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The Underground has its origins in the Metropolitan Railway, opening on 10 January 1863 as the world's first underground passenger railway.[6] It is now part of the Circle, District, Hammersmith & City and Metropolitan lines. The first line to operate underground electric traction trains, the City & South London Railway in 1890, is now part of the Northern line.[7]

The idea of an underground railway linking the City of London with the urban centre was proposed in the 1830s,[21] and the Metropolitan Railway was granted permission to build such a line in 1854.[22] To prepare construction, a short test tunnel was built in 1855 in Kibblesworth, a small town with geological properties similar to London. This test tunnel was used for two years in the development of the first underground train, and was later, in 1861, filled up.[23] The world's first underground railway, it opened in January 1863 between Paddington and Farringdon using gas-lit wooden carriages hauled by steam locomotives.[24] It was hailed as a success, carrying 38,000 passengers on the opening day, and borrowing trains from other railways to supplement the service.[25] The Metropolitan District Railway (commonly known as the District Railway) opened in December 1868 from South Kensington to Westminster as part of a plan for an underground "inner circle" connecting London's main-line stations.[26] The Metropolitan and District railways completed the Circle line in 1884,[27] built using the cut and cover method.[28] Both railways expanded, the District building five branches to the west reaching Ealing, Hounslow,[29] Uxbridge,[30] Richmond and Wimbledon[29] and the Metropolitan eventually extended as far as Verney Junction in Buckinghamshire, more than 50 miles (80 km) from Baker Street and the centre of London.[31]

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Yerkes soon had control of the District Railway and established the Underground Electric Railways Company of London (UERL) in 1902 to finance and operate three tube lines, the Baker Street and Waterloo Railway (Bakerloo), the Charing Cross, Euston and Hampstead Railway (Hampstead) and the Great Northern, Piccadilly and Brompton Railway, (Piccadilly), which all opened between 1906 and 1907.[41][42] When the "Bakerloo" was so named in July 1906, The Railway Magazine called it an undignified "gutter title".[42] By 1907 the District and Metropolitan Railways had electrified the underground sections of their lines.[43]

In 1933, most of London's underground railways, tramway and bus services were merged to form the London Passenger Transport Board, which used the London Transport brand.[54] The Waterloo & City Railway, which was by then in the ownership of the main line Southern Railway, remained with its existing owners.[55] In the same year that the London Passenger Transport Board was formed, Harry Beck's diagrammatic tube map first appeared.[56]

In 1979 another new tube, the Jubilee line, named in honour of Queen Elizabeth's Silver Jubilee, took over the Stanmore branch from the Bakerloo line, linking it to a newly constructed line between Baker Street and Charing Cross stations.[76] Under the control of the GLC, London Transport introduced a system of fare zones for buses and underground trains that cut the average fare in 1981. Fares increased following a legal challenge but the fare zones were retained, and in the mid-1980s the Travelcard and the Capitalcard were introduced.[77]

On 18 November 1987, fire broke out in an escalator at King's Cross St Pancras tube station. The resulting fire cost the lives of 31 people and injured a further 100. London Underground was strongly criticised in the aftermath for its attitude to fires underground, and publication of the report into the fire led to the resignation of senior management of both London Underground and London Regional Transport.[80] Following the fire, substantial improvements to safety on the Tube were implemented - including the banning of smoking, removal of wooden escalators, installation of CCTV and fire detectors, as well as comprehensive radio coverage for the emergency services.[81]

Fifty-five per cent of the system runs on the surface. There are 20 miles (32 km) of cut-and-cover tunnel and 93 miles (150 km) of tube tunnel.[1] Many of the central London underground stations on deep-level tube routes are higher than the running lines to assist deceleration when arriving and acceleration when departing.[108] Trains generally run on the left-hand track. In some places, the tunnels are above each other (for example, the Central line east of St Paul's station), or the running tunnels are on the right (for example on the Victoria line between Warren Street and King's Cross St. Pancras, to allow cross-platform interchange with the Northern line at Euston).[107][109]

London Underground trains come in two sizes, larger sub-surface trains and smaller deep-tube trains.[126] Since the early 1960s all passenger trains have been electric multiple units with sliding doors[127] and a train last ran with a guard in 2000.[128] All lines use fixed-length trains with between six and eight cars, except for the Waterloo & City line that uses four cars.[129] New trains are designed for maximum number of standing passengers and for speed of access to the cars and have regenerative braking and public address systems.[130] Since 1999 all new stock has had to comply with accessibility regulations that require such things as access and room for wheelchairs, and the size and location of door controls. All underground trains are required to comply with The Rail Vehicle Accessibility (Non Interoperable Rail System) Regulations 2010 (RVAR 2010) by 2020.[131]

The current standard tube map shows the Docklands Light Railway, London Overground, IFS Cloud Cable Car, London Tramlink and the London Underground;[271] a more detailed map covering a larger area, published by National Rail and Transport for London, includes suburban railway services.[201] The tube map came second in a BBC and London Transport Museum poll asking for a favourite UK design icon of the 20th century[272] and the underground's 150th anniversary was celebrated by a Google Doodle on the search engine.[273][274]

After the UERL was absorbed into the London Passenger Transport Board in 1933, it used forms of the roundel for buses, trams and coaches, as well as the Underground. The words "London Transport" were added inside the ring, above and below the bar. The Carr-Edwards report, published in 1938 as possibly the first attempt at a graphics standards manual, introduced stricter guidelines.[281] Between 1948 and 1957 the word "Underground" in the bar was replaced by "London Transport".[282] As of 2013[update], forms of the roundel, with differing colours for the ring and bar, are used for other TfL services, such as London Buses, Tramlink, London Overground, London River Services and Docklands Light Railway.[283] Crossrail will also be identified with a roundel.[284] The 100th anniversary of the roundel was celebrated in 2008 by TfL commissioning 100 artists to produce works that celebrate the design.[285][286] Roundels are featured outside many underground stations; they are commonly mounted on a white pole known as a "Venetian mast".[287] In some cases, the mast is merely a decorative feature, with the roundel being mounted to a wall, and the mast supported by the roundel, rather than vice versa.[citation needed]

These differences can be interpreted in a straightforward way as adaptations to a subterranean life: the ability to breed in confined spaces is a prerequisite; the greater warmth underground during winter means that there is no need for a diapause; nutrients are more readily obtained in the larval stage or from mammals rather than from birds. These differences are heritable (Pasteur, 1977), and presumably maladaptive in the above-ground environment. Interbreeding with above-ground populations is additionally hampered by the physical separation of the two habitats.

The lack of demonstrable gene flow between the two forms in London may, in part, be explained by the implications of the breeding experiments, in which no offspring were recorded from crosses between Underground and surface populations. The ability to mate in confined spaces (stenogamy) may be a major contribution to premating reproductive isolation. Intriguingly, the Oval population, which seems to be an essentially overground population living underground (but below open sky) produced one (F1) egg raft. The F2 eggs, however, failed to hatch.

The data are more consistent with local populations being the source of molestus populations in the Underground. However, Fig. 1 shows that there is no association between individual Underground populations and local surface populations, which suggests that multiple independent colonizations are unlikely or, at least, that such events are rare. The most parsimonious explanation is a single colonization. This would explain the fixation of a single allele at multiple allele loci such as the esterases. The similarity of the Underground populations to the more distant Beckton population (nearest Underground line 4 km) raises the possibility that adaptation to the underground environment may have occurred only once, and that the established subterranean populations are the source for subsequent colonizations.

This combination of factors can explain why gene flow is negligible or absent between the two forms in the populations that were studied. As it appears plausible that colonization of the London Underground has occurred only once, the reproductive isolation could have accumulated in the few hundred generations since then. Conversely, in the south of the range, where climate promotes more frequent gene flow, this may be sufficient to prevent the establishment of reproductive isolation, even though the pleiotropic effects of selection in the underground environment are the same.

Comparing our results with influenza-like illnesses (ILI) data collected by Public Health England (PHE) in London boroughs, shows a correlation between the use of public transport and the spread of ILI. Specifically, we show that passengers departing from boroughs with higher ILI rates have higher number of contacts when travelling on the underground. Moreover, by comparing our results with other demographic key factors, we are able to discuss the role that the Underground plays in the spread of airborne infections in the English capital. ff782bc1db