Edinburgh Zoo's male giant panda Yang Guang has been castrated following a tumor scare, dealing a huge blow to the only panda breeding program in the United Kingdom.

Representatives from the zoo said that Yang Guang - which means sunshine in English - is recovering well from surgery to remove both testicles due to the presence of tumors. The operation took place in mid-November after group consultation by British and Chinese giant panda specialists.

Darren McGarry, head of living collections at Edinburgh Zoo, said the zoo will now hold talks with its Chinese partners to discuss "future breeding possibilities".

The panda and his mate Tian Tian - which means sweetie in English - were brought to Edinburgh from Beijing Zoo in 2011. They are the only giant pandas in the UK.

"Testicular tumors have previously been reported in giant pandas," said McGarry. "The tumors we removed developed recently, so they have not been a factor in Yang Guang and Tian Tian not having had a cub. We will discuss next steps with our partners in China over coming weeks."

Yang Guang's infertility is a setback for captive breeding efforts, and also means that Scotland will miss out on the significant returns a panda cub could generate for the economy.

Beijing and Edinburgh zoos struck a deal that the pandas would stay in Scotland for 10 years for an annual fee of 600,000 pounds ($767,000).

That fee has been eclipsed by the money the pandas have generated for the zoo and the Scottish economy at large. During the bears' inaugural year in Edinburgh, the zoo experienced an increase of 5 million pounds in its annual turnover. The zoo's income jumped 53 percent to almost 15 million pounds in 2012.

An economic impact study conducted by business development body Scottish Enterprise before the bears arrived predicted that the animals would generate 28 million pounds in visitor expenditure in Edinburgh, and 20 million pounds for the wider Scottish economy, over a 10-year period.

In 2013, Scottish Enterprise speculated that the birth of a cub could generate a further 50 million pounds for Scotland, factoring in repeat visits, renewed public interest and increased media attention.

As part of the agreement between the two zoos, any resulting offspring would remain in Edinburgh. Tian Tian is believed to be have been pregnant on several occasions, but has thus far produced no cubs.

Successfully breeding pandas in captivity is a notoriously difficult endeavor. Females only ovulate once a year, and are only able to conceive for a few days during ovulation. Finding pandas that are compatible is also a tricky business -- many captive pairs show disinterest or incompetence when it comes to copulation.

Despite these challenges, a record 42 giant pandas were born in 2017 at two breeding stations in China operated by the China Conservation and Research Center for the Giant Panda.

Conservation efforts have also seen wild panda numbers rebound, from 1,112 in 1990 to 1,864 pandas in 2014 when the last census was conducted.