This study provides firm support for the hypothesis that the silencing effect reported by Suchow and Alvarez (
2011) resulted from a combination of motion integration and crowding. The effect clearly depends on motion, with a well-defined threshold. The motion threshold was 0.2 rotations per second (average linear speed around 10°/s) under the conditions used by Suchow and Alvarez. The motion thresholds varied with dot spacing, being much lower for closely spaced dots than broadly spaced, consistent with crowding. Thresholds for dot spacing were independent of dot size but depended on the spacing of their centers, as occurs with crowding (Bouma,
1970; Levi & Carney,
2009; Tripathy & Cavanagh,
2002). The Bouma constants for the silencing effects ranged between 0.4 and 0.6, well within the normal range reported in the literature (Pelli & Tillman,
2008; Whitney & Levi,
2011).
There is very good evidence that neural mechanisms integrate motion signals over large and complex trajectories. Neurons in the dorsal portion of area medial superior temporal area that respond specifically to various types of flow motion, including rotation, have very large receptive fields, often extending over more than 90° (Duffy & Wurtz,
1991; Tanaka & Saito,
1989; Tanaka, Fukada, & Saito,
1989). Psychophysical studies also show compulsory integration of flow motion (Morrone, Burr, & Vaina,
1995), again over very large areas (Burr, Morrone, & Vaina,
1998). It seems plausible that this integration process subsumes all dynamic signals within the area, including those associated with the changes in stimulus size.
There are several examples of global motion obscuring local motion, or transients. A field of coherently oriented dipole dot-pairs appearing continuously in random positions gives a strong impression of circular motion (Ross, Badcock, & Hayes,
2000;
Supplementary Movie S6). So strong and smooth is the sense of motion that it is hard to believe that the dipoles in fact are appearing and disappearing at random. There is little sense of the dynamics of the individual dipoles; this is all consumed by the global sense of circular motion. A more mundane example is the “limited-lifetime” stimuli that most of us use routinely for motion studies. We are typically totally unaware that the dots continually drop out and reappear at random positions; yet if the display is stopped, the continuous flicker becomes obvious (
Supplementary Movie S7). Saiki and Holcombe (
2012) have provided an even more dramatic example of our inability to detect color changes in individual dots. Clouds of dots, half red and half green, rotating in an apparent sphere can all switch color, and the switch goes unnoticed, provided the summary statistics (such as red/green ratio) remain unchanged. This clearly shows that the visual system does not monitor independently the behavior of every single dot in multidot displays.
However, it is also clear that motion integration is not the entire solution. It is easy to track the changes in size (or other dimensions) in isolated or sparse arrays of dots in motion. One of the clearest examples is Boi, Ogmen, Krummenacher, Otto, and Herzog's (
2009) demonstration of “nonretinotopic” processing, being able to process rotational motion in apparently moving objects, correctly with respect to the moving reference plane (Pooresmaeili, Cicchini, Morrone, & Burr,
2012). This is a more taxing problem than simply detecting that size or color has changed. Motion silencing also fails when the displays are sparse (see
Supplementary Movie S5), leading to the suggestion that crowding may be involved. When the display is stationary, the dynamic change-signals of each element breaks through crowding, in the same way that temporal transients are known to cause ‘pop-out,' reaching awareness without active attention; but if the signal-changes are subsumed by global motion mechanisms, then we would have to be able to individuate the dots to be able to detect changes in each.
Crowding refers to the fact recognition of target in the periphery is difficult when surrounded by other stimuli. The critical parameter for crowding is not the distance between object contours but the distance between their centers; and this
critical spacing is proportional to eccentricity (Bouma,
1970), approximately half the target eccentricity. This relationship is so solid, it is generally termed Bouma's law, although the precise value varies depending on stimulus characteristics and task requirements (Whitney & Levi,
2011). An important consequence of this law is that the size of the interference zone is independent of target (Bouma,
1970; Levi & Carney,
2009; Pelli & Tillman,
2008; Tripathy & Cavanagh,
2002; but see also Manassi, Sayim, & Herzog,
2012). The evidence presented here strongly supports the suggestion that motion silencing occurs only in crowded displays. Speed thresholds depended strongly on spacing of the stimulus rings. The
critical spacing for motion silencing was about half the eccentricity of the rings, for two eccentricities (3.5° and 7°), well within the rage of parameters normally reported. Importantly, the critical spacing did not vary with a halving of object size, a signature of crowding.
It is difficult to relate the results of the first experiment, with 100 dots randomly positioned within a 5° × 8° annulus, with the second series, where the dots were uniformly positioned around three equispaced rings. The first experiment (with randomly positioned dots) yielded velocity thresholds around 0.2 rotations per second.
Figure 4 suggests that with regularly spaced dots, the threshold spacing should be around 3°. The average spacing between the random dots of Experiment 1 was 1°: but as they were randomly positioned, this varied, so that 15% had spacing greater than 3°. Perhaps the discrimination (under forced choice) was achieved by this small percentage of uncrowded dots?
Crowding occurs for moving objects, with a similar dependence of eccentricity and size of critical zones as those observed for stationary objects (Bex & Dakin,
2005; Bex, Dakin, & Simmers,
2003). Crowding does not increase with speed, consistent with the fact that we find Bouma constants similar to those reported for stationary targets. Interestingly, there is also evidence that crowding occur after motion has been processed, as it is the perceived rather than actual physical position determines crowding in stimuli moving within a stationary window (Maus, Fischer, & Whitney,
2011).
In summary we believe that two mechanisms are involved in motion silencing: integration of global motion to absorb the dynamic change signals of the individual dots, and crowding mechanisms to prevent perceptual isolation of individual dots, allowing their individual changes to be monitored.