University of Washington Ray Hilborn, a professor at the University of Washington School of Aquatic and Fishery Sciences, is under fire from Greenpeace over his alleged failure to disclose the corporate interests backing his research.

A prominent fisheries scientist who has challenged the need for marine conservation is under investigation by his university after the environmental group Greenpeace accused him of failing to reveal seafood industry funding.

Ray Hilborn, a professor at the University of Washington’s School of Aquatic and Fishery Sciences, received at least $3.56 million over 12 years from fishing and seafood industry groups for research and private consulting, according to documents cited by Greenpeace. The group submitted a complaint to the University of Washington on Wednesday, asking it to investigate whether Hilborn adequately disclosed industry backing and whether the funding is a conflict of interest.

Greenpeace said Hilborn's research shows he is a "denier of overfishing."

"The seafood industry has given millions of dollars to Ray Hilborn in an attempt to undermine the broad scientific consensus that poor fisheries management has resulted in depleted fish populations and damaged ecosystems," Greenpeace USA Oceans campaign director John Hocevar said in a statement. He said readers of Hilborn's work "should at the very least know that corporate interests are underwriting his commentary."

University of Washington spokesman Norman G. Arkans said the school takes the accusations "very seriously" and would investigate whether Hilborn had breached policies. (See UPDATE below.)

Greenpeace files complaint on industry funding for UW fishery professor: https://t.co/VALRXaxitp #OverfishingDenier pic.twitter.com/zrOKGEXPdg — Greenpeace USA (@greenpeaceusa) May 12, 2016

Hilborn, 68, whose research has earned multiple awards -- including the 2016 International Fisheries Science Prize -- told HuffPost he "absolutely rejects" the criticism. He said Greenpeace targeted him because his work doesn't fit the narrative that all marine ecosystems need conservation to prevent overfishing.

"Greenpeace is unable to attack the science I and my collaborators do; science that threatens their repeated assertions that overfishing is universal and that the oceans are being emptied," Hilborn wrote on his blog.

Ron Wurzer via Getty Images A study on salmon populations is one of the papers that Greenpeace has taken issue with.

The $3.56 million that Hilborn received from industry groups from 2003 to 2015 is just 22 percent of all the funding the scientist brought to the university, according to the Seattle Times.

Hilborn's funding sources included companies like Trident Seafoods and Peter Pan Seafoods, as well as the industry group National Fisheries Institute. In some of his scientific papers, Hilborn failed to disclose industry funding, Greenpeace alleged.

Hilborn acknowledged receiving the $3.56 million in funding, but said the figure includes money from Alaskan community groups that depend on fishing.

Pascal Rossignol / Reuters Greenpeace bills trawling as a damaging fishing practice that is depleting fish stores.

Greenpeace cited is a 2006 paper published in the Canadian Journal of Fisheries and Aquatic Sciences on the orange roughy in New Zealand, which said the fish population needed no changes in fishery management. Hilborn failed to disclose $58,000 in funding from the New Zealand Seafood Industry Council around the time of the study, Greenpeace said.

Greenpeace also pointed to a 2007 article in the journal Ecosystems, which said current fishing management was working in some places, but not in others. Hilborn should have disclosed in that paper that he was receiving funding from Trident, Peter Pan, the New Zealand Seafood Industry Council and the Bering Sea Fishermen's Association, Greenpeace said.

Hilborn argued in his blog post that if he had to disclose every group that contributed to his research on every paper, the list would be "as long as some of the papers." He said he acknowledges funders of research that is the main subject of each paper.

Industry funding, he said, helps support student and staff salaries, and pays for field expenses.

Hilborn said more of his funding has come from environmental foundations than from industry, including the Society for Conservation Biology, the Natural Resources Defense Council and the Environmental Defense Fund.

Greenpeace seems to think fishing industry should not fund research ... here is how I fund my research program. pic.twitter.com/0gmdiRCQju — Ray Hilborn (@hilbornr) May 12, 2016

"I am a vocal advocate for where marine conservation has worked, and identifying where it is not working," Hilborn said.

"In fact, it is in the financial interest of fishing communities and industries to find solutions that are sustainable and provide for healthy stocks into the future," he wrote on his blog. "And funding from these groups should be considered part of an inclusive, transparent and honest research process."

"The major threat to sustainable jobs, food, recreational opportunity and revenue from U.S. marine fisheries is no longer overfishing, but underfishing," Hilborn wrote in a 2013 testimony submitted to Congress.

@danagoodyear Note abundance of U.S. east coast fish stocks increasing, not declining pic.twitter.com/wH9XjaHymZ — Ray Hilborn (@hilbornr) November 3, 2015

Here is the trend in Atlantic bluefin tuna abundance showing rapid recovery after effective management. @the_klute pic.twitter.com/w2LLvLb14d — Ray Hilborn (@hilbornr) October 7, 2015

Industry funding of scientific research has stirred significant debate in recent years. A 2015 New York Times report exposed scientists backed by Coke for downplaying the link between soda and obesity.

But as public funding for science declines, researchers are turning to industry and other groups for financial support. Some scientific journals, including some that have published Hilborn's papers, have created or toughened policies for disclosure of that funding.

UPDATE: June 8 -- The University of Washington and the journal Proceedings of the National Academy of Sciences both concluded that Hilborn properly disclosed his funding sources. Additionally, the university said he didn't violate its policies on conflicts of interest or outside work.

Handout . / Reuters Sockeye salmon in Bristol Bay, Alaska.