Volker H. Franz

Grasping Visual Illusions: No Evidence for a Dissociation Between Perception and Action

Neuropsychological studies prompted the theory that the primate visual system might be organized into two parallel pathways, one for conscious perception and one for guiding action. Supporting evidence in healthy subjects seemed to come from a dissociation in visual illusions: In previous studies, the Ebbinghaus (or Titchener) illusion deceived perceptual judgments of size, but only marginally influenced the size estimates used in grasping. Contrary to those results, the findings from the present study show that there is no difference in the sizes of the perceptual and grasp illusions if the perceptual and grasping tasks are appropriately matched. We show that the differences found previously can be accounted for by a hitherto unknown, nonadditive effect in the illusion. We conclude that the illusion does not provide evidence for the existence of two distinct pathways for perception and action in the visual system.

Action does not resist visual illusions

Recent TICS articles discussed the psychophysical evidence in favor of Goodale and Milners action vs. perception hypothesis. Carey argued that most of the studies investigating the effects of visual illusions on grasping can be reconciled with the notion that the action system resists visual illusions. Bruno suggested a new interpretation of the action vs. perception hypothesis in order to incorporate most of the empirical findings. Here, I argue that action does not resist visual illusions. Even more, the effects on the motor system seem to be comparable to the effects on the perceptual system. This challenges the action vs. perception hypothesis in its current form.

Grasping visual illusions: Consistent data and no dissociation.

The finding that the Ebbinghaus/Titchener illusion deceives perception but not grasping is usually seen as strong evidence for Goodale and Milners (1992) notion of two parallel visual systems, one being conscious and deceived by the illusion (vision-for-perception) and the other being unconscious and not deceived (vision-for-action). However, this finding is controversial and led to studies with seemingly contradictory results. We argue that these results are not as contradictory as it might seem. Instead, studies consistently show similar effects of the illusion on grasping. The perceptual effects are strongly dependent on the specific perceptual measure employed. If, however, some methodological precautions are used, then these diverse perceptual results can be reconciled and point to a single internal size estimate that is used for perception and for grasping. This suggests that the Ebbinghaus illusion deceives a common representation of object size that is used by perception and action.

When is grasping affected by the Müller-Lyer illusion?: A quantitative review

Milner and Goodale (1995) [Milner, A. D., & Goodale, M. A. (1995). The visual brain in action. Oxford, UK: Oxford University Press] proposed a functional division of labor between vision-for-perception and vision-for-action. Their proposal is supported by neuropsychological, brain-imaging, and psychophysical evidence. However, it has remained controversial in the prediction that actions are not affected by visual illusions. Following up on a related review on pointing (see Bruno et al., 2008 [Bruno, N., Bernardis, P., & Gentilucci, M. (2008). Visually guided pointing, the Müller-Lyer illusion, and the functional interpretation of the dorsal-ventral split: Conclusions from 33 independent studies. Neuroscience and Biobehavioral Reviews, 32(3), 423-437]), here we re-analyze 18 studies on grasping objects embedded in the Müller-Lyer (ML) illusion. We find that median percent effects across studies are indeed larger for perceptual than for grasping measures. However, almost all grasping effects are larger than zero and the two distributions show substantial overlap and variability. A fine-grained analysis reveals that critical roles in accounting for this variability are played by the informational basis for guiding the action, by the number of trials per condition of the experiment, and by the angle of the illusion fins. When all these factors are considered together, the data support a difference between grasping and perception only when online visual feedback is available during movement. Thus, unlike pointing, grasping studies of the Müller-Lyer (ML) illusion suggest that the perceptual and motor effects of the illusion differ only because of online, feedback-driven corrections, and do not appear to support independent spatial representations for vision-for-action and vision-for-perception.

Visual illusions, delayed grasping, and memory: No shift from dorsal to ventral control

We tested whether a delay between stimulus presentation and grasping leads to a shift from dorsal to ventral control of the movement, as suggested by the perception-action theory of Milner and Goodale (Milner, A.D., & Goodale, M.A. (1995). The visual brain in action. Oxford: Oxford University Press.). In this theory the dorsal cortical stream has a short memory, such that after a few seconds the dorsal information is decayed and the action is guided by the ventral stream. Accordingly, grasping should become responsive to certain visual illusions after a delay (because only the ventral stream is assumed to be deceived by these illusions). We used the Müller-Lyer illusion, the typical illusion in this area of research, and replicated the increase of the motor illusion after a delay. However, we found that this increase is not due to memory demands but to the availability of visual feedback during movement execution which leads to online corrections of the movement. Because such online corrections are to be expected if the movement is guided by one single representation of object size, we conclude that there is no evidence for a shift from dorsal to ventral control in delayed grasping of the Müller-Lyer illusion. We also performed the first empirical test of a critique Goodale (Goodale, M.A. (2006, October 27). Visual duplicity: Action without perception in the human visual system. The XIV. Kanizsa lecture, Triest, Italy.) raised against studies finding illusion effects in grasping: Goodale argued that these studies used methods that lead to unnatural grasping which is guided by the ventral stream. Therefore, these studies might never have measured the dorsal stream, but always the ventral stream. We found clear evidence against this conjecture.

Manual size estimation: a neuropsychological measure of perception?

Manual size estimation (participants indicate the size of an object with index finger and thumb) is often interpreted as a measure of perceptual size information in the visual system, in contrast to size information used by the motor system in visually guided grasping. Because manual estimation is a relatively new measure, I compared it to a more traditional perceptual measure (method of adjustment). Manual estimation showed larger effects of the Ebbinghaus (or Titchener) illusion than the traditional perceptual measure. This inconsistency can be resolved by taking into account that manual estimation is also unusually responsive to a physical variation of size. If we correct for the effect of physical size, manual estimation and the traditional perceptual measure show similar illusion effects. Most interestingly, the corrected illusion effects are also similar to the illusion effects found in grasping. This suggests that the same neuronal signals which generate the illusion in the traditional perceptual measure are also responsible for the effects of the illusion on manual estimation and on grasping.

Grasp effects of the Ebbinghaus illusion: obstacle avoidance is not the explanation.

The perception-versus-action hypothesis states that visual information is processed in two different streams, one for visual awareness (or perception) and one for motor performance. Previous reports that the Ebbinghaus illusion deceives perception but not grasping seemed to indicate that this dichotomy between perception and action was fundamental enough to be reflected in the overt behavior of non-neurological, healthy humans. Contrary to this view we show that the Ebbinghaus illusion affects grasping to the same extent as perception. We also show that the grasp effects cannot be accounted for by non-perceptual obstacle avoidance mechanisms as has recently been suggested. Instead, even subtle variations of the Ebbinghaus illusion affect grasping in the same way as they affect perception. Our results suggest that the same signals are responsible for the perceptual effects and for the motor effects of the Ebbinghaus illusion. This casts doubt on one line of evidence, which used to strongly favor the perception-versus-action hypothesis.

Effects of auditory stimulus intensity on response force in simple, go/no-go, and choice RT tasks

In four experiments, increasing the intensities of both relevant and irrelevant auditory stimuli was found to increase response force (RF) in simple, go/no-go, and choice reaction time (RT) tasks. These results raise problems for models that localize the effects of auditory intensity on purely perceptual processes, indicating instead that intensity also affects motor output processes under many circumstances. In Experiment 1, simple RT, go/no-go, and choice RT tasks were compared, using the same stimuli for all tasks. Auditory stimulus intensity affected both RT and RF, and these effects were not modulated by task. In Experiments 2–4, an irrelevant auditory accessory stimulus accompanied a relevant visual stimulus, and the go/no-go and choice tasks were used. The intensity of the irrelevant auditory accessory stimulus was found to affect RT and RF, although the sizes of these effects depended somewhat on the temporal predictability of the accessory stimulus.

Standard errors and confidence intervals in within-subjects designs: Generalizing Loftus and Masson (1994) and avoiding the biases of alternative accounts

Repeated measures designs are common in experimental psychology. Because of the correlational structure in these designs, the calculation and interpretation of confidence intervals is nontrivial. One solution was provided by Loftus and Masson (Psychonomic Bulletin & Review 1:476–490, 1994). This solution, although widely adopted, has the limitation of implying same-size confidence intervals for all factor levels, and therefore does not allow for the assessment of variance homogeneity assumptions (i.e., the circularity assumption, which is crucial for the repeated measures ANOVA). This limitation and the method’s perceived complexity have sometimes led scientists to use a simplified variant, based on a per-subject normalization of the data (Bakeman & McArthur, Behavior Research Methods, Instruments, & Computers 28:584–589, 1996; Cousineau, Tutorials in Quantitative Methods for Psychology 1:42–45, 2005; Morey, Tutorials in Quantitative Methods for Psychology 4:61–64, 2008; Morrison & Weaver, Behavior Research Methods, Instruments, & Computers 27:52–56, 1995). We show that this normalization method leads to biased results and is uninformative with regard to circularity. Instead, we provide a simple, intuitive generalization of the Loftus and Masson method that allows for assessment of the circularity assumption.

Memory mechanisms in grasping

The availability of visual information influences the execution of goal-directed movements. This is very prominent in memory conditions, where a delay is introduced between stimulus presentation and execution of the movement. The corresponding effects could be due to a decay of the visual information or to different processing mechanisms used for movements directed at visible (dorsal stream) and remembered (ventral stream) objects as proposed by the two visual systems hypothesis. In three experiments, the authors investigated grasping under full vision and three different delay conditions with increasing memory demands. Results indicate that the visuomotor information used for grasping decays rapidly. No evidence was found for qualitative changes in movement kinematics and the use of different representations for visually guided and memory guided movements. Findings rather suggest that delayed grasping is similar to grasping directed to larger objects under full vision. Therefore, the authors propose that grasping after a delay is guided by classic memory mechanisms and that this is reflected in an increasing maximum grip aperture in grasping.