WASHINGTON — Former FBI director James Comey testified Wednesday before a Republican-led Senate committee seeking to discredit the investigation he opened during the 2016 election into ties between Donald Trump’s campaign and Russia.

With another presidential election looming, Republicans on the Senate Judiciary Committee were eager to portray Trump as a victim of a politically motivated smear by the FBI that unfairly cast a shadow over his presidency. And they contended that Comey was the ringleader.

Comey strongly defended the FBI’s handling of the investigation, including his decision to open it. But he acknowledged, as he has before, that his initial claims were wrong that a wiretap of a former Trump campaign adviser, Carter Page, was properly handled and conceded that the bureau had been sloppy on that aspect of the broader inquiry.

He testified by video from his home.

Sen. Lindsey Graham of South Carolina, the Judiciary Committee chairman, renewed his criticism of the FBI’s investigation of ties between Russian election interference and the Trump campaign.

The panel has for months pounded away at the inquiry, building its work on an investigation by the Justice Department’s independent inspector general, Michael E. Horowitz, that found evidence of negligence and errors in one narrow aspect of the investigation: the FBI’s applications to wiretap Page. But where the inspector general concluded there was no evidence of illegal activity or a politically motivated plot by senior department officials, Graham insists there may have been.

Comey signed off on some of the certifications for the warrant applications and, as director, was the top bureau official responsible for the investigation until he was fired by Trump in the spring of 2017.

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But in an opening statement, Graham more narrowly trained his focus on the secret wiretap warrants and made nary a mention of Comey.

“I’m saying this to my Democratic friends: If it happened to us, it could happen to you. Every American should be worried about this,” Graham said. “This is not just an abuse of power against Mr. Page and the Trump campaign. This is a system failure.”

The committee has already publicly questioned two former deputy attorneys general, Rod Rosenstein and Sally Yates, who oversaw the Russia investigation and signed off on the applications for the secret wiretap warrants targeting Page. Both expressed regret for errors identified by Horowitz but dismissed assertions by Republicans on the panel that their actions were politically motivated or that Trump’s campaign need not have been investigated.

Democrats have opposed Graham at every turn, accusing him of abusing his Senate powers to help Trump and take attention from the continuing Russian threat. On Wednesday, they said he was unfairly trying to discredit the entire investigation based on one small aspect of it, a dossier of unverified information compiled by a British former spy, Christopher Steele, that investigators relied in part on to secure court permission for the Page wiretaps.

“Those errors were serious, but the errors and the so-called Steele dossier — and this is important — played no part in the broader Russia investigation,” said Sen. Dianne Feinstein of California, the panel’s top Democrat.

She noted that of the 10 people interviewed in the committee’s investigation, not one had claimed anything different.

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Comey, who had not testified before Congress since Horowitz’s report was released in December, remained steadfast in his decision to open the investigation, arguing that the FBI had sufficient reason to scrutinize the ties between the Trump campaign and Russia.

“In the main, it was done by the book, it was appropriate and it was essential that it be done,” Comey said under questioning by Graham. “Overall I am proud of the work, but there are parts of it we will talk about that are concerning.”

Comey described the problems with the wiretap as sloppy and problematic and, pressed by Graham, said he would not have signed off on the warrant applications knowing what he does now.

“The collection of omissions, failure to consider updates, to communicate between the team trying to figure out what’s true or not true in the Steele material and the team investigating Carter Page — it’s embarrassing, it’s sloppy, I’ve run out of words,” Comey said.

“There is no indication — the inspector general would say it if he found it — that people were doing bad things on purpose,” he added, “but that doesn’t make it any less concerning or embarrassing.”

He noted earlier that the wiretap applications were a small part of the larger inquiry into links between the Trump campaign and Russia.

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“The overarching investigation was very important — the Page slice of it, far less given the scope,” Comey said.

Comey also decried Attorney General William Barr’s denunciations of the investigation, including his assertion that the FBI lacked sufficient reason to open it.

“He says that a lot — I have no idea what on earth he’s talking about,” Comey said, noting that the special counsel who took over the investigation, Robert Mueller, secured dozens of indictments and that a bipartisan Senate Intelligence Committee inquiry found that onetime Trump campaign chairman Paul Manafort shared proprietary campaign information with a known Russian intelligence officer.

“The notion that the attorney general believes that was an illegitimate endeavor to investigate? That mystifies me,” Comey added.

Comey said that he had closely followed the Justice Department’s request in May that a federal judge throw out the case against Trump’s former national security adviser Michael T. Flynn, who had twice pleaded guilty to lying to the FBI about his contacts with the Russian ambassador.“The department’s conduct in handling it is deeply concerning,” Comey said, adding, “It’s deeply concerning because it’s this guy is getting treated in a way that nobody’s ever been treated before.”

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Republicans grew increasingly frustrated when Comey repeatedly said he could not remember details surrounding the investigative process and demurred on their questions.

“With all due respect, Mr. Comey, you don’t seem to know anything about an investigation that you ran,” said an irate Sen. Mike Lee, R-Utah.

Graham sought to focus on unverified intelligence on Russia.

As Comey defended the broader Russia investigation and played down concerns about the Page wiretap applications, Graham shifted focus to a different theory that he argued demonstrated the FBI’s bias, muddying in the process the fact that U.S. intelligence agencies concluded that Russia interfered in 2016 to aid Trump.

Graham built his case around newly released unverified intelligence made public a day earlier by John Ratcliffe, the director of national intelligence, in an apparent bid to help Trump politically. The years-old intelligence, rejected by other investigators, suggested that Russian intelligence officers had acquired information that Hillary Clinton had approved a campaign plan to elevate concerns about Trump’s ties to Russia.

Pressed by Graham, Comey said he was not aware of the intelligence. Graham inaccurately insisted it was part of a pattern suggesting that Democrats had helped manufacture material to justify the FBI’s investigation of the Trump campaign, and then exploited that investigation for political gain.

“I’m beginning to understand there was a two-tiered system here,” Graham said. “When it came to Trump, there were no rules, plow ahead, ignore everything, lie if you need to, alter documents. When it came to Clinton, it seems to be a completely different standard.”

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Comey disagreed.

“I have read Mr. Ratcliffe’s letter, which frankly I have trouble understanding,” he said.

Graham’s characterization was misleading. The FBI opened its Russia investigation independently of Steele’s work, which played a role in only the Page wiretap applications, not the other parts of the inquiry, in which investigators secured more than 230 orders for communications records.

Former officials said the newly released unverified intelligence has little credibility and could have also been either Russian misinformation or merely Russian analysis of U.S. politics. Sen. Sheldon Whitehouse, D-R.I., said the unverified intelligence suggested that Clinton’s campaign was simply sounding an alarm about the Trump campaign’s ties to Russia.

“It’s really hard to call a campaign’s effort to blow the whistle on the other side’s communications with Russia and Russia’s efforts to support that candidate with the actual efforts to do that,” Whitehouse said. “They are not the same thing.”

Mueller and the Republican-led Senate Intelligence Committee had both previously rejected the intelligence, and career intelligence strenuously officials were said to oppose Ratcliffe’s disclosure of it.

“I am really concerned that we are treating this Ratcliffe letter as something at all serious or credible,” Whitehouse said.

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Comey denounced an unfounded assertion pushed by Republicans — including Trump at the first presidential debate on Tuesday — that Biden came up with the legal theory for prosecuting Flynn in an Oval Office meeting in the final days of the Obama administration.

The Justice Department has released handwritten FBI notes that Trump and his allies claim as supporting evidence. But former FBI officials say Republicans are misrepresenting the notes and taking them out of context.

Comey, who attended the meeting, said that Biden had played no role in directing the investigation and that Obama had instructed Comey to follow procedure in handling it.

“I would remember it because it’d be highly inappropriate for a president or vice president to suggest prosecution or investigation of anyone — and it did not happen,” Comey said.Comey said Trump’s debt could make him vulnerable to foreign influence.

Sen. Dick Durbin, D-Ill., briefly steered the discussion to a recent New York Times report that found, based on years of Trump’s tax information, that the president is personally responsible for loans and debts totaling $421 million.

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Comey said that such a scenario could pose a national security threat because foreign adversaries can use government officials’ personal finances as leverage over them and would be the type of information that law enforcement and intelligence officials consider when deciding whether to grant officials access to sensitive or classified government information.

“A person’s financial situation could make them vulnerable to coercion to an adversary, and allow an adversary to do what we try to do to foreign government officials that we find are indebted: that is, recruit them to our side,” Comey said.

“So as a general matter, are there serious risks when someone with hundred of millions of dollars in debt, personal debt, has access, as the president does, to all of the country’s classified and sensitive information?” Durbin asked.

Comey said he could not speak to the specifics of Trump’s case, which the president has denied, but added: “It’s a serious concern when anyone seeking or with a clearance has that kind of financial vulnerability.”