estimate put the figure at 110,000, equivalent to about 10 percent of the Scottish male population aged between sixteen and fifty, and probably to about 15 per cent of total British war dead — the sacrifice was higher in proportionate terms than for any other country in the Empire." Second World War Scottish soldiers fought in many battles in World War II, in both the Pacific and European theatres. The Cold War & The End of Empire Defence establishments in Scotland Army In the wake of the Jacobite risings, several fortresses were built throughout the Highlands in the 18th century by General Wade in order to pacify the region, including Fort George, Fort Augustus and Fort William. The Ordnance Survey was also commissioned to map the region. Later, due to their topography and perceived remoteness, parts of Scotland have housed many sensitive defence establishments, some controversial. During World War II, Allied and British Commandos trained at Achnacarry in the Highlands and the island of Gruinard was used for an exercise in biological warfare. Regular British Army Garrisons currently operational in Scotland are: Fort George near Inverness; Redford Barracks and Dreghorn Barracks in Edinburgh; and Glencorse Barracks at Penicuik. Royal Naval Between 1960 and 1991, the Holy Loch was a base for the US Navy's fleet of Polaris-armed ballistic missile submarines. Today, HM Naval Base Clyde, west of Glasgow, is the base for the four Trident-armed ballistic missile submarines which are armed with approximately 200 Trident nuclear warheads. Since the decommissioning of free-falling bombs in 1998, the Trident SLBM system is the UK's only nuclear deterrent. HMS Caledonia at Rosyth in Fife is the support base for navy operations in Scotland and also serves as the Naval Regional Office (NRO Scotland and Northern Ireland). The Royal Navy's LR5 and Submarine Rescue Service is based in Renfrew, near Glasgow. The Royal Navy's submarine nuclear reactor development establishment, is located at Vulcan NTRE, adjacent to Dounreay, which was the site of the UK's fast breeder nuclear reactor programme. RM Condor at Arbroath, Angus is home to 45 Commando, Royal Marines, part of 3 Commando Brigade. Also, the Fleet Protection Group Royal Marines is based at HMNB Clyde. Since 1999, the Scottish Government has had devolved responsibility over fisheries protection duties in Scotland's exclusive economic zone, carried out by the Scottish Fisheries Protection Agency, which consists of a fleet of four offshore patrol vessels