Opposed to this flat, unrelieved style was the work of a follower, Agatharchos of Samos (fl. end of fifth century B.C.). He was a scene-painter, and by the necessities of his craft was led toward nature. Stage effect required a study of perspective, variation of light, and a knowledge of the laws of optics. The slight outline drawing of his predecessor was probably superseded by effective masses to create illusion. This was a distinct advance toward nature. Apollodorus (fl. end of fifth century B.C.) applied the principles of Agatharchos to figures. According to Plutarch, he was the first to discover variation in the shade of colors, and, according to Pliny, the first master to paint objects as they appeared in nature. He had the title of _skiagraphos_ (shadow-painter), and possibly gave a semi-natural background with perspective. This was an improvement, but not a perfection. It is not likely that the backgrounds were other than conventional settings for the figure. Even these were not at once accepted by the painters of the period, but were turned to profit in the hands of the followers. After the Peloponnesian Wars the art of painting seems to have flourished elsewhere than in Athens, owing to the Athenian loss of supremacy. Other schools sprang up in various districts, and one to call for considerable mention by the ancient writers was the IONIAN SCHOOL, which in reality had existed from the sixth century. The painters of this school advanced upon the work of Apollodorus as regards realistic effect. Zeuxis, whose fame was at its height during the Peloponnesian Wars, seems to have regarded art as a matter of illusion, if one may judge by the stories told of his work. The tale of his painting a bunch of grapes so like reality that the birds came to peck at them proves either that the painter's motive was deception, or that the narrator of the tale picked out the deceptive part of his picture for admiration. He painted many subjects, like Helen, Penelope, and many _genre_ pieces on panel. Quintilian says he originated light-and-shade, an achievement credited by Plutarch to Apollodorus. It is probable that he advanced light-and-shade. In illusion he seems to have been outdone by a rival, Parrhasios of Ephesus. Zeuxis deceived the birds with painted grapes, but Parrhasios deceived Zeuxis with a painted curtain. There must have been knowledge of color, modelling, and relief to have produced such an illusion, but the aim was petty and unworthy of the skill. There was evidently an advance technically, but some decline in the true spirit of art. Parrhasios finally suffered defeat at the hands of Timanthes of Kythnos, by a Contest between Ajax and Ulysses for the Arms of Achilles. Timanthes's famous work was the Sacrifice of Iphigenia, of which there is a supposed Pompeian copy.