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He injected a new naturalism into painting. Plants in the first “Virgin of the Rocks” are depicted in almost microscopic detail. Portrait subjects had traditionally been depicted in profile, but Leonardo turned them so they look toward the viewer. He strove to capture his subjects’ thoughts and emotions on their faces. “The Lady With an Ermine,” a depiction of his patron’s 16-year-old mistress, has a teasing, inward expression, while “The Belle Ferronniere” _ based on the duke’s wife _ looks boldly and challengingly at the viewer.

“These are much more than simple likenesses of people,” Syson said.

In “The Madonna Litta,” on loan from the Hermitage in St. Petersburg, the Virgin’s idealized beauty is complemented by the breast-feeding baby’s cheeky gaze toward the viewer.

Later, Leonardo began to seek perfection, seeing himself as “somebody whose talent is, in a sense, fed by God himself” and striving to depict a kind of absolute beauty.

“I think 100 years later he would have got into trouble with the Inquisition,” because of his unorthodox ideas on art and religion, Syson said.

“He believed painting could represent not just everything that was visible, but all that was invisible in the universe.”