With just a fortnight to go before the historic referendum that will decide whether Britain remains a member of the European Union, Johnson was accused on Thursday night of misleading the public and putting his own career before the good of the country – including by his Conservative colleague energy secretary Amber Rudd.
The Scottish first minister described it as a “scandal” that Johnson’s Vote Leave campaign bus still has the controversial claim that Britain sends £350m a week to Brussels emblazoned on the side of it.
The figure has been called “potentially misleading” by the independent Statistics Authority, for failing to take into account the UK’s rebate from the EU. Sturgeon accused Johnson – who repeatedly defended the number during the debate – as “driving around the country in a bus with a giant whopper painted on the side”.
Labour’s Angela Eagle, also making the case for remain, joined Sturgeon and Rudd on the attack, in another of the extraordinary combinations that have characterised the referendum campaign. At one point Eagle jabbed her finger at Johnson, saying, “get that lie off your bus!”.
Rudd, who is regarded by some in her party as a potential future leader, also gave a punchy performance, describing Johnson as, “the life and soul of the party, but not the man you want driving you home at the end of the evening”.
She also suggested he was keener on furthering his own career than standing by his principles. “We need to look at the numbers – I fear the only number Boris is interested in is the one that says No 10”.
However, the former mayor of London appeared unruffled by the attacks on his character, and stuck to the line that a vote to leave would “take back control of our democracy” – a sentiment that was warmly received by the studio audience.
At one point during Thursday night’s debate, the former mayor of London urged the opposing panellists not to “reduce this argument to a lot of personal stuff” – and he was applauded by the audience when he hit back at Sturgeon, accusing her of being “more keen on being ruled from Brussels than ruled from Westminster”.
With Johnson widely regarded as the leave campaign’s strongest advocate, and the polls suggesting the result on 23 June will be too close to call, the Britain Stronger in Europe campaign, being coordinated from Downing Street, is keen to ramp up the pressure on him.
Stronger In will be unveiling a deliberately provocative poster on Friday morning, picturing Johnson gambling in a casino alongside the justice secretary Michael Gove, and with Ukip leader Nigel Farage, clutching a cigar.
Sturgeon offered a strong defence of immigration, saying the free movement of people that is a fundamental principle of the EU single market is a “two-way street”, which benefits UK citizens who can travel abroad, as well as EU migrants who can come to the UK and work.
“Immigration is causing pressure in some areas, Sturgeon said. “But the answer to that is to invest in our public services. If you are struggling to access public services, blame the politicians not the immigrants.”
She added: “The impact of austerity on our public services is much greater than the impact of immigration.”
And while Vote Leave has repeatedly claimed that exiting the EU would free up more resources to spend on public services including the NHS, Sturgeon said she “wouldn’t trust Boris Johnson with the NHS any further than I could throw Boris Johnson”, and said he wasn’t interested in protecting Britain’s workers, but only in seizing “David Cameron’s job”.
Rudd insisted that tackling immigration was a “complex issue”, for which there was no “silver bullet” – but that leaving the EU would hit the economy.
The shadow business secretary also hinted at her own leadership ambitions, when Johnson asked whether she could be the first woman leader of the Labour party, warning him to, “beware of the blonde bombshell”.
The overwhelming majority of Labour MPs support remaining in the EU, but outspoken backbencher John Mann announced on Thursday night that he would back Brexit. In an open letter to the Sun newspaper, Mann said Labour is “going to get a big shock across the country,” when the referendum result comes in on 24 June
Iain Duncan Smith, the former work and pensions secretary – who was spinning for Vote Leave, told the Guardian he believed the attacks on Johnson during the TV debate were carefully scripted.
“Tonight I thought was stark,” he said. “The remain side came on with the usual old scare stories about Britain not being good enough but what really added to that, was lacing its way through that, was just personal abuse. One after another. You could see their heads dip down to read the line: ‘Now time to abuse Boris Johnson’.”
In his closing statement, Johnson accused the remain camp of scaremongering, by repeatedly highlighting the risks of leaving the EU.
“The last couple of hours and the whole campaign is the contrast between this side offering hope and that side offering nothing but fear about life outside,” he said.
“They say we can’t do it on our own. We say we are a great country. We say we can. They say we have absolutely no choice but to stay locked in the back of the EU car driven in the wrong direction, driven in a destination we do not want to go.”
Andrea Leadsom, a Conservative who was participating for leave, received a warm response when she described Britain’s relationship with the EU as “a takeover by an unelected superstate”. A junior minister in the department for energy and climate change, Leadsom also had a spiky exchange with Rudd, her boss, who criticised her tone on immigration.
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