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Question 5·Hard·Standard English Conventions

To investigate whether early exposure to artificial light shapes later navigation, entomologist Priya Nanduri reared two groups of hawk moths, one beneath amber streetlamps and the other in darkness. During release trials, the moths that had developed beneath the lamps, unlike those raised in natural ______ toward a bright ultraviolet source even when moonlight was visible.

Which choice completes the text so that it conforms to the conventions of Standard English?