Direction selective neurons in macaque primary visual cortex are narrowly tuned for orientation, and are thus afflicted by the aperture problem. At the next stage of motion processing, in the middle temporal (MT) area, some cells appear to solve this problem, responding to the pattern motion direction of plaids. Models have been proposed to account for this computation, but they do not replicate the diversity of responses observed in MT. We recorded from 386 cells in area MT of two male macaques, while presenting a wide range of random-line stimuli and their compositions into noise plaids. As we broadened the range of stimuli used to probe the cells, yielding ever more challenging conditions for extracting pattern motion, the diversity of the responses observed increased, and the fraction of cells that faithfully encoded pattern motion direction shrank. However, we show here that a pattern motion signal is present at the population level. We identified four mechanisms, one never proposed before, that toget...