What is a number? The number sense hypothesis suggests that numerosity is "a primary visual property" like color, contrast, or orientation. However, exactly what attribute of a stimulus is the primary visual property and determines numbers in the number sense? To verify the invariant nature of numerosity perception, we manipulated the numbers of items connected/enclosed in arbitrary and irregular forms while controlling for low-level features (e.g., orientation, color, and size). Subjects performed discrimination, estimation, and equality judgment tasks in a wide range of presentation durations and across small and large numbers. Results consistently show that connecting/ enclosing items led to robust numerosity underestimation, with the extent of underestimation increasing monotonically with the number of connected/enclosed items. In contrast, grouping based on color similarity had no effect on numerosity judgment. We propose that numbers or the primitive units counted in numerosity perception are influenced by topological invariants, such as connectivity and the inside/outside relationship. Beyond the behavioral measures, neural tuning curves to numerosity in the intraparietal sulcus were obtained using functional MRI adaptation, and the tuning curves showed that numbers represented in the intraparietal sulcus were strongly influenced by topology.W hat is a number? The answer to this age-old and fundamental question of philosophy has increasingly benefited from recent scientific investigation using psychology and neuroscience. The number sense hypothesis (1, 2) suggests that a number is "a basic property of the environment" (3) and particularly, because of its remarkable adaptation effect, "a primary visual property" (2), like color, contrast, or orientation (2-8). However, exactly what attribute in the environment is the primary property and determines numbers in the number sense, or more concretely, what is counted in numerosity perception? Consider the invariant nature of numerosity perception. It is self-evident that numerosity is invariant to specific features (e.g., orientation, size, shape, and color) of individual items to be counted. In other words, the primitive units to be counted must be invariant with variation in form dimensions and other visual features (2, 3, 7, 9-11). Then, the critical question becomes how to define precisely such abstract and invariant attributes.
ResultsGeneralizing Connection to a Topological Invariant: Connectivity. We designed arbitrary and irregular shapes of connecting line segments (Fig. 1A, Upper) to test the invariant effect of connection on numerosity judgement independent of the concrete forms or manners of connection (12, 13). Three conditions of connection were constructed: zero, one, and two connected pairs of dots in the test patterns (Fig. 1A). A typical numerosity discrimination task was adopted, in which two visual patterns of dots were briefly presented on opposite sides of fixation (one serving as a reference and the other one serving as a test), and subj...