The brilliance of Leonardo da Vinci has been variously attributed to his meticulous attention to detail, his vast knowledge of art, science and optics, and the fact he painted with his left hand.
But new research has shown that an intermittent flaw in his eyesight may also have helped him perfectly capture three-dimensional form.
Professor Christopher Tyler, a visual neuroscientist from City, University of London, studied two sculptures, two oil paintings and two drawings that featured the artist - either in the form of a self-portrait or as the posing model.
They included Leonardo's Salvator Mundi, Vitruvian Man and the bronze statue of David by Andrea del Verrocchio, his teacher.
In each case, the alignments of the pupils in the eye apertures diverged by up to 13.2 degrees, suggesting Leonardo had a condition known as exotropia, where one eye looks directly at an object while the other gazes off to one side.