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SAN FRANCISCO — Koko the gorilla, whose remarkable sign-language ability and motherly attachment to pet cats helped change the world’s views about the intelligence of animals and their capacity for empathy, has died at 46.

Koko was taught sign language from an early age as a scientific test subject and eventually learned more than 1,000 words, a vocabulary similar to that of a human toddler.

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She became a celebrity who played with the likes of William Shatner, Sting, Leonardo DiCaprio, Robin Williams and Mr. Rogers. At her home preserve, where she was treated like a queen, she ran around with Williams’ eyeglasses and unzipped Roger’s famous cardigan sweater.

In so doing, Koko showed the American public that a giant ape didn’t have to be scary but wanted to be tickled and hugged.

The Gorilla Foundation said the 280-pound western lowland gorilla died in her sleep at the foundation’s preserve in California’s Santa Cruz mountains on Tuesday.