Moscow -- Russia on Tuesday buried the legendary human rights activist, Lyudmila Alexeyeva, whose fearless efforts to force the Soviet Union and later Russia's government to observe its citizens’ rights made her a national icon.

Alexeyeva died on Saturday at the age of 91.

One of the last well-known Soviet-era dissidents, Alexeyeva was the doyenne of Russia’s human rights movement and a symbol of the long struggle against political repression in the country. She had continued to campaign until her death and had appeared at rallies well into her eighties, denouncing the new turn back to authoritarianism under Vladimir Putin.

Even Putin could not ignore Alexeyeva’s stature and influence. He paid his respects at Tuesday's in-state ceremony.

But in a mark of the political realties that Alexeyeva was still fighting against, her long-time collaborator, 77-year-old Lev Ponomarev, was unable to attend his friend's funeral because he is currently serving a 16-day sentence for a Facebook post related to a protest over Putin's rule. A court refused his request to be allowed out for the funeral.

Putin’s presence at the funeral produced a strange juxtaposition, as he sat among a group of Russia’s best-known human rights defenders, many of whom have dedicated their lives to defending the rights and political freedoms that his government has sought to suppress. Many of them have also been detained at rallies protesting Putin.

Alexeyeva’s open casket lay on a stage in a hall, surrounded by wreaths and flanked by a rotating honor guard of her fellow rights activists, most of them also now aged.

Putin arrived on the stage after they had gathered and, after laying flowers and touching the casket, sat with Alekseeva’s son Mikhail, talking with him. After around 10 minutes, Putin excused himself and left.

Those at the event greeted Putin politely or looked on silently. In the eulogies that followed, some of Alexeyeva’s colleagues made veiled criticisms of Putin and his system.

Henry Reznik, a lawyer who had worked with Alexeyeva for decades, told how she had returned from a comfortable exile in the United States to campaign for human rights in Russia.

With a fury in his voice, he criticized "dishonest politicians" who "just do PR for themselves" with patriotic shows.

Alexeyeva had sought to find a language to communicate with Putin and had maintained good relations with him. She joined Putin's presidential council on human rights, though she abandoned it for a time over concerns it was becoming a puppet body. Putin had been careful to display respect to her, last year visiting her at home to wish her happy 90th birthday. She had used the occasion to press Putin to release a senator jailed in a controversial case.

Alexeyeva held a unique status among Russia's human rights defenders, universally admired for her unfailing principles and efforts to aid victims of persecution. Her funeral saw the leading figures of Russia's present-day opposition also pay their respects, including Alexey Navalny as well as a senior Putin ally, Vyacheslav Volodoin.

Alexeyeva began her dissident career in the 1960s, volunteering to type out the Chronicle of Current Events, a banned journal that detailed judicial abuses by the Soviet authorities. Such activities carried severe risk -- the Soviet Union remained a totalitarian police state and many dissidents were jailed in labor camps or forcibly imprisoned in psychiatric institutions. In 1976, she helped found the Moscow Helsinki Group, the USSR’s first human rights organization, joining a tiny group that stood up publicly to the Soviet regime. A year later, she was forced into exile, leaving for the U.S. where she spent 16 years.

She returned to Russia in 1993 and as Putin began to restore authoritarianism in a softer form, she turned her campaigns against him, warning that Putin was seeking to destroy Russia's civil society.

After Tuesday's ceremony, Putin attended a session of the presidential rights council, whose annual meeting fell on the same day. "I think all of you will agree with me that this was a shining, brave and strong person. She defended justice as her conscience guided her," Putin said of Alexeyeva at the council meeting that was televised.

Putin was asked by one of the council members to re-examine the jailing of Alexeyeva's colleague, Ponomarev, who was unable to attend her funeral. Putin promised to request it be looked into but also noted that Ponomarev was charged in relation to unauthorised political rallies and then referred to the chaotic protests in Paris.

"I, like you, also don't want events like those in Paris, where they are tearing up paving stones, burning everything, and then the country sinks into a state of emergency," Putin said.

Besides her unyielding belief, Alexeyeva was known for her mischievous sense of humor and quick tongue. She was arrested at a protest in 2009 at the age of 82, while dressed as a snow maiden. In their eulogies, several colleagues described how the tiny, kind-eyed elderly woman could deliver iron words when they were needed. They described how she would call the presidential administration and pressure officials there to act on specific political cases.

Alexeyeva has helped hundreds, perhaps thousands of people targeted in political cases over the years.

“She was an intercessor,” said Svetlana Gannushkina, a celebrated rights defender who in 2017 was awarded the Right Livelihood Award, sometimes called the "alternative Nobel Prize."

She was also an uncomfortable icon for Russia in recent years, as the Kremlin painted human rights NGOs as a Western instrument to weaken Russia.

Alexeyeva's own Moscow Helsinki Group was threatened by a 2012 law labeling human rights NGOs "foreign agents."

Her friend Reznik pledged that her efforts would be continued

"The job of human rights defending will be continued. Without a doubt," Reznik told mourners. "All those young souls -- those who feel that feeling that only on a basis of freedom and dignity can this life be fixed -- they will be proud of this remarkable woman."