Harambe, the 17-year-old Western lowland gorilla shot dead at the Cincinnati Zoo late last month after a 3-year-old boy fell into his enclosure, may be physically gone, his tissues harvested for research and his sperm extracted to help diversify the captive breeding gene pool.

Yet the 440-pound silverback leaves another metaphorical gorilla in the room, raising questions that extend far beyond the particulars of the case, including whether the zoo or the boy’s mother were more to blame for Harambe’s death.

For primatologists and conservationists who devote their lives to studying the great apes and to doing what they can to help protect the rapidly vanishing populations of the primates in the wild, a linked set of ethical and practical dilemmas looms almost unbearably large.

As research continues to reveal the breadth of our genetic, emotional and cognitive kinship with the world’s four great apes — gorillas, chimpanzees, bonobos and orangutans — many primatologists admit to feeling frankly uncomfortable at the sight of a captive ape on display, no matter how luxe or “natural” the zoo exhibit may be.