto.ca

© Koninklijke Brill NV, Leiden, 2018 DOI:10.1163/22134808-00002615

456 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

1. Introduction
Upper-limb movements are involved in most activities of daily living. Sensory
systems are used to gather information from the environment to plan and ex-
ecute these voluntary movements. Additionally, sensory information can be
used to make corrections to ongoing body movements. Specific to the current
study, although the initial portion of a limb trajectory ought to reflect the ac-
tion plan (e.g., Wolpert and Ghahramani, 2000), corrections can be made to
that initial plan, in order to accurately complete the movement (i.e., online
control: e.g., Woodworth, 1899). Such online control processes can be neces-
sary considering the limb trajectory may not be accurately planned (e.g., lack
of experience), subject to noise during its implementation (e.g., tremor due to
a disease or aging) or simply requiring an amendment due to a shift of the rel-
ative positions between the limb and the target (e.g., confronting an opponent
in a sport setting).
Recently, Elliott and colleagues (2010) described a theoretical model of
upper-limb online control, consisting of multiple sub-processes that occur
during a typical discrete reaching movement. Namely, impulse regulation pro-
cesses were identified as the early comparisons of the actual and expected
movement-related sensory inputs. Additionally, limb–target regulation pro-
cesses were identified as the comparison of the relative locations of the limb vs.
the target. Studies that contributed to the elaboration of the multiple-processes
model also indicated that online limb trajectory amendments could require the
use of visual and proprioceptive information (e.g., Ghez et al., 1995; Grierson
and Elliott, 2008, 2009; Scott et al., 2015). The current study aimed to fur-
ther investigate the contribution of vision and proprioception to online control
mechanisms during voluntary action.
1.1. The Importance of Vision
Visual feedback is known to be the more dominant sense to gather spatial
estimations (e.g., Rock and Victor, 1964), which are crucial to both plan and
execute the movement. In terms of the online control of upper-limb reaches,
vision has been deemed to be critical for both impulse regulation and limb–
target regulation processes (see Elliott et al., 2010). According to Elliott et al.,
(2010: see also Elliott et al., 1991), vision can be utilized to implement online
control processes throughout most of the trajectory. More recently, support for
the multiple-processes model was found although the use of visual feedback
for online control mechanisms appeared to be important for very brief periods,
early in the limb trajectory.
In line with the idea that limb velocity is a critical component for the prepa-
ration and control of goal-directed action (e.g., Ashe and Georgeopoulos,
1994; Churchland et al., 2006; Tremblay and de Grosbois, 2015), Tremblay
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 457

and colleagues recently manipulated vision based on real-time limb velocity

in a series of experiments. In Tremblay et al. (2013), vision was provided
above or below velocity criteria that ranged from 0.8 m/s to 1.4 m/s. In that
study, some dependent variables were deemed to be more closely associated
with impulse regulation processes (e.g., endpoint variability, which is calcu-
lated irrespective of the target location) while other variables were deemed
to be more associated with limb–target regulation processes (e.g., correla-
tion coefficients between limb position during the trajectory vs. position at
movement end: see Fisher2 variable in the Section 2.1). Specifically, vision
manipulations influenced endpoint variability the most when the velocity cri-
teria changed from 0.8 to 0.9 m/s whereas vision manipulations influenced the
limb vs. target position correlation coefficients the most when the velocity cri-
teria changed from 1.0 to 1.1 m/s. Subsequently, to determine which particular
portion of the trajectory was vision most relevant to online control processes
(i.e., before vs. after peak limb velocity), Kennedy et al. (2015) provided three
brief windows of vision to participants, all presented when the limb was trav-
elling above 0.8 m/s. As compared to when vision was provided throughout
the entire movement, participants exhibited comparable endpoint variability
with the early (0.8 to 1.4 m/s, before peak limb velocity) and the middle win-
dow of vision (above 1.4 m/s, including peak limb velocity). However, the
middle window also yielded longer times spent during the limb deceleration
phase (i.e., time after peak limb velocity). As such, the late window yielded
worse endpoint variability than the control vision condition and both the mid-
dle and late windows yielded longer limb deceleration durations. In contrast,
only the early window of vision — which lasted a mere 43 ms — yielded
comparable endpoint variability and timing than when vision was available
throughout the trajectory. Overall, Kennedy et al. (2015) provided evidence
that the visual samples that contribute the most to online control processes
are typically very early in the movement. Further, in the second experiment of
Tremblay et al. (2017), brief visual samples were provided at 0.6 m/s, 1.0 m/s
and 1.4 m/s, while limb–target regulation processes were perturbed via un-
perceivable target jumps on one third of the trials (see Goodale et al., 1986).
Although one could propose that obtaining unconscious information about the
displaced target should provide more time to correct the limb trajectory when
gathered earlier in the movement (i.e., 0.6 m/s), participants only significantly
corrected their movement endpoint distributions when the window of vision
was provided upon reaching the 1.0 m/s threshold. These results strength-
ened Tremblay et al.’s (2013) proposal that limb–target regulation processes
are mainly implemented when the limb velocity reaches 1.0 m/s. Altogether,
there is a large body of literature that is consistent with Elliott et al.’s (2010)
multiple-processes model (e.g., Cressman et al., 2010; Elliott et al., 1991; Gri-
erson and Elliott, 2008, 2009; Proteau et al., 2009; Sarlegna and Mutha, 2015;
458 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

Saunders and Knill, 2003) and the use of real-time movement-dependent vi-
sion manipulations provided a significant amount of additional support (i.e.,
Kennedy et al., 2015; Tremblay et al., 2013, 2017).

1.2. The Importance of Proprioception

Although it is evident that visual information is useful during upper-limb

reaching movements, it has been proposed that other sensory modalities are
important for sensorimotor control, such as the information regarding muscle
length and rate of change of muscle length (i.e., proprioception: type Ia spin-
dle afferents: e.g., Gardner and Johnson, 2013). Such proprioceptive feedback
is thought to contribute to the online control of limb movements, which is
supported by the notion that people can complete movements in the absence
of any visual feedback (Woodworth, 1899). Evidence for the contribution of
proprioceptive information to sensorimotor control was provided in a large
number of studies. For example, Rossetti and colleagues (1995) used optical
prisms to displace the seen environment, which led to corrections for upper-
limb trajectories. They reported that movement direction was biased towards
the visual information but that participants only partially adjusted for the op-
tical displacement. That is, the average final finger location was somewhere
between the visual displaced hand and the actual target. Thus, it was deemed
that proprioception had played a significant role in determining the finger tra-
jectory. As a second example, Sainburg et al. (1993) employed a bread slicing
pantomime task with deafferented human patients. Patients were capable to
perform the task when vision was available, while removal of visual feed-
back permitted performance but with large performance decrements. As such,
in line with the multiple processes model of online control, proprioceptive
feedback is thought to be important to control upper-limb reaches. It must be
noted, however, that in both provided examples, proprioception could also be
contributing to the planning of voluntary movement. The current study aims
to focus on the online control of discrete action.
While it is easy to manipulate visual information (i.e., occlude the eyes
and/or the target), it is more challenging to manipulate proprioceptive feed-
back. The aforementioned experiments investigating the use of visual and
proprioceptive information either indirectly perturbed proprioception (e.g.,
visual–proprioceptive mismatching task in Rossetti et al., 1995) or employed
proprioceptive manipulations that can also elicit reflexive muscle contractions
(e.g., external limb perturbations in Grierson and Elliott, 2008). As a result,
the current study aimed to assess the contributions of vision and propriocep-
tion to the online control of an upper-limb goal-directed action by employing
direct manipulations for both modalities. Specifically, liquid-crystal goggles
were used to directly perturb vision (see Milgram, 1987). Also, two high in-
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 459

tensity tendon vibrators were used to directly perturb proprioceptive feedback

(see Roll et al., 1989), although such manipulations were implemented using
a relatively novel approach.
Tendon vibration has been used to directly perturb proprioceptive feedback,
at least since the well-cited study by Goodwin et al. (1972). Vibrations applied
to a muscle tendon have been shown to affect the majority of Type Ia muscle
spindles situated within the associated muscle (i.e., proprioceptive feedback:
Roll and Vedel, 1982; Roll et al., 1989). The use of tendon vibration during
reaching movements has also yielded evidence that proprioceptive information
is important during voluntary actions (e.g., Capaday and Cooke, 1981; Redon
et al., 1991). Redon and colleagues (1991) specifically used tendon vibration
to assess proprioceptive feedback during various parts of a discrete movement,
and discovered that tendon vibration yielded the largest endpoint biases when
applied during the later stages of movements. However, because these studies
manipulated proprioception during the movements, the results of these ma-
nipulations could be explained by the muscle reflex activity arising from the
tendon vibration. This potential modulation is the reason why the current study
employed a proprioceptive after-effect arising from tendon vibration.
Ribot-Ciscar and colleagues (1998) discovered that tendon vibration
elicited altered Ia muscle spindle activity after the associated tendon (i.e.,
tibialis anterior, extensor digitorum longus and lateral peroneal) has been vi-
brated. Specifically, immediately after the removal of the tendon vibration,
73% of Ia muscle spindles exhibited a significantly smaller firing rate while
13% exhibited a significantly greater firing rate. As a result, the firing rates
exhibited by the Ia muscle spindles after the presentation of tendon vibra-
tion elicit noisier proprioceptive signals than when no vibration is presented.
Building upon this knowledge, a recent study in our laboratory tested the use
of between-trial tendon vibration in a simple limb-matching paradigm, to in-
vestigate the ability for the aftereffects of vibration to perturb proprioceptive
feedback. Results revealed that presenting vibration to both the biceps and
triceps brachii muscles simultaneously between movements did elicit greater
variability in a limb-matching task (Goodman, 2015). In a second experi-
ment, a goal-directed upper-limb reach was employed and the vibration again
yielded increased variability at movement endpoint (Goodman, 2015). This
method could be advantageous over other direct or indirect proprioceptive ma-
nipulation methods (e.g., air pulse or visual mismatch respectively) because
it is not intuitive for individuals to determine that the vibration should yield
significant limb position variability and/or the direction of endpoint biases that
the vibration should elicit (cf. forward vs. backward air pulse in Grierson and
Elliott, 2008).
460 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

1.3. The Current Study

The purpose of the current study was to further explore the contribution of
proprioceptive and visual information to the online control of goal-direction
action, specifically in reference to the multiple processes model of online con-
trol. A first experiment was performed to investigate the contributions of vision
and proprioception to the online control of action when reaching to a single
target (i.e., single-target experiment). If both vision and proprioception con-
tribute to online control mechanisms, then endpoint variability should increase
with both visual occlusion and tendon vibration manipulations (see Tremblay
et al., 2013). This could be interpreted as evidence that both vision and pro-
prioception are important for impulse regulation mechanisms. As well, if both
modalities are important for online control processes, then limb trajectories
should be corrected to a lesser extent and appear to be more stereotypical when
either manipulation is applied (see Tremblay et al., 2013). Such a result would
support the idea that vision and proprioception are important for limb–target
regulation processes (Elliott et al., 2010). In contrast, if only visual occlu-
sion yields increased endpoint variability and more stereotypical trajectories,
then such results could be interpreted as evidence that only vision significantly
contributes to online impulse regulation and limb–target regulation processes.
Also, because Elliott et al.’s (2010) multiple-processes model explicitly posits
that vision and proprioception contribute to limb–target regulation processes,
a second experiment was performed to directly test this proposal by utiliz-
ing a target jump protocol, with and without tendon vibration (i.e., target
jump experiment, see Tremblay et al., 2017). If proprioception significantly
contributes to online limb–target regulation processes, then target jump cor-
rections should be evidently corrected in the absence of tendon vibration but
less or not corrected when tendon vibration is applied between trials. In con-
trast, if only vision significantly contributes to online limb–target regulation
processes (or if vision is weighed significantly more than proprioception), then
limb trajectory endpoints should adjust to target jumps even in the presence of
tendon vibration between trials.

2. Single-Target Experiment
2.1. Methods
Fourteen (10 female) participants were recruited from the University of
Toronto community (mean age = 20.5 years). All participants were right-
handed with normal or corrected-to-normal vision and passed a brief neuro-
logical questionnaire (i.e., self-declared neurologically intact). The participant
was seated at a table facing a custom aiming console, which was equipped
with a translucent polymer surface and a Velcro™ home position (7 mm
square). The participants lined up the body midline with the home position.
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 461

The polymer surface hid the target LED (5 mm in diameter) until illuminated.
The participant was outfitted with liquid-crystal goggles (Translucent Tech-
nologies Inc., Toronto, ON, Canada: Milgram, 1987), which manipulated the
entire visual field. An infra-red emitting diode (IRED) was affixed to the tip of
the participant’s index finger and was digitized by an Optotrak Certus motion
tracking system (Northern Digital Inc., Waterloo, ON, Canada) to record 3D
kinematic data at 250 Hz. A piezoelectric buzzer (Mallory Sonalert Products
Inc., Indianapolis, IN, USA, Model SC628, tone frequency of 2900 Hz) was
used to signal movement start and end. Finally, two cylindrical tendon vibra-
tors (VB100, Dynatronic, Valence, France; diameter: 30 mm; length: 75 mm;
weight: 125 g; vibration: 100 Hz, 0.5 mm) were placed on the distal tendons
of the biceps brachii and triceps brachii muscles with polyethylene foam wrap.
Optotrak data gathering and device control were performed via a custom Mat-
Lab script (The MathWorks Inc., Natick, MA, USA) on a standard personal
computer equipped with a PCI6024-E analog-to-digital board (National In-
struments Inc., Austin, TX).
Participants were instructed to perform a 30 cm movement from the home
position (i.e., lined up with their midline) to the target, which was located at
45° clockwise of the participant’s midline (see Fig. 1). Movements were per-
formed as accurately as possible, while falling within a movement time band-
width of 290–400 ms. The data collection protocol involved a familiarization
phase and two experimental sessions of 60 trials each. These sessions were to
divide the presentation of the vibration (Vib) and no-vibration (NoVib) con-
ditions and the order was counterbalanced across participants. A mandatory
15-minute break was instituted between sessions, to diminish any effects of the
tendon vibration if presented in the first session and to include the same break
if tendon vibration was presented in the second session. The tendon vibrators
were attached to the participant for both sessions. Each session included two
blocks, one each for the full vision (FV) and the no-vision (NV) conditions,
which were also counterbalanced across participants. Prior to the initiation
of the vibration session, the tendon vibrators were activated for a total of
60 s to ensure the activation of the Ia fibers as well as the hypo- and hyper-
sensitization following the end of vibration (see Ribot-Ciscar et al., 1998).
A typical trial began with the participant’s finger on the home position with
the goggles closed (i.e., no-vision of the environment). Then, a 4 s foreperiod
took place, which included tendon vibration, if in the appropriate session. The
start of the movement was then signaled by the occurrence of an 800 Hz tone
lasting 50 ms, generated by the piezoelectric buzzer, in conjunction with the
opening of the goggles (i.e., goggles turned transparent). Depending on the
vision condition of the trial, the goggles either remained transparent (i.e., FV)
or turned translucent (i.e., NV) upon movement start, which was defined as the
point in time when the velocity of the IRED (located on the finger) surpassed
462 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

Figure 1. Depiction of the experimental set-up, including the aiming board and the liquid-
crystal goggles and tendon vibrators worn by the participant. The simple target experiment only
employed the 30 cm target (filled circle) while the target jump experiment also employed the
27 cm target (dashed circle) used during a target jump trial. The arrows indicate the direction of
the positive values along primary (Prim) and secondary (Sec) movement axes.

0.03 m/s for two consecutive samples in a row. Following the completion of
the movement, which was determined as the point in time when the velocity
of the IRED fell below 0.03 m/s for three consecutive samples in a row, the
goggles were turned translucent (provided the trial was a FV condition). Once
the kinematic data collection period (1 s) concluded, another single tone was
presented, to indicate to the participant to return to the home position without
visual feedback. The goggles were kept occluded at movement end as well
as during the return to the home position to minimize terminal feedback and
practice effects (see Salmoni et al., 1984).
Multiple variables were calculated from the IRED position data. These vari-
ables included the average location (i.e., constant error, CE) and variability
(i.e., variable error, VE) at movement end, both along the primary (Prim) and
the secondary (Sec) axes (see Fig. 1). Temporal variables were also calculated:
movement time (MT) — the time from movement start to movement end; time
to peak velocity (TtPV) — the time from movement start to peak velocity; time
after peak velocity (TaPV) — the time from peak velocity to movement end. In
addition, peak velocity (PV) was the highest velocity value along the primary
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 463

Table 1.
(Top) Means and between-participant standard deviations for kinematic variables in the single-
target experiment: endpoint accuracy (End), endpoint variability (VE) in the primary axis (Prim)
and in the secondary axis (Sec). Movement time (MT), time to peak velocity (TtoPV), time
after peak velocity (TafterPV), and peak velocity (PV) values. (Bottom) Fisher2 values for the
correlations between movement time proportions and movement end point

No-vibration Vibration
Vision No-vision Vision No-vision
M (SD) M (SD) M (SD) M (SD)

MT 338 (12) 347 (18) 339 (9) 342 (18)

TtoPV 143 (13) 146 (14) 143 (12) 142 (10)
TafterPV 195 (13) 201 (19) 196 (10) 201 (19)
PV 1.81 (0.1) 1.73 (0.2) 1.81 (0.09) 1.84 (0.3)
pPV 146 (9) 140 (14) 146 (8) 147 (15)
VeSec 6 (1) 8 (2) 6 (1) 8 (2)
EndSec −3 (5) 10 (12) −4 (6) 2 (17)
VePrim 9 (2) 14 (5) 9 (3) 13 (3)
EndPrim 299 (4) 293 (19) 301 (5) 306 (34)
Fisher2 25% 0.05 (0.05) 0.12 (0.13) 0.10 (0.11) 0.08 (0.09)
Fisher2 50% 0.16 (0.14) 0.50 (0.49) 0.16 (0.19) 0.34 (0.34)
Fisher2 75% 1.99 (0.86) 3.52 (1.64) 2.01 (1.08) 3.50 (1.12)

axis between movement onset and offset. Finally, correlation coefficients of

the limb position in the primary movement axis at 25, 50 and 75% vs. 100% of
movement time were calculated, Fisher-transformed and squared (Fisher2 ) as
a measure of online control. As suggested by Heath (2005), more stereotyped
trajectories can be interpreted as evidence of fewer trajectory amendments
(see also Bernier et al., 2007). Although we compared the limb position at
movement end with the limb position at three movement time proportions, the
contrasts between 75% and 100% of movement time are of greatest interest
because these have been strongly associated with online visuomotor processes
(see de Grosbois and Tremblay, 2015). All dependent variables were submit-
ted to a 2 vision (FV, NV) × 2 vibration (NoVib, Vib) repeated-measures
ANOVA. All post-hoc analyses were conducted using Bonferroni-corrected
t-tests. Means and standard deviations for all variables are reported in Table 1.
In the interest of conciseness, only the significant effects are discussed. How-
ever, all inferential statistics can be found in Table 2.
2.2. Results
All statistical data for the single-target experiment can be seen in Table 2.
Please refer to the tables for all data values. For clarification, the variables
 464

Table 2.
Inferential statistics from the single-target experiment

MT TtoPV TafterPV PV pPv EndSec VeSec EndPrim VePrim

Vision F -ratio 4.679 0.394 5.371 0.174 0.296 9.381 18.899 0.074 32.530
P -value 0.051 0.542 0.309 0.684 0.597 0.010 0.001 0.790 0.000
Effect size 0.281 0.032 0.309 0.014 0.024 0.439 0.612 0.006 0.731
Vibration F -ratio 0.295 1.945 0.075 4.330 4.085 3.972 0.012 6.301 0.008
P -value 0.597 0.188 0.788 0.060 0.066 0.069 0.915 0.027 0.928
Effect size 0.024 0.139 0.006 0.265 0.254 0.249 0.001 0.344 0.001
Vision* Vibration F -ratio 1.153 1.714 0.103 3.081 8.619 5.176 0.192 3.846 0.993
P -value 0.304 0.215 0.754 0.105 0.012 0.042 0.669 0.073 0.339
Effect size 0.088 0.125 0.009 0.204 0.418 0.301 0.016 0.243 0.076

All significant effects are underlined.

R. Goodman et al. / Multisensory Research 31 (2018) 455–480
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 465

of interest were: finger position at the end of the movement in the primary
axis (EndPrim), variability of finger position at the end of movement in the
primary axis (VePrim), finger position at the end of the movement in the sec-
ondary axis (EndSec), variability of finger position at the end of movement
in the secondary axis (VeSec), movement time (MT), time to peak veloc-
ity (TtPV), maximum velocity achieved (PV), time spent after reaching peak
velocity (TaPV), position of finger at peak limb velocity (pPV), and Fisher2
transforms of the correlation coefficient values between limb position at vari-
ous movement time proportions and limb position at movement end.
Vision main effects: A main effect of vision was significant for final finger
position in EndSec, indicating a rightward (i.e., lateral) reaching bias for the
no-vision conditions (Fig. 2A, B). Also, a main effect of vision was present
for VePrim, VeSec, TaPV, indicating less endpoint variability and shorter de-
celeration phase durations in Full Vision, as compared to no-vision. There was
also a significant effect of vision on the Fisher2 values, as participants exhib-
ited greater Fisher2 values in the no-vision condition (i.e., more stereotyped
trajectories), compared to when vision was provided. This was true for both
the primary and secondary axes.
Vibration main effects: Main effect of vibration was only present in the
final finger position in the primary axis (EndPrim) (Fig. 2 C, D), indicating
movement endpoints further in the vibration than in the no-vibration condition.
Vision by vibration interaction: The interaction between the manipulations
was significant for EndSec, however the associated post-hoc analysis revealed
that endpoints were positively biased in the no-vision as compared to the full
vision condition, but only in the absence of tendon vibration (Fig. 2A). There
was also an interaction between the manipulations for pPV. The associated
post-hoc analysis yielded a significant larger movement amplitude at peak
limb velocity with vibration compared to without vibration, and that is, only
in the no-vision conditions.

2.3. Discussion

Vision occlusion yielded shorter movement amplitudes, larger endpoint vari-

ability, and more stereotyped limb trajectories, which all represent further
evidence for the importance of vision for planning and online control mech-
anisms. Critically, the results of the latter two variables (VePrim and Fisher2 )
could be interpreted as further evidence for the importance of online visual
feedback for impulse and limb–target regulation mechanisms, respectively
(see Tremblay et al., 2013). In contrast, proprioceptive manipulations (i.e.,
tendon vibration) had a significant influence on the limb trajectories (i.e.,
EndPrim) but these main effects were not related to variables associated with
online control processes (i.e., no main effects or interactions associated with
466 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

Figure 2. Results from the single-target experiment. Significant differences for endpoint ampli-
tude in the secondary axis between the no-vibration (A) and vibration condition (B), as well as
in the primary axis (C and D).

vibration for VePrim or Fisher2 ). As well, all the significant results and inter-
actions associated with tendon vibration were related to average limb position
data. Critically, the effect of vibration on movement amplitude was observed
earlier in the trajectory (i.e., at peak limb velocity) only when vision was not
available. As such, it is possible to infer that the emergence of movement am-
plitude differences between peak velocity and movement end in the vision
conditions can be explained by online proprioceptive feedback. In line with
the changes in sensitivity of the type Ia afferent fibers immediately after ten-
don vibration (re.: Ribot-Ciscar et al., 1998), the curvilinear limb trajectories
and endpoints in the presence of vibration can be interpreted as underestima-
tions of the actual distance travelled by the limb. Interestingly, the effects of
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 467

tendon vibration on the endpoint biases in the primary axis were observed ir-
respective of the vision condition at movement end, but only in the no-vision
conditions at peak limb velocity. As a result, it is possible to explain the larger
movement amplitudes with than without vision, to planning mechanisms for
the no-vision condition. However, the larger movement amplitudes observed
by movement end with than without vibration in the vision conditions could be
due to online control mechanisms. Taken together, these results have demon-
strated a significant influence of tendon vibration after-effects on performance
and reaffirmed the importance of proprioception for the execution of voluntary
actions.
Overall, the results from the single-target experiment provided evidence for
the contribution of vision and proprioception to the online control of action.
Elliott et al.’s (2010) model clearly states that both vision and proprioception
contribute to limb–target regulation processes. In order to elicit a relatively
greater contribution of limb–target regulation processes and to provide a more
direct assessment of the use of vision and proprioception for this specific type
of online control, a target jump paradigm was employed in a second experi-
ment (see Goodale et al., 1986; Tremblay et al., 2017).

3. Target Jump Experiment

3.1. Methods
Seventeen participants were recruited from the University of Toronto com-
munity for the second experiment. However, there was a data collection error
(i.e., file overwriting) for two participants, yielding a final sample of fifteen
(6 female; mean age = 22.6 years). There was no overlap in participant re-
cruitment between the first and second experiment. The experimental set-up
and inclusion criteria were the same as in the single-target experiment, with
the exception of the increased sampling rate of the position data to 500 Hz.
This increase was done to ensure that the visual manipulations delay (i.e., due
to the sampling frequency of the Optotrak data, the Matlab processing delays,
the acquisition board transmission delays, and the goggles status change de-
lays) were less than 10 ms. As a result, the visual manipulations were deemed
to take place in real time, at least from a sensorimotor perspective.
Other differences from the single-target experiment included the addition
of a target located at a distance of 27 cm from the home position. That 27 cm
target was inserted for the target jump protocol, creating a 3 cm target jump to-
wards the participant (Fig. 1). In the target jump experiment, the participants
were provided with a 1 s preview of the environment prior to the go signal,
whereas visual information during the movement was limited to a 20 ms win-
dow (see below).
468 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

The target jump experiment was divided in two vibration sessions (i.e., Vib
and NoVib, see above). Vibration was applied to the biceps brachii and triceps
brachii tendons simultaneously (as in the single-target experiment). Prior to
these sessions, participants were provided with 40 familiarization trials in a
full vision (FV) condition and six familiarization trials in the window condi-
tions (see below for description). This relatively long familiarization phase
was employed to ensure that the performance stabilized before the exper-
imental trials, hence increasing the likelihood of detecting limb trajectory
differences between the no-jump and jump trials. Specifically, the initial 20 FV
familiarization trials allowed participants to adjust to the set-up and movement
time constraints (i.e., 350–400 ms), while the latter 20 FV familiarization tri-
als were used in conjunction with electrooculography (EOG) to ensure that
participants looked at the home position prior to each movement and prevent
them from consciously perceiving the target jump. Because EOG data collec-
tion increased processing delays for the manipulations (i.e., real-time visual
occlusion and target jump), it was only employed during the familiarization
phase. The experimental trials consisted of 10 jump (27 cm) and 20 no-jump
(30 cm) target conditions at each of the three velocity criteria (0.6, 1.0, 1.4 m/s)
employed to trigger the 20 ms visual windows (i.e., 90 total trials). Both the
velocity criteria and the target jumps were randomized with the limitation of
not presenting the same condition more than three times in a row.
A typical trial began with the participants placing their right index finger on
the home position without vision of the environment. In the tendon vibration
session, a 60 s vibration period was provided prior to the first trial and also
for 5 s between each trial. Upon the termination of the vibration period, a 1 s
preview of the environment was provided during which the participants were
instructed to look at the home position, to ensure that any target jump would
not be consciously perceived (see Goodale et al., 1986). Upon the go signal,
which was the appearance of the 30 cm target and a 50 ms tone (800 Hz), par-
ticipants were asked to perform a saccade to the target and initiate the reaching
movement. The goggles closed (i.e., vision was removed) 50 ms after the go
signal, so the gaze and finger were still on the home position at that time.
During the trajectory, the IRED velocity was used to trigger the start of the
20 ms visual window (i.e., goggles turned transparent). Depending on the trial
condition, the trigger velocity was set to 0.6, 1.0, or 1.4 m/s, initiating the
window of vision once the velocity has surpassed the trigger velocity for two
consecutive samples. As such, the actual limb velocities during the window
were higher than the criteria (see Tremblay et al., 2017). On one third of the
trials, the 27 cm target was illuminated during the window while the 30 cm
target was illuminated on the remaining two thirds of the experimental trials.
After the reaching movement, a double tone signaled the participant to return
to the home position, without visual information of the environment.
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 469

The Target Jump Experiment employed the same dependent variables and
post-hoc analyses as the single-target experiment. All dependent variables
were subjected to a 2 vibration (No Vib, Vib) × 2 target jump (No Jump —
30 cm, Jump — 27 cm) × 3 window condition (0.6, 1.0, 1.4 m/s) repeated-
measures ANOVA. Also, because the corrections for the target jumps were
hypothesized to maximally take place in the 1.0 m/s window condition, at least
in the absence of tendon vibration (Tremblay et al., 2017), a-priori planned
comparisons were completed between levels of target jumps for both vibra-
tion conditions. Means and standard deviations for each dependent measure
are all reported in Table 3. In the interest of brevity, only significant effects are
reported. However, the inferential statistics for the analyses of all dependent
variables are reported in Table 4.
It was hypothesized that if target jumps elicit limb–target regulation pro-
cesses, especially at 1.0 m/s (Tremblay et al., 2017), then endpoint distri-
butions in the primary movement axis should be closer to the 27 cm target,
at least in the 1.0 m/s window condition. Further, if limb–target regulation
processes rely upon online visual and proprioceptive feedback (Elliott et al.,
2010), then the correction for the target jumps should be significantly smaller
in the absence of tendon vibration. In contrast, if proprioception does not
significantly contribute to online limb–target regulation processes, then the
presence of vibration should have no influence on limb trajectory corrections
arising from target jumps.
3.2. Results
3.2.1. Main Effects of Target Jump
The ANOVAs revealed main effects associated with the target jump manip-
ulation for the following variables: EndPrim, MT, and TaPV. Participants
exhibited shorter reaching amplitudes in the jump trials as compared to the
no-jump trials. The jump trials also yielded shorter MTs and TaPVs than the
no-jump trials.
3.2.2. Main Effects of Vibration
Tendon vibration yielded no significant main effects for any of the dependent
variables.
3.2.3. Two-Way Interactions
The ANOVAs revealed multiple two-way interactions for the vibration × tar-
get comparisons. MT, TAPV, and VePrim yielded a significant two-way inter-
action. Post-hoc analyses were performed on the two-way interactions using a
Bonferroni correction. Post-hoc testing of MT and TAPV solely revealed that
participants completed their movements in less time and spent less time in the
deceleration phase following a target jump, both without and with tendon vi-
bration. Further, the post-hoc contrasts for VePrim and PV failed to yield any
Table 3. 470
Means and between-participant standard deviations for kinematic variables in the target jump experiment: endpoint accuracy (End), endpoint variability
(VE) in the primary axis (Prim) and in the secondary axis (Sec). Movement time (MT), time to peak velocity (TtoPV), time after peak velocity
(TafterPV), and peak velocity (PV) values

No-vibration Vibration
0.6 m/s 1.0 m/s 1.4 m/s 0.6 m/s 1.0 m/s 1.4 m/s
M (SD) M (SD) M (SD) M (SD) M (SD) M (SD)

MT Jump 330 (40) 325 (42) 332 (41) 328 (34) 329 (37) 337 (38)
No-jump 346 (40) 351 (51) 353 (46) 337 (37) 341 (34) 343 (39)
TtoPV Jump 145 (23) 145 (22) 145 (22) 145 (19) 142 (15) 144 (20)
No-jump 145 (19) 145 (19) 145 (20) 143 (16) 145 (20) 145 (17)
TafterPV Jump 185 (23) 180 (28) 187 (24) 183 (25) 187 (26) 193 (25)
No-jump 202 (25) 206 (36) 207 (33) 194 (26) 196 (20) 199 (25)
PV Jump 1.77 (0.2) 1.77 (0.2) 1.75 (0.2) 1.76 (0.2) 1.76 (0.2) 1.76 (0.2)
No-jump 1.74 (0.2) 1.76 (0.3) 1.74 (0.2) 1.77 (0.2) 1.76 (0.2) 1.77 (0.2)
pPV Jump 151 (14) 146 (13) 146 (13) 144 (16) 143 (15) 145 (14)
No-jump 149 (14) 149 (14) 149 (13) 146 (8) 143 (12) 146 (13)
VeSec Jump 5.6 (2.5) 6.5 (3.1) 6.0 (2.4) 6.9 (2.2) 7.2 (2.4) 6.6 (2.4)
No-jump 6.0 (1.9) 6.2 (1.8) 6.2 (1.9) 6.6 (2.0) 6.6 (1.7) 6.8 (1.7)
EndSec Jump −10 (13) −8 (13) −7 (12) −42 (70) −42 (71) −42 (70)
No-jump −8 (14) −7 (11) −6 (11) −41 (71) −41 (70) −42 (70)
R. Goodman et al. / Multisensory Research 31 (2018) 455–480

VePrim Jump 12.0 (4.9) 10.4 (3.6) 11.3 (4.4) 13.8 (6.0) 13.1 (3.6) 13.0 (6.5)
No-jump 12.2 (3.9) 10.8 (3.2) 12.4 (4.8) 11.9 (3.7) 12.3 (3.8) 12.6 (4.6)
EndPrim Jump 311 (24) 308 (22) 316 (26) 309 (26) 307 (22) 312 (26)
No-jump 317 (23) 319 (25) 319 (26) 314 (23) 315 (27) 316 (26)
Table 4.
Inferential statistics from the target jump experiment

MT TtoPV TafterPV PV pPV VeSec EndSec VePrim EndPrim

Vibration F -ratio 0.088 0.029 0.098 0.016 2.091 2.062 3.492 1.028 0.124
P -value 0.772 0.867 0.758 0.902 0.170 0.173 0.083 0.328 0.730
Effect size 0.006 0.002 0.007 0.001 0.130 0.128 0.200 0.068 0.009
Window F -ratio 2.116 0.208 2.547 0.072 1.793 1.101 0.623 1.069 1.928
P -value 0.139 0.813 0.096 0.931 0.185 0.347 0.491 0.357 0.164
Effect size 0.131 0.015 0.154 0.005 0.114 0.073 0.043 0.071 0.121
Target F -ratio 36.658 0.181 43.911 0.065 2.270 0.086 16.425 0.567 58.281
P -value 0.000 0.677 0.000 0.803 0.154 0.774 0.001 0.464 0.000
Effect size 0.724 0.013 0.758 0.005 0.140 0.006 0.540 0.039 0.806
Vibration*Window F -ratio 0.134 0.002 0.208 0.241 1.501 0.321 2.118 0.647 0.124
P -value 0.875 0.998 0.813 0.711 0.240 0.728 0.139 0.531 0.884
Effect size 0.009 0.000 0.015 0.017 0.097 0.022 0.131 0.044 0.009
Vibration*Target F -ratio 13.888 0.111 12.504 4.096 0.061 0.410 0.272 18.306 0.720
P -value 0.002 0.744 0.003 0.063 0.808 0.532 0.610 0.001 0.410
Effect size 0.498 0.008 0.472 0.226 0.004 0.028 0.019 0.567 0.049
Vision*Target F -ratio 1.360 0.778 0.747 0.072 0.502 0.859 1.808 0.630 5.363
P -value 0.273 0.430 0.483 0.820 0.611 0.434 0.183 0.540 0.024
Effect size 0.089 0.053 0.051 0.005 0.035 0.058 0.114 0.043 0.277
R. Goodman et al. / Multisensory Research 31 (2018) 455–480

Vibration*Window*Target F -ratio 0.493 0.359 1.386 0.244 0.684 0.134 0.188 0.127 0.715
P -value 0.616 0.702 0.267 0.785 0.513 0.875 0.830 0.881 0.469
Effect size 0.034 0.025 0.090 0.017 0.047 0.010 0.013 0.009 0.049

** Note: Effect sizes are reported as partial η2 values.

All significant effects and interactions are underlined.
471
472 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

significant and meaningful differences. The ANOVA’s also revealed one two-
way interaction for the vision × target comparisons for the EndPrim variable.
Post-hoc analyses revealed that in the jump condition, people corrected more
in the 1.0 m/s window than in the 1.4 m/s window.
3.2.4. Three-Way Interactions
The 2 vibration × 2 target × 3 window interaction analyses did not reach
significance for any of the dependent variables.
The a-priori contrasts examining performance in the 1.0 m/s window condi-
tions across levels of target jump provided evidence of a significant correction
to target jumps irrespective of vibration condition in terms of both EndPrim
(i.e., Vib: M = 307, SD = 21; NoVib: 308, SD = 21), and TaPV (i.e., Vib:
M = 186, SD = 26; NoVib: M = 180, SD = 27). That is, the EndPrim results
yielded the expected shorter movement amplitudes in the Jump as compared
to no-jump trials, in the 1.0 m/s window condition. Further, irrespective of the
application of tendon vibration was present, both MT and TaPV values were
shorter in the jump compared to the no-jump trials only for the 1.0 m/s window
condition (Table 4). Moreover, there were significant differences in the jump
condition between 1.0 m/s and 1.4 m/s windows, lending further evidence for
a 1.0 m/s ‘sweet spot’ (Tremblay et al., 2017).
Further, the contrast of the EndPrim between the jump and no-jump trials
in the 1.0 m/s window condition when tendon vibration was present yielded a
p-value of 0.0052 while the Bonferroni-corrected value was 0.0083. Analyses
of the three-way interactions for the MTs and the TaPVs yielded significantly
shorter movement durations and movement deceleration phase durations in the
jump compared to the no-jump trials across all window conditions in the ab-
sence of tendon vibration (Fig. 3C and D). In contrast, when tendon vibration
was present, the TaPVs were shorter in the jump compared to the no-jump
trials only for the 1.0 m/s window condition (Table 2).
3.3. Discussion
Overall, the target jumps elicited the anticipated shifts in endpoint distribu-
tions towards the briefly presented 27 cm target. That is, in both the presence
and the absence of vibration, participants significantly corrected their ongoing
trajectories in response to a target jump. To further quantify and evaluate the
magnitude of these responses to target jumps, the effect sizes associated with
the main effect of target were calculated and found to be comparable across
vibration conditions (Vib: Cohen’s D = 0.23; No Vib: Cohen’s D = 0.29).
As such, while the effects of target jumps were observed as expected when
tendon vibration was not present, participants also significantly corrected for
the target jumps in the vibration condition. These results represent evidence
that participants were able to effectively implement online control processes.
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 473

Figure 3. Results from the target jump experiment. Endpoint amplitude in the no-vibration (A)
and vibration conditions (B), reveal significant differences at 1.0 m/s. Time after peak velocity
revealed differences in all window conditions for the no-vibration (C), and at 1.0 m/s in the
vibration condition (D).

Additionally, the effects of the target jumps on the temporal characteristic also
yielded significant evidence of the contribution of online control processes in
response to target jumps.
As expected, the jump trials yielded shorter movement durations and shorter
limb deceleration phases durations than the no-jump trials. Further, the target
jumps failed to yield movement time differences whereas the limb decelera-
tion phase durations were shorter for the jump than the no-jump trials. As such,
474 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

because the later portions of the trajectory are thought to reflect online control
processes (e.g., Chua and Elliott, 1993; Woodworth, 1899), the results of the
target jump experiment yielded further evidence that visual feedback strongly
contributes to the online control of goal-directed movements. In contrast, little
evidence was observed for the contributions of proprioception to the online
control of action. Notably, three dependent variables yielded an interaction
between target jump and vibration condition. However, the post-hoc analyses
revealed that target jump yielded shorter movements times and deceleration
phases, both without and with tendon vibration. Such an explanation supports
the hypothesis that a greater weight is assigned to vision, as compared to pro-
prioception, to implement online limb–target regulation processes, which is
further supported when contrasting both experiments of the current study.

4. General Discussion
The current study aimed to directly assess the online utilization of vision
and proprioception during rapid upper-limb movements. Between-trial dual-
muscle tendon vibration was used to directly perturb proprioception, while
liquid-crystal goggles were used to directly perturb vision. Two experiments
were performed to assess the mechanisms of control, but more importantly, to
evaluate relative contributions of visual and proprioceptive feedback to online
control processes, especially the limb–target regulation processes. The results
of the current study were considered relative to Elliott and colleagues (2010)
multiple-processes model.
Known effects regarding online visual feedback utilization were replicated.
Results from the single-target experiment indicated that visual information
was likely used both for impulse regulation processes (i.e., variable error) and
limb–target regulation processes (i.e., Fisher2 ). Likewise, results from the tar-
get jump experiment revealed that with or without vibration, people were able
to adjust online for the unconscious target jump, even if visual information
was provided for only 20 ms. The current study also lent further support to the
multiple processes model, because online visual feedback could be utilized to
correct for the target jump (i.e., limb–target regulation: see also Tremblay et
al., 2017). In contrast, this study raises some questions about the use of propri-
oception for online limb–target regulation processes, which may contrast with
the extant literature because of the methodological approach employed.
As previously mentioned, the studies confirming the specific use of proprio-
ceptive feedback primarily either employed visual–proprioceptive mismatches
or online perturbations to the limb that can elicit reflexive activity. The current
study employed the after-effects of tendon vibration, which are known to al-
ter the resting firing rates of Ia spindles (see Ribot-Ciscar et al., 1998). Such
after-effects have been shown to decrease the reliability of the proprioceptive
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 475

signals in a limb-matching task (Goodman, 2015). With this between-trial vi-

bration method, the current study showed that the manipulation and associated
influence proprioception yielded clear effects on movement endpoint distribu-
tions, yet no evidence of effects on limb–target regulation processes.
The results from the single-target experiment showed that between-trial
tendon vibration yielded endpoint biases even when vision was available.
This result provides some evidence for visual–proprioceptive integration for
the control of voluntary reaching movements. In accordance to the maxi-
mum likelihood estimate model, both visual and proprioceptive sensory cues
should contribute to performance (Ernst and Bülthoff, 2004). As well, the
single-target experiment yielded evidence of trajectory biases (i.e., at peak
limb velocity and movement end) that were only significant when vision was
not available, which also suggests that participants were adequately altering
the weight assigned to each sensory modality based on its availability (e.g.,
Tremblay and Proteau, 1998). Such results in the no-vision conditions could
also be explained by motor planning differences. In contrast, the larger move-
ment amplitudes observed with than without vibration in the vision condition
were likely the result of online proprioceptive feedback utilization because the
mean amplitude differences at peak limb velocity was negligible (i.e., less than
0.1 mm). Limb displacements associated with voluntary actions may lead to
noisier proprioceptive inputs, which could decrease the reliance on these sig-
nals. As such, multisensory integration may still contribute to online control
processes but with smaller contributions from the proprioceptive system as
compared to the visual system, and that is, depending on whether vision is
available or not.
Importantly, the findings of the current study are consistent with studies
involving a deafferented individual wherein the individual retained the abil-
ity to correct ongoing reached to shifts in visual-target position, even in the
absence of vision of the limb (e.g., Bard et al., 1999; Sarlegna et al., 2006).
Similarly, it has been shown that when monkeys learned a motor task and
were subsequently deafferented (i.e., via bilateral dorsal rhizotomy), a certain
level of movement accuracy was retained following deafferentation (Polit and
Bizzi, 1978, 1979). Although the current study attempted to create noise in
a functional proprioceptive system, which is not fully comparable to deaf-
ferentation studies, all results support the core interpretation regarding the
relatively greater importance of vision as compared to proprioception for the
control of goal-directed action. Altogether, it is clear that tendon vibration did
have an impact on the voluntary reaches although one could question which
sub-processes were influenced by such proprioceptive perturbation. It is hoped
that future investigations will elucidate the relative weighing of vision and pro-
prioception for the implementation of online control processes.
476 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

Tendon vibration was delivered before movement onset and one could thus
argue that such proprioceptive perturbations affected planning mechanisms.
Indeed, proprioceptive inputs were admittedly altered during both the plan-
ning and execution of the movement (Ribot-Ciscar et al., 1998). One challenge
to this motor planning explanation were that the times taken to reach peak
velocity and the actual peak velocity did not significantly differ between vibra-
tion conditions, which are variables typically associated with motor planning
mechanisms (e.g., Chua and Elliott, 1993). In contrast, some of the effects of
tendon vibration on the temporal characteristics of the reaching movements
were observed in the deceleration phase.
In the target jump experiment, tendon vibration did hamper the influence of
the target jump on the deceleration phase durations (i.e., TaPV). Because the
deceleration phase duration has been associated with online control processes
(e.g., Chua and Elliott, 1993), the current study thus provided some evidence
that proprioception contributes to online trajectory amendments. One possi-
ble explanation for such effects is based on the expected decreased sensitivity
of most Ia afferent fibers, following tendon vibration (re.: Ribot-Ciscar et al.,
1998). Such decreased sensitivity could have yielded proprioceptive underesti-
mations of the displacement and rate of displacement of the limb. So, if tendon
vibration yielded the sensation of a smaller displacement or rate of displace-
ment of the finger than the one gathered by the visual input, an appropriate
online correction could be to slow down the movement and thus reduce the
deceleration phase duration. Although the main effects of tendon vibration did
not yield shorter deceleration phase durations per se, the effects sizes asso-
ciated with the target jump for the deceleration phase were starkly different
between the vibration conditions (Vib: Cohen’s D = 0.36; No Vib: Cohen’s
D = 0.80). Although more work is needed to elucidate the contribution of
proprioception to the online control of voluntary action, it remains clear that
tendon vibration and the associated direct proprioceptive perturbation failed to
yield evidence of the use of proprioception for online limb–target regulation
processes (cf. Elliott et al., 2010).
Tendon vibration did not significantly influence the main variable asso-
ciated with limb–target regulation processes (i.e., Fisher2 ). Moreover, target
jumps presented well after movement onset led to trajectory amendments re-
gardless of whether proprioception was perturbed or not. As a result, there was
no significant evidence that proprioception was utilized to implement the in-
duced online limb–target regulation processes (cf. Elliott et al., 2010), elicited
via target jumps. The relative contributions of vision and proprioceptive to the
control of action has been studied and it is evident that both modalities are im-
portant in some capacity (Bagesteiro et al., 2006; Grierson and Elliott, 2008,
2009; Khan and Franks, 2003; Rossetti et al., 1995). Some have suggested that
while vision is important for action planning, while proprioception contributes
 R. Goodman et al. / Multisensory Research 31 (2018) 455–480 477

largely to the control of action, later in the movement (e.g., Redon et al., 1991).
However, the empirical evidence employed to support the proposal that both
vision and proprioception are important for limb–target regulation processes
have either indirectly manipulated proprioceptive inputs or the direct proprio-
ceptive manipulations employed likely induced reflexive muscle activity that
should not be attributed to online sensorimotor control processes. Although
the direct proprioceptive perturbation methodology employed also presents
its own shortcomings, the current study provided compelling evidence for the
presence of online limb–target regulation processes (e.g., correction for the tar-
get jump) and that such processes took place even when employing a method
known to perturb the proprioceptive inputs. Because the proprioceptive per-
turbation elicited endpoint amplitude differences that could be explained by
online control mechanisms, at least in the vision conditions, it is possible that
the multiple processes model of online control (Elliott et al., 2010) needs to
be revised. Specifically, the contribution of proprioception to online control
mechanisms could be solely significant for impulse regulation processes but
not for limb–target regulation processes.
Although the findings of the current study are novel and compelling, future
experiments have been planned to corroborate the current findings. Firstly,
a target jump paradigm with a target shifting further along the trajectory (i.e.,
33 cm target) could be useful to confirm that tendon vibration and the associ-
ated proprioceptive disruption does not significantly interference with limb–
target regulation processes. Because the tendon vibration is thought to yield
underestimations of the displacement and rate of displacement of the limb, a
target jump requiring a limb re-acceleration will require a longer deceleration
phase period, which should hypothetically disambiguate the effects of spatial
endpoint vs. temporal characteristics. Additionally, in order to further inves-
tigate proprioceptive control, a paradigm including proprioceptive target may
also help provide evidence to the multiple-processes model because people
should plan and execute movement differently when controlling movements
towards a proprioceptive vs. a visual target (e.g., Sober and Sabes, 2003).
Both scenarios are relevant to activities of daily living. Finally, a paradigm em-
ploying a sham proprioceptive perturbation may help to persuade arguments
against conscious planning and control alterations for something that may be
changing feedback.
The importance of online sensory feedback availability is relevant to volun-
tary movements lasting at least 200 ms and involving accuracy requirements
corresponding to index of difficulty (ID: see Fitts, 1954) of 3.58 bits or more
(Wallace and Newell, 1983). Such a task difficulty corresponds to reaching
to a target of 2.5 cm in diameter or less, located 15 cm away from the start
position, and that is in at least 200 ms. Therefore, the current study and the
associated literature can be deemed relevant and ecologically valid, consider-
478 R. Goodman et al. / Multisensory Research 31 (2018) 455–480

ing that a significant proportion of activities of daily living involve movement

characteristics with longer durations and amplitudes combined with smaller
target sizes.

5. Conclusion
Overall, the current study provides further evidence for the importance of
vision for online control mechanisms (e.g., Elliott et al., 2010) and for the
importance of proprioceptive feedback for the production of accurate volun-
tary actions (e.g., Rossetti et al., 1995). As well, the current study provided
evidence that proprioceptive information can be utilized to implement online
trajectory amendments, even when vision was available. However, despite
clear evidence for the implementation of online limb–target regulation pro-
cesses (e.g., Tremblay et al., 2017), the current study yielded evidence of the
use of proprioception for online control processes but failed to yield signifi-
cant influence of the use of proprioception to implement limb–target regulation
processes (cf. Elliott et al., 2010).

Acknowledgements
This research was supported in part by grants from the Natural Sciences and
Engineering Research Council of Canada (NSERC), Canadian Foundation for
Innovation (CFI), Ontario Research Fund (ORF), Ontario Graduate Scholar-
ship (OGS), and the University of Toronto Graduate Student Scholarship.

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