Another woman recalled witnessing the pandas’ growth journey. “Witnessing their growth, especially since they were so small, has been such a joy,” she said.

Some visitors were pictured sobbing as they bade the bears goodbye.

Xiao Xiao and Lei Lei were born in 2021 at the Ueno zoo to their mother Shin Shin and their father Ri Ri, both of whom were on loan to Japan for breeding research.

More recently, China’s panda loans have coincided with major trade agreements. In 2011, the loan of two pandas to Edinburgh Zoo in Scotland was agreed during negotiations over contracts to supply salmon meat, Land Rover vehicles and energy technology to China.

Many pandas have been returned to China in recent years – a loan agreement typically lasts 10 years although extensions are common.

However, the prospects of a new panda loan to Japan have remained uncertain amid the escalating row.

Japanese prime minister Sanae Takaichi’s comments on Taiwan angered Beijing which views the self-governed island as part of its territory and has not ruled out the use of force to “reunite” with it.

Both sides have since engaged in increasingly hostile actions and rhetoric towards each other. Early this month, China tightened restrictions on exports of rare earth-related products to Japan.