In an undated photo from Penn State, the biologist Victoria Braithwaite in 2012. Braithwaite, who studied the behavior of fish, including the effect of environmental stimulation that they experience in tanks, died on Sept. 30, 2019, at her home in Boalsburg, Pa. She was 52. (Penn State University via The New York Times) -- NO SALES; FOR EDITORIAL USE ONLY WITH NYT STORY OBIT BRAITHWAITE BY CORNELIA DEAN FOR NOV. 1, 2019. ALL OTHER USE PROHIBITED. --

In his 2005 novel, “Saturday,” the acclaimed writer Ian McEwan describes what his protagonist, Henry Perowne, sees when he visits a London seafood market to buy ingredients for a fish stew: crates of crabs and lobsters, still moving, and marble slabs arrayed with “bloodless white flesh, and eviscerated silver forms.”

While there, he ponders reports of recent scientific research demonstrating that fish can feel pain. It is a good thing that sea creatures “have no voice,” Perowne thinks. “Otherwise there’d be howling from those crates.”

Perowne’s thoughts may have been fictional, but the reports were real. They were written by Victoria Braithwaite, then a senior lecturer at the University of Edinburgh, who died on Sept. 30 at her home in Boalsburg, Pennsylvania. She was 52.

Her death, from pancreatic cancer, was confirmed by a spokeswoman for Penn State University, where she had been a professor of fisheries and biology since 2007.

In two papers, one published in Proceedings of the Royal Society and the other in The Journal of Pain, Braithwaite and her colleagues attracted public attention with their reports that fish had neural cells called nociceptors, analogous to cells in humans that transmit pain; that these cells react to injuries; and that fish exposed to unpleasant stimuli — vinegar added to their water, for example — acted differently from fish not similarly exposed.

In the book “Do Fish Feel Pain?” (2010), she discussed these findings and argued that fish should be accorded the same protections commonly applied to birds and mammals, such as humane slaughter. Animal rights activists praised her work and publicized it on social media.

But many neuroscientists and fish biologists responded to Braithwaite’s findings with skepticism, or even disdain.

In 2013, a team of seven scientists from Europe, Canada, Australia and the United States assessed research on pain in fish and concluded that fish do not have the neurophysiological capacity for conscious feelings of pain. For example, they said in the journal Fish and Fisheries, the fish brain lacks a neocortex, which in mammals is involved in brain functions like sensory perception. Fish may respond to unpleasant stimuli, they concluded, but they would feel pain only if they had consciousness — which they said Braithwaite’s work did not demonstrate.

One of those scientists, Robert Arlinghaus, an expert on fish and fisheries with appointments at Humboldt University and the Leibniz Institute of Freshwater Ecology and Inland Fisheries, both in Berlin, said in an email that Braithwaite was “an outstanding fish behavior scientist,” but that “her work on pain I do not find convincing.”

Still, he and the other researchers noted in their report that people are morally responsible for justifying their use of fish, and for minimizing stress and other harm to the creatures.

Braithwaite had been appointed to head the Leibniz Institute when she received her cancer diagnosis.

Neuroscientists note that much is unknown about brain function, whether mammalian, avian, reptilian or piscine. Pain, too, remains mysterious, and highly subjective.

Carl Saab, a neuroscientist at Brown University, said that, among other things, researchers are still seeking to identify measurable substances, or biomarkers, that are indications of pain. “What do we mean when we say a rat or a human has pain,” he asked, “and how to measure that pain objectively?”

Victoria Anne Braithwaite was born on July 19, 1967, in Halifax, Yorkshire, England, the sixth of seven children of Alan and June (Pickles) Braithwaite. Her father was a textile executive, her mother a magistrate.

She studied zoology as both an undergraduate and graduate student at Oxford, where she wrote a doctoral dissertation on navigation by pigeons. She took a postdoctoral appointment at the University of Glasgow, where she began her work on cognition in fish.

She was on the faculty of the University of Edinburgh from 1995 until joining Penn State in 2007. She was also a visiting professor of biology at the University of Bergen, Norway, for several years.

In 1992 she married Andrew Read, a researcher on infectious diseases, who also joined the faculty of Penn State; they divorced in 2015. She is survived by two sons, James and Matthew, and two grandchildren.

In recent years, Braithwaite had begun researching the effect of environmental stimulation that fish experience in tanks. She concluded that they did better when their surroundings varied and contained things like plants.

In a 2014 television interview for the BBC program “Newsnight,” Braithwaite discussed the treatment of wild-caught and farmed fish with Bertie Armstrong, who was the chief executive of the Scottish Fishermen’s Federation at the time.

Braithwaite said that she felt worse for farmed fish, which she said were often raised in overcrowded, disease-ridden pens, and that regulators should insist on more humane treatment for them. “We know how to more humanely kill fish on farms,” she said.

By contrast, she said, fish caught in the wild may have lived a full life. She said their suffering upon capture was relatively short — though she praised fishers in Denmark and Norway who she said pumped nets of fish like mackerel into saltwater tanks on their boats.

Armstrong did not address her assertions that fish feel pain. Instead, he noted that fish are an important source of protein for the world. “It is just not possible to catch that volume of fish and handle them individually,” he said.

Perhaps that is what Ian McEwan’s protagonist is thinking as he debates how much empathy to have for the creatures at the fish market.

“The trick, as always, the key to human success and domination, is to be selected in your mercies,” Perowne tells himself. He buys prawns, monkfish tails, and the bones and heads of two skates, and heads home.

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