title: Trump–Ukraine scandal

text: Marie Yovanovitch and Bill Taylor resigned from their positions in the State Department. Jennifer Williams left her position to take up a new post. Alexander Vindman was dismissed from his position in the White House following Trump's acquittal by the Senate. Vindman's twin brother Yevgeny—who was not involved in the case—was also dismissed. Both Vindman brothers were reported to have been physically escorted from the White House. Gordon Sondland was also recalled from his position as ambassador. The White House claimed that the dismissals were necessary, but Trump was criticized for seeking revenge against those who had testified against him. Trump was also reported to have labelled Williams and Alexander Vindman as "Never Trumpers". Trump suggested that the Pentagon should seek disciplinary action against Vindman, but the Army declined to investigate. John Rood, the top Pentagon policy advisor who, on July 25, 2019, warned Defense Secretary Mark Esper against withholding military aid to Ukraine, was forced to resign on February 19, 2020. In May 2019 he had certified to Congress that Ukraine was eligible for the aid.


title: Trump–Ukraine scandal

text: Subsequently, during a panel discussion held on February 11, 2020, at the Atlantic Council, the president's national security advisor, Robert C. O'Brien said that it was his decision to transfer both Vindman brothers back to the Army for re-assignment and denied that the move was ordered by Trump in retaliation for Vindman's testimony. "I can absolutely tell you that they were not retaliated against", O'Brien told the panel. O'Brien also disputed the move as being characterized as "fired" since both brothers remain on active duty. O'Brien noted that their transfer was part of a larger NSA staff reduction. It was later reported that the firings and dismissals were part of a wider purge of the Trump administration that targeted people who were perceived as not being loyal enough to Trump and his agenda, including intelligence officials who might be part of Trump's claims of a "deep state" conspiracy against him.


title: Trump–Ukraine scandal

text: Michael Atkinson was fired from his position as Inspector General of the Intelligence Community in April 2020. The White House issued a statement that Trump had lost confidence in Atkinson's ability to perform his duties. The decision to fire Atkinson was criticized because his role as Inspector General was supposed to be independent and because Trump chose to fire him during the coronavirus pandemic, which led to claims that Trump had tried to use the pandemic to distract from Atkinson's firing.


title: Trump–Ukraine scandal

text: As the impeachment hearings and trial unfolded, Rudy Giuliani returned to Ukraine to conduct his own investigation into Joe and Hunter Biden. This was widely criticized as a further attempt to undermine Biden's election campaign, for promoting widely debunked conspiracy theories about the Bidens, for Giuliani being a likely target for misinformation spread by Russian intelligence services, and because Giuliani is himself under investigation by American authorities. Giuliani went on to claim that he had found evidence that Barack Obama and Joe Biden had previously contacted Ukrainian officials looking to open an investigation into Paul Manafort, the chair of Trump's 2016 presidential campaign, who had lobbied to American lawmakers on behalf of former Ukrainian president Viktor Yanukovych.


title: Trump–Ukraine scandal

text: In February 2020, Attorney General William Barr announced that the Justice Department would receive any information gathered by Giuliani. Barr had previously announced that all investigations into foreign donations and interference into the 2020 presidential elections would require his personal approval.


title: Trump–Ukraine scandal

text: During the impeachment process, Trump denied having sent Giuliani to Ukraine, but in an interview that aired on February 13, 2020, he reversed his prior denials and openly admitted sending Giuliani to Ukraine, praising him as a "crime fighter" and "the best prosecutor."


title: Trump–Ukraine scandal

text: In February 2020, United States attorney John Durham was appointed to lead an investigation into the origins of the Mueller inquiry. It was reported that the investigation was focusing on former CIA director and Trump critic John Brennan and whether he had mishandled evidence during the early stages of the inquiry into Russian meddling in the 2016 elections. Mick Mulvaney linked Durham's investigation to the Ukraine scandal, stating that Durham had sought help from Ukraine and interviewed Ukrainian citizens. The Durham inquiry has been described as an "inquiry into its own Russia investigation". and "investigating the investigators" of the Russian interference.<ref name="Sullivan_10/2/2019"></ref><ref name="Berenson_10/4/2019"></ref>


title: Trump–Ukraine scandal

text: Alexander Vindman, the former Director for European Affairs who supported the whistleblower's testimony, claimed Trump's intervention weakened Ukrainian efforts to counteract Russian aggression in a number of ways in an interview with VICE News, saying "It’s because of Trump’s corruption that we have a less capable, less prepared Ukraine".


title: Trump–Ukraine scandal

text: Trump and his allies had since 2017 promoted a conspiracy theory that Ukraine, rather than Russia, had interfered with the 2016 election, which American intelligence believes has been promoted by Russia in order to frame Ukraine. Some speculate Maria Zakharova at Russia's Ministry of Foreign Affairs was the source of this campaign. The conspiracy theory included allegations that Democrats, CrowdStrike and the FBI had conspired to frame Russia in the 2016 hacking of a Democratic National Committee server. Trump has repeatedly insisted without evidence that an unnamed Ukrainian oligarch was behind the conspiracy to frame Russia and that Ukraine is in possession of the DNC server.


title: Trump–Ukraine scandal

text: Shortly before Trump took office, top American intelligence officials briefed him on their evidence—including from their hacking of Russian intelligence networks and information provided by a high-level Kremlin mole—that Russia was behind the hacking and other election interference, on the personal orders and orchestration of Vladimir Putin. In December 2019, the Trump-appointed FBI director Christopher Wray stated, "we have no information that indicates that Ukraine interfered with the 2016 presidential election," adding, "there's all kinds of people saying all kinds of things out there. I think it's important for the American people to be thoughtful consumers of information and to think about the sources of it and to think about the support and predication for what they hear."


title: Trump–Ukraine scandal

text: During the July 25, 2019, phone call between Trump and Zelenskyy, Trump referred to a far-right conspiracy theory pushed by internet trolls, right-wing blogs, right-wing news websites and Russian state media. This conspiracy theory concerns CrowdStrike, the cybersecurity and internet security firm that first investigated the 2015–2016 hacking of the Democratic National Committee (DNC) network and determined that Russian military intelligence (GRU) was behind these cyber attacks.


title: Trump–Ukraine scandal

text: Tom Bossert, Trump's former homeland security advisor, said in an interview with ABC News' George Stephanopoulos that Trump was repeatedly warned by his staff that the CrowdStrike conspiracy theory was "completely debunked". Bossert blamed Giuliani for Trump's fixation upon the conspiracy theory.


title: Trump–Ukraine scandal

text: The overarching theme of the CrowdStrike conspiracy theory is that the DNC fabricated evidence to implicate Russia in the cyber attacks. CrowdStrike's co-founder, Dmitri Alperovitch, is a naturalized American citizen born in the Soviet Union. According to the hoax, Alperovitch is a Ukrainian who was ordered by the DNC to discredit Russia for the election interference, and he was personally motivated to get even with Vladimir Putin. Also, according to the theory, CrowdStrike is owned by a rich Ukrainian, and the actual server involved in the cyber attack is in Ukraine.


title: Trump–Ukraine scandal

text: CrowdStrike is actually a publicly traded company headquartered in California that the National Republican Congressional Committee has also hired for cyber security services. "The server" is actually 140 servers, decommissioned and located in the United States. The theory additionally says FBI agents were not allowed to examine the server because such action would expose the DNC plot, although in fact—and as documented in the Mueller Report—system images and traffic logs of the DNC servers were provided to the FBI. This conspiracy theory originated from a "GRU persona, 'Guccifer 2.0', created to cast doubt on Russia's culpability in the DNC [intrusion]".


title: Trump–Ukraine scandal

text: Various right-wing commentators speculated the whistleblower had help from others, perhaps constituting a coordinated conspiracy. Speculation centered around Adam Schiff, the press, Fusion GPS, Media Matters, a team of lawyers or a research firm, and the intelligence community in general. After the whistleblower had informed the CIA's general counsel of his concerns, he grew troubled by "how that initial avenue for airing his allegations through the CIA was unfolding", according to "The New York Times". He then contacted an aide for the House Intelligence Committee and provided a vague statement. The aide then followed standard procedure and advised the whistleblower to find a lawyer and file a complaint with the Intelligence Community inspector general (IC IG). Neither Rep. Schiff nor the other members of the Committee saw the complaint until the night before they released it publicly, and the Committee was not involved in writing the complaint. Schiff and the Committee had no role in helping the whistleblower select an attorney.


title: Trump–Ukraine scandal

text: According to Mark S. Zaid, a member of the whistleblower's "pro bono" legal team: "The whistleblower took the advice to find an attorney and did what most people do, they asked around to trusted friends as to who they should contact. Andrew [Bakaj]'s name was provided and he was retained. Exactly how it happens every day." Andrew P. Bakaj is the Lead Attorney representing the whistleblower. During a news conference on October 2, Trump claimed that "The New York Times" article proved Schiff had helped write the whistleblower complaint, prompting one of the reporters who wrote the story to reply on Twitter that their story said no such thing and that Schiff had not even known the whistleblower's identity.


title: Trump–Ukraine scandal

text: In late September, Trump tweeted a conspiracy theory that whistleblower rules were changed before the whistleblower complaint was submitted. Senator Lindsey Graham, and Trump's lawyers Jay Sekulow and Rudy Giuliani made similar claims. Trump's claim was based on an article from "The Federalist" which incorrectly stated that the IC IG "secretly eliminated a requirement that whistleblowers provide direct, first-hand knowledge of alleged wrongdoings", by revising their complaint form sometime between May 2018 and August 2019, removing a section from the old form containing the sentence: "If you think wrongdoing took place, but can provide nothing more than secondhand or unsubstantiated assertions, IC IG will not be able to process the complaint or information for submission as an ICWPA." "The Federalist" article failed to mention that the old form had checkboxes where the whistleblower could indicate that their information was "direct" or from either "other employees" or other indirect sources.


title: Trump–Ukraine scandal

text: The IC IG responded the whistleblower's complaint was submitted with the old form (before the forms changed), and that the whistleblower's complaint was based on both "direct knowledge of certain alleged conduct" and knowledge from other employees. The IC IG also said the old form had been under review, and that "in response to recent press inquiries regarding the instant whistleblower complaint", the form was changed because "certain language in those forms and, more specifically, the informational materials accompanying the forms, could be read—incorrectly—as suggesting that whistleblowers must possess first-hand information in order to file an urgent concern complaint". The IC IG also said that by law a complainant is not required to have "first-hand information" themselves, and that their office "cannot add conditions to the filing of an urgent concern that do not exist in law".


title: Trump–Ukraine scandal

text: The "rules" for whistleblowing arise from Intelligence Community Directive 120, last updated in 2016. The directive states that the requirement for a complaint is to be one which the whistleblower "reasonably believes evidences a violation of any law, rule or regulation". The burden of obtaining and evaluating first-hand knowledge for credibility is placed on the IC IG, who has 14 days to conduct an investigation to do so. In this case, the preliminary review done by the IC IG did find more information to support the allegations as credible. Tom Devine of the Government Accountability Project, a non-profit watchdog organization, said only around 10% of all credible whistleblower complaints have firsthand information.


title: Trump–Ukraine scandal

text: Republican senator Chuck Grassley, a prominent author and advocate of whistleblower laws, spoke out against the conspiracy theory, saying the whistleblower appeared to have acted in accordance with the law and deserved to be heard.


title: Trump–Ukraine scandal

text: In late-September television appearances, Giuliani asserted without offering any evidence that George Soros, a frequent subject of conservative conspiracy theories, was running an anti-Trump scheme in Ukraine while Biden was protecting Soros from prosecution there. Lawyers Joseph diGenova and Victoria Toensing appeared as guests on "The Sean Hannity Show" to promote the conspiracy theory that Soros funded the whistleblower. They cited the whistleblower's footnote references to the Organized Crime and Corruption Reporting Project, an organization that has received grants from Soros' Open Society Foundations among multiple other funding sources. Soros was also invoked, again without evidence, by the media organization "Breitbart News".


title: Trump–Ukraine scandal

text: Discredited allegations against Soros were also part of a "packet of propaganda and disinformation" that had been circulating within the State Department since May 2019, until being revealed to Congress on October 2.


title: Davy Jones (character)

text: Davy Jones is a fictional character in the "Pirates of the Caribbean" film series, portrayed by Bill Nighy.


title: Davy Jones (character)

text: He is first mentioned in "", appears in "" and "", and makes a cameo appearance in "". Once a mortal pirate of Ulster Scots descent, Davy Jones is the tyrannical captain of the Flying Dutchman (based on the ghost ship of the same name) and the dark lord of the Seven Seas.


title: Davy Jones (character)

text: The computer-generated imagery used to complete Jones was named by "Entertainment Weekly" as the tenth favorite computer-generated film character in film history, behind King Kong in 2007. The work on Davy Jones by Industrial Light and Magic earned them the 2006 Academy Award for Visual Effects for "Dead Man's Chest".


title: Davy Jones (character)

text: Before officially casting Bill Nighy, producers also met Jim Broadbent, Iain Glen and Richard E. Grant for the role.


title: Davy Jones (character)

text: Like the entire crew of the "Flying Dutchman" (except "Bootstrap Bill"), Davy Jones' physical appearance is completely computer-generated. Nighy's performance was recorded using motion capture during actual filming on the set, with Nighy wearing several markers in both a grey suit and his face, rather than in a studio during post-production. Nighy wore make-up around his eyes, since the original plan was to use his real eyes in the digital character if necessary to get the proper lighting. In addition, he also wore make-up on his lips and around his mouth, to assist in the motion capture of his character's Scottish accent. Davy Jones briefly appears as a human for a single scene in the third film, played by Nighy in costume. Several reviewers have in fact mistakenly identified Nighy as wearing prosthetic makeup or a latex mask due to the computer-generated character's photorealism.


title: Davy Jones (character)

text: The physique of Davy Jones and the other crew of the "Flying Dutchman" was designed by the films' producers to be a mixture of various aquatic flora and fauna features. In the third film, Tia Dalma reveals it is the fate of the captain and crew of the "Dutchman" should they fail or abandon their duties on board.


title: Davy Jones (character)

text: Jones' most striking feature is his cephalopod-like head: prehensile octopus-like appendages give the illusion of a thick beard, a prominent sac bulges from under his barnacle-encrusted tricorne, and he breathes through a siphon located on the left side of his face in the place of nasal features. Jones has a crustacean-style claw for his left arm, a long tentacle in place of the index finger on his right hand, and the right leg of a crab (resembling a pegleg). It is revealed in the bonus features of the Special Edition DVD that Jones' skin color was partly inspired by a coffee-stained styrofoam cup which was then scanned into ILM's computers to be used as such.


title: Davy Jones (character)

text: Jones speaks with a clearly distinguishable, albeit thick, Scottish accent that's slightly altered to account for his lack of a nose, and presumably, a nasal cavity and/or sinuses. Originally, director Gore Verbinski wanted Jones to be Dutch, as he is the captain of the "Dutch-man". Nighy however responded, "I don't do Dutch. So I decided on Scottish." Nighy later revealed that the Scottish sitcom "Still Game" influenced his choice of accent, stating: "I had to find an accent no one else had. Although Alex Norton is Scottish, mine was slightly different. We wanted something that was distinctive and authoritative...I have seen "Still Game" and I am a fan. The sort of extremity of the accent was inspired in that area."


title: Davy Jones (character)

text: Davy Jones, a mortal Scottish pirate and a great sailor, fell in love with Calypso, a sea goddess. She entrusted him with the task of ferrying the souls of those who died at sea to the next world. Calypso gave him the "Flying Dutchman" to accomplish this task. She swore that after ten years, she would meet him and they would spend one day together before he returned to his duties. However, when Jones returned to shore after ten years, Calypso failed to appear. Believing Calypso had betrayed him, a heartbroken and enraged Davy Jones turned the Pirate Brethren against her, saying that if she were removed from the world, they would be able to claim the seas for themselves. They assembled in the First Brethren Court and Jones successfully taught them how to imprison her into her human form.


title: Davy Jones (character)

text: Despite betraying her, Jones still loved Calypso, and in despair and guilt for what he had done, he carved out his own heart from his body and placed it in the "Dead Man's Chest". The Chest was sealed and placed within a larger wooden chest, along with Jones' numerous love letters to Calypso and all other items having to do with her, except his matching musical locket. The chest was then buried on Isla Cruces. Jones kept the chest's key with him at all times, leaving the locket beside the massive pipe organ aboard the "Dutchman". With Calypso gone, Jones abandoned his duties and returned to the Seven Seas. As a result of this, Jones gradually became monstrous, his physical appearance merging with various aquatic fauna. Sailors everywhere would fear him to the death, for Davy Jones had turned fierce and cruel, with an insatiable taste for all things brutal. Jones recruits dying sailors by promising them a reprieve from death in exchange for 100 years of service aboard the "Dutchman". During this time, he comes to command the Kraken, a feared mythological sea monster.


title: Davy Jones (character)

text: In the about Jack Sparrow's earlier adventures, Davy Jones shows interest in the Sword of Cortes, also sought by Jack. He is a minor character, but appears in the seventh book as Jack and his crew encounter the "Flying Dutchman".


title: Davy Jones (character)

text: Jones also appears in the about Jack's first years as a captain. He helps the Brethren Court to identify the traitor among them, who turns out to be Borya Palachnik, the Pirate Lord of the Caspian Sea.


title: Davy Jones (character)

text: Before the events of the first film, Davy Jones approaches Sparrow with a deal: Jones will raise the "Black Pearl" back from Davy Jones' Locker, allowing Sparrow to be captain for 13 years if Sparrow agrees to serve on the "Dutchman" for 100 years. This event, referenced in the films, also appears in the book series.


title: Davy Jones (character)

text: Davy Jones first appears in the second film, "", in which he attempts to collect on his bargain with Jack Sparrow, sending Bootstrap Bill to threaten Jack and give him the black spot to attract the Kraken. Previously, Davy Jones had raised the "Black Pearl" from the sea for Sparrow, in exchange for Sparrow's soul after captaining the "Pearl" for 13 years. Attempting to get out of the situation, Sparrow argues that he was the captain for only two years before Hector Barbossa committed mutiny. Jones rejects this explanation, explaining that despite the mutiny, Jack still stubbornly gave himself the title "Captain". Sparrow then attempts to escape the deal by providing Will Turner as a substitute for himself. Jack falsely strikes a new deal with Jones; Jack will be spared enslavement on the "Dutchman" if he brings Jones one hundred souls to replace his own within the next three days (in reality he plans to recruit a new crew to help him find the "Dead Man's Chest" to save Will and break his debt). Jones accepts, removes the black spot from Jack's hand, and retains Will, keeping him as a "good faith payment."


title: Davy Jones (character)

text: While on the "Dutchman", Will challenges Jones at a game of liar's dice. They wager Will's soul for an eternity of service against the key to the "Dead Man's Chest". Bootstrap Bill joins the game and purposefully loses to save Will. During the game, Will learns where Jones keeps the key, being his real purpose in the game. The next morning, Jones realizes the key is gone and summons the Kraken to destroy the ship carrying Turner, who actually survives. The "Dutchman" then sails to Isla Cruces to stop Sparrow from getting the Chest.


title: Davy Jones (character)

text: Arriving, Jones sends his crew to retrieve the Chest; they return to him with it. The "Dutchman" then chases after the "Black Pearl", but is outrun. Jones summons the Kraken, which drags Jack Sparrow and the "Pearl" to Davy Jones's Locker. He afterward opens the Chest only to find his heart missing; it having been taken by James Norrington, who gives it to Lord Cutler Beckett, the chairman of the East India Trading Company.


title: Davy Jones (character)

text: In the third film "", Jones is under the control of Beckett, under the threat of death. Beckett orders Jones to sink pirate ships, but is infuriated when Jones leaves no survivors; Beckett wants prisoners to interrogate about the Brethren Court. Beckett also orders Jones to kill the Kraken. Later, he orders Jones attack the Pirate Lord Sao Feng; Jones subsequently kills Sao and captures Elizabeth Swann, who had been named captain by Sao Feng upon his death. When Admiral James Norrington dies on board the "Dutchman" while helping Elizabeth escape, Jones claims Norrington's sword (originally crafted by Will Turner). Jones then attempts mutiny against the EITC. However, Mercer successfully defends the Chest, forcing Jones to continue under Beckett's service.


title: Davy Jones (character)

text: Beckett later summons Jones to his ship, the "Endeavour". Jones confronts Will Turner and divulges his past with Calypso while learning of Jack Sparrow's escape from the Locker. The three men then arrive at Shipwreck Cove.


title: Davy Jones (character)

text: Jones confronts Calypso, locked in the brig of the "Black Pearl". The two former lovers discuss Calypso's betrayal and Jones's curse. Calypso temporarily lifts his curse, allowing him to be seen briefly in his original human form. Jones tells her that his heart will always belong to her. Calypso, unaware that Jones betrayed her to the first Brethren Court, says that after her release, she will fully give her love to him.


title: Davy Jones (character)

text: Jones participates in a parley in which the EITC trades Turner for Sparrow with the latter actually planning to stab the heart. After Calypso is freed, Will reveals that Jones betrayed her. She escapes, refusing to aid either the pirates or Jones. Her fury creates a monstrous maelstrom. The "Dutchman" and the "Pearl" enter it and battle.


title: Davy Jones (character)

text: During the battle, Jones suffocates Mercer to death with his tentacles and retrieves the key to the Chest. Sparrow and Jones fight for control of the chest in the rigging of the "Dutchman". Jack acquires both the Chest and the key while Jones battles Will and Elizabeth. Jones quickly overpowers Elizabeth, and is subsequently impaled through the back by Will. Jones, unharmed, holds Will at sword-point. Jack threatens to stab the heart, and Jones cruelly stabs Will. Remembering Will as his son, Bootstrap Bill briefly fights and overpowers Jones, but is quickly defeated. Jones attempts to kill Bootstrap, but Jack helps Will stab the heart. Jones then calls out for Calypso, before tumbling to his death in the maelstrom.


title: Davy Jones (character)

text: In the post-credits scene of the fifth film "", Will (no longer bound to the "Flying Dutchman" after the destruction of the Trident of Poseidon) and Elizabeth are sleeping in their bed together when their room is entered by the silhouette of an apparently resurrected Davy Jones. Just as Will sees Jones raising his clawed arm to strike at the couple, Will awakens and sees that the room is empty. Assuming Jones' appearance to be a nightmare, Will goes back to sleep, oblivious to the presence of barnacles on the floor amid a small puddle of seawater that reveals that Jones is possibly back and it may not be a nightmare.


title: Davy Jones (character)

text: Jones and Calypso possess a matching set of heart-shaped music-box lockets that play a distinct melody (the latter is a half-step above the former's), and Jones is known to play this melody on his pipe organ. The melody is also Jones' theme as heard throughout the films featuring him; it comes in two variations: one heard only in the "" soundtrack, and a simpler variant played in the films on several occasions (including briefly after his appearance in "").


title: Davy Jones (character)

text: Davy Jones is identified as the Devil by his crew: he is cruel, cunning, ruthless, greedy, murderous and careless of other people; these traits make him similar to Cutler Beckett. Evil, proud and possibly nihilistic, he also tried to force Jack Sparrow to leave Will with him and bring him 100 souls knowing that Sparrow is against slavery. Excluding those who manage to reach an agreement with him, Davy Jones presents himself as an extremely formidable enemy, without hesitation or reservation. Unlike the cowardly Beckett, Jones is brave and daring, and he can be very scary and quick-tempered to prove he is serious. It becomes more dangerous when he realizes that he has been cheated: in fact when Jack Sparrow revealed that he had not recruited 100 souls to give to him in exchange for his (not being able to force himself to do a horrible thing), but that he had instead gone to get his heart to blackmail him into leaving him and his crew alone. Jones began shooting at the Black Pearl, trying to sink it again to give Sparrow pain, knowing how much he cares about the ship. Although he does not tolerate the violation of his contracts, he never respects his promises. Probably due to the betrayal of his beloved Calypso, Jones is extremely convinced of the cruelty and unhappiness of life; in fact, his phrase is explanatory of this vision: "Life is cruel! Why should the afterlife be any different?". His passionate nature is rarely shown to others, such as when he plays her theme on the pipe organ as he sheds a tear on Calypso and eventually meets her aboard the Black Pearl. In times of intense emotional distress, Jones uses his organ to vent his anger and suffering. Despite the wickedness, anger and bitterness that characterize him, Davy Jones has a repressed feeling of love and goodness: even after being betrayed by Calypso, Jones still loves her, and her name is in fact the last word that pronounce before dying.


title: Davy Jones (character)

text: Davy Jones speaks with a strong Scottish accent, accentuated by various vocal and facial tics, which make it even more disturbing. His typical laugh is greasy and mocking.


title: Davy Jones (character)

text: Davy Jones possesses a large number of supernatural abilities. Jones is capable of teleportation onboard ships at sea, and he can pass through solid objects. His crew are bound to his every whim, and his enchanted ship, the "Flying Dutchman", is capable of traveling underwater.


title: Davy Jones (character)

text: Jones is immortal, capable of surviving injuries that would be fatal to mortals, though not impervious to pain. In particular, he was able to cut out his heart and leave it disembodied; this soon becomes his only real weakness, as the heart can be possessed by others and used to kill him or leverage him under their control.


title: Davy Jones (character)

text: Jones can also track any soul that is owed to him using the black spot. Any member of his crew can mark a victim with the black spot, but only Jones himself can remove it. Jones also has the power to control and call forth the Kraken, a sea monster that can destroy ships upon command.


title: Davy Jones (character)

text: Jones is unable to step on dry land except once every ten years. However, he can stand and even walk given his path has buckets of seawater for him to place his feet in (as seen during the parley on a sandbar in "At World's End"), and he can also send his crew ashore in his stead.


title: Davy Jones (character)

text: In a physical confrontation, Jones' mutated physiology also gives him various advantages. His facial tentacles allow him to manipulate objects with the dexterity of a cephalopod and with greater versatility, as seen when he masterfully plays his pipe organ. As he uses his tentacles, his non-human hands can be thus left free to accomplish other tasks, such as when he is able to restrain Mercer's arms while creatively smothering him with his tentacles. His tentacle finger allows him to exert a much stronger grip and control his sword more quickly and precisely than a normal hand could, and his crab claw hand possesses enough strength to halt melee attacks and bend or sever sword blades. He also demonstrates more general superhuman strength when he throws Jack off the crossbeam using only his right arm.


title: Davy Jones (character)

text: Davy Jones was a part of Series One of the "Pirates of the Caribbean: Dead Man's Chest" action figure set produced by NECA. Although the initial run of figures had a sticker on the box that proclaimed that the figure came with the Dead Man's Chest and Jones' heart, both props (as well as the key) were released with the Bootstrap Bill figure in Series Two.


title: Davy Jones (character)

text: Many different sized figurines of Jones were also produced by the toy company Zizzle in 2006 and 2007 including an appearance as a smaller figure with crew members Angler, Wheelback and Penrod as well as in a 3 figure pack with Hector Barbossa and a limited edition gold Jack Sparrow for "At World's End".


title: Davy Jones (character)

text: Davy Jones and his ship, the "Flying Dutchman", were produced as a Mega Blocks set for the movies "Dead Man's Chest" and "". Although his Minifigure counterpart in the "Dead Man's Chest" set has more bluish tentacles then his counterpart in the "At World's End" set, which has more greenish tentacles.


title: Davy Jones (character)

text: A Lego minifigure of Jones was released in November 2011 in the "Lego Pirates of the Caribbean" theme.


title: Davy Jones (character)

text: Hot Toys also announced plans to make a 1:6 version of Davy Jones, which became available in 2008. It is widely regarded as more detailed than those produced by NECA.


title: Cerebellum

text: The cerebellum (Latin for "little brain") is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebellum plays an important role in motor control. It may also be involved in some cognitive functions such as attention and language as well as emotional control such as regulating fear and pleasure responses, but its movement-related functions are the most solidly established. The human cerebellum does not initiate movement, but contributes to coordination, precision, and accurate timing: it receives input from sensory systems of the spinal cord and from other parts of the brain, and integrates these inputs to fine-tune motor activity. Cerebellar damage produces disorders in fine movement, equilibrium, posture, and motor learning in humans.


title: Cerebellum

text: Anatomically, the human cerebellum has the appearance of a separate structure attached to the bottom of the brain, tucked underneath the cerebral hemispheres. Its cortical surface is covered with finely spaced parallel grooves, in striking contrast to the broad irregular convolutions of the cerebral cortex. These parallel grooves conceal the fact that the cerebellar cortex is actually a continuous thin layer of tissue tightly folded in the style of an accordion. Within this thin layer are several types of neurons with a highly regular arrangement, the most important being Purkinje cells and granule cells. This complex neural organization gives rise to a massive signal-processing capability, but almost all of the output from the cerebellar cortex passes through a set of small deep nuclei lying in the white matter interior of the cerebellum.


title: Cerebellum

text: In addition to its direct role in motor control, the cerebellum is necessary for several types of motor learning, most notably learning to adjust to changes in sensorimotor relationships. Several theoretical models have been developed to explain sensorimotor calibration in terms of synaptic plasticity within the cerebellum. These models derive from those formulated by David Marr and James Albus, based on the observation that each cerebellar Purkinje cell receives two dramatically different types of input: one comprises thousands of weak inputs from the parallel fibers of the granule cells; the other is an extremely strong input from a single climbing fiber. The basic concept of the Marr–Albus theory is that the climbing fiber serves as a "teaching signal", which induces a long-lasting change in the strength of parallel fiber inputs. Observations of long-term depression in parallel fiber inputs have provided some support for theories of this type, but their validity remains controversial.


title: Cerebellum

text: At the level of gross anatomy, the cerebellum consists of a tightly folded layer of cortex, with white matter underneath and a fluid-filled ventricle at the base. Four deep cerebellar nuclei are embedded in the white matter. Each part of the cortex consists of the same small set of neuronal elements, laid out in a highly stereotyped geometry. At an intermediate level, the cerebellum and its auxiliary structures can be separated into several hundred or thousand independently functioning modules called "microzones" or "microcompartments".


title: Cerebellum

text: The cerebellum is located in the posterior cranial fossa. The fourth ventricle, pons and medulla are in front of the cerebellum. It is separated from the overlying cerebrum by a layer of leathery dura mater, the tentorium cerebelli; all of its connections with other parts of the brain travel through the pons. Anatomists classify the cerebellum as part of the metencephalon, which also includes the pons; the metencephalon is the upper part of the rhombencephalon or "hindbrain". Like the cerebral cortex, the cerebellum is divided into two cerebellar hemispheres; it also contains a narrow midline zone (the vermis). A set of large folds is, by convention, used to divide the overall structure into 10 smaller "lobules". Because of its large number of tiny granule cells, the cerebellum contains more neurons than the total from the rest of the brain, but takes up only 10% of the total brain volume. The number of neurons in the cerebellum is related to the number of neurons in the neocortex. There are about 3.6 times as many neurons in the cerebellum as in the neocortex, a ratio that is conserved across many different mammalian species.


title: Cerebellum

text: The unusual surface appearance of the cerebellum conceals the fact that most of its volume is made up of a very tightly folded layer of gray matter: the cerebellar cortex. Each ridge or gyrus in this layer is called a folium. It is estimated that, if the human cerebellar cortex were completely unfolded, it would give rise to a layer of neural tissue about 1 meter long and averaging 5 centimeters wide—a total surface area of about 500 square cm, packed within a volume of dimensions 6 cm × 5 cm × 10 cm. Underneath the gray matter of the cortex lies white matter, made up largely of myelinated nerve fibers running to and from the cortex. Embedded within the white matter—which is sometimes called the "arbor vitae" (tree of life) because of its branched, tree-like appearance in cross-section—are four deep cerebellar nuclei, composed of gray matter.


title: Cerebellum

text: Connecting the cerebellum to different parts of the nervous system are three paired cerebellar peduncles. These are the superior cerebellar peduncle, the middle cerebellar peduncle and the inferior cerebellar peduncle, named by their position relative to the vermis. The superior cerebellar peduncle is mainly an output to the cerebral cortex, carrying efferent fibers via thalamic nuclei to upper motor neurons in the cerebral cortex. The fibers arise from the deep cerebellar nuclei. The middle cerebellar peduncle is connected to the pons and receives all of its input from the pons mainly from the pontine nuclei. The input to the pons is from the cerebral cortex and is relayed from the pontine nuclei via transverse pontine fibers to the cerebellum. The middle peduncle is the largest of the three and its afferent fibers are grouped into three separate fascicles taking their inputs to different parts of the cerebellum. The inferior cerebellar peduncle receives input from afferent fibers from the vestibular nuclei, spinal cord and the tegmentum. Output from the inferior peduncle is via efferent fibers to the vestibular nuclei and the reticular formation. The whole of the cerebellum receives modulatory input from the inferior olivary nucleus via the inferior cerebellar peduncle.


title: Cerebellum

text: Based on the surface appearance, three lobes can be distinguished within the cerebellum: the anterior lobe (above the primary fissure), the posterior lobe (below the primary fissure), and the flocculonodular lobe (below the posterior fissure). These lobes divide the cerebellum from rostral to caudal (in humans, top to bottom). In terms of function, however, there is a more important distinction along the medial-to-lateral dimension. Leaving out the flocculonodular lobe, which has distinct connections and functions, the cerebellum can be parsed functionally into a medial sector called the spinocerebellum and a larger lateral sector called the cerebrocerebellum. A narrow strip of protruding tissue along the midline is called the cerebellar vermis. ("Vermis" is Latin for "worm".)


title: Cerebellum

text: The smallest region, the flocculonodular lobe, is often called the vestibulocerebellum. It is the oldest part in evolutionary terms (archicerebellum) and participates mainly in balance and spatial orientation; its primary connections are with the vestibular nuclei, although it also receives visual and other sensory input. Damage to this region causes disturbances of balance and gait.


title: Cerebellum

text: The medial zone of the anterior and posterior lobes constitutes the spinocerebellum, also known as paleocerebellum. This sector of the cerebellum functions mainly to fine-tune body and limb movements. It receives proprioceptive input from the dorsal columns of the spinal cord (including the spinocerebellar tract) and from the cranial trigeminal nerve, as well as from visual and auditory systems. It sends fibers to deep cerebellar nuclei that, in turn, project to both the cerebral cortex and the brain stem, thus providing modulation of descending motor systems.


title: Cerebellum

text: The lateral zone, which in humans is by far the largest part, constitutes the cerebrocerebellum, also known as neocerebellum. It receives input exclusively from the cerebral cortex (especially the parietal lobe) via the pontine nuclei (forming cortico-ponto-cerebellar pathways), and sends output mainly to the ventrolateral thalamus (in turn connected to motor areas of the premotor cortex and primary motor area of the cerebral cortex) and to the red nucleus. There is disagreement about the best way to describe the functions of the lateral cerebellum: It is thought to be involved in planning movement that is about to occur, in evaluating sensory information for action, and in a number of purely cognitive functions, such as determining the verb which best fits with a certain noun (as in "sit" for "chair").


title: Cerebellum

text: Two types of neuron play dominant roles in the cerebellar circuit: Purkinje cells and granule cells. Three types of axons also play dominant roles: mossy fibers and climbing fibers (which enter the cerebellum from outside), and parallel fibers (which are the axons of granule cells). There are two main pathways through the cerebellar circuit, originating from mossy fibers and climbing fibers, both eventually terminating in the deep cerebellar nuclei.


title: Cerebellum

text: Mossy fibers project directly to the deep nuclei, but also give rise to the following pathway: mossy fibers → granule cells → parallel fibers → Purkinje cells → deep nuclei. Climbing fibers project to Purkinje cells and also send collaterals directly to the deep nuclei. The mossy fiber and climbing fiber inputs each carry fiber-specific information; the cerebellum also receives dopaminergic, serotonergic, noradrenergic, and cholinergic inputs that presumably perform global modulation.


title: Cerebellum

text: The cerebellar cortex is divided into three layers. At the bottom lies the thick granular layer, densely packed with granule cells, along with interneurons, mainly Golgi cells but also including Lugaro cells and unipolar brush cells. In the middle lies the Purkinje layer, a narrow zone that contains the cell bodies of Purkinje cells and Bergmann glial cells. At the top lies the molecular layer, which contains the flattened dendritic trees of Purkinje cells, along with the huge array of parallel fibers penetrating the Purkinje cell dendritic trees at right angles. This outermost layer of the cerebellar cortex also contains two types of inhibitory interneuron: stellate cells and basket cells. Both stellate and basket cells form GABAergic synapses onto Purkinje cell dendrites.


title: Cerebellum

text: The top, outermost layer of the cerebellar cortex is the molecular layer. This layer contains the flattened dendritic trees of Purkinje cells, and the huge array of parallel fibers, from the granular layer, that penetrate the Purkinje cell dendritic trees at right angles. The molecular layer also contains two types of inhibitory interneuron: stellate cells and basket cells. Both stellate and basket cells form GABAergic synapses onto Purkinje cell dendrites.


title: Cerebellum

text: Purkinje cells are among the most distinctive neurons in the brain, and one of the earliest types to be recognized—they were first described by the Czech anatomist Jan Evangelista Purkyně in 1837. They are distinguished by the shape of their dendritic tree: The dendrites branch very profusely, but are severely flattened in a plane perpendicular to the cerebellar folds. Thus, the dendrites of a Purkinje cell form a dense planar net, through which parallel fibers pass at right angles. The dendrites are covered with dendritic spines, each of which receives synaptic input from a parallel fiber. Purkinje cells receive more synaptic inputs than any other type of cell in the brain—estimates of the number of spines on a single human Purkinje cell run as high as 200,000. The large, spherical cell bodies of Purkinje cells are packed into a narrow layer (one cell thick) of the cerebellar cortex, called the "Purkinje layer". After emitting collaterals that affect nearby parts of the cortex, their axons travel into the deep cerebellar nuclei, where they make on the order of 1,000 contacts each with several types of nuclear cells, all within a small domain. Purkinje cells use GABA as their neurotransmitter, and therefore exert inhibitory effects on their targets.


title: Cerebellum

text: Purkinje cells form the heart of the cerebellar circuit, and their large size and distinctive activity patterns have made it relatively easy to study their response patterns in behaving animals using extracellular recording techniques. Purkinje cells normally emit action potentials at a high rate even in the absence of the synaptic input. In awake, behaving animals, mean rates averaging around 40 Hz are typical. The spike trains show a mixture of what are called simple and complex spikes. A simple spike is a single action potential followed by a refractory period of about 10 ms; a complex spike is a stereotyped sequence of action potentials with very short inter-spike intervals and declining amplitudes. Physiological studies have shown that complex spikes (which occur at baseline rates around 1 Hz and never at rates much higher than 10 Hz) are reliably associated with climbing fiber activation, while simple spikes are produced by a combination of baseline activity and parallel fiber input. Complex spikes are often followed by a pause of several hundred milliseconds during which simple spike activity is suppressed.


title: Cerebellum

text: A specific, recognizable feature of Purkinje neurons is the expression of calbindin. Calbindin staining of rat brain after unilateral chronic sciatic nerve injury suggests that Purkinje neurons may be newly generated in the adult brain, initiating the organization of new cerebellar lobules.


title: Cerebellum

text: Cerebellar granule cells, in contrast to Purkinje cells, are among the smallest neurons in the brain. They are also the most numerous neurons in the brain: In humans, estimates of their total number average around 50 billion, which means that about 3/4 of the brain's neurons are cerebellar granule cells. Their cell bodies are packed into a thick layer at the bottom of the cerebellar cortex. A granule cell emits only four to five dendrites, each of which ends in an enlargement called a "dendritic claw". These enlargements are sites of excitatory input from mossy fibers and inhibitory input from Golgi cells.


title: Cerebellum

text: The thin, unmyelinated axons of granule cells rise vertically to the upper (molecular) layer of the cortex, where they split in two, with each branch traveling horizontally to form a parallel fiber; the splitting of the vertical branch into two horizontal branches gives rise to a distinctive "T" shape. A human parallel fiber runs for an average of 3 mm in each direction from the split, for a total length of about 6 mm (about 1/10 of the total width of the cortical layer). As they run along, the parallel fibers pass through the dendritic trees of Purkinje cells, contacting one of every 3–5 that they pass, making a total of 80–100 synaptic connections with Purkinje cell dendritic spines. Granule cells use glutamate as their neurotransmitter, and therefore exert excitatory effects on their targets.


title: Cerebellum

text: Granule cells receive all of their input from mossy fibers, but outnumber them by 200 to 1 (in humans). Thus, the information in the granule cell population activity state is the same as the information in the mossy fibers, but recoded in a much more expansive way. Because granule cells are so small and so densely packed, it is difficult to record their spike activity in behaving animals, so there is little data to use as a basis for theorizing. The most popular concept of their function was proposed in 1969 by David Marr, who suggested that they could encode combinations of mossy fiber inputs. The idea is that with each granule cell receiving input from only 4–5 mossy fibers, a granule cell would not respond if only a single one of its inputs were active, but would respond if more than one were active. This combinatorial coding scheme would potentially allow the cerebellum to make much finer distinctions between input patterns than the mossy fibers alone would permit.


title: Cerebellum

text: Mossy fibers enter the granular layer from their points of origin, many arising from the pontine nuclei, others from the spinal cord, vestibular nuclei etc. In the human cerebellum, the total number of mossy fibers has been estimated at 200 million. These fibers form excitatory synapses with the granule cells and the cells of the deep cerebellar nuclei. Within the granular layer, a mossy fiber generates a series of enlargements called "rosettes". The contacts between mossy fibers and granule cell dendrites take place within structures called glomeruli. Each glomerulus has a mossy fiber rosette at its center, and up to 20 granule cell dendritic claws contacting it. Terminals from Golgi cells infiltrate the structure and make inhibitory synapses onto the granule cell dendrites. The entire assemblage is surrounded by a sheath of glial cells. Each mossy fiber sends collateral branches to several cerebellar folia, generating a total of 20–30 rosettes; thus a single mossy fiber makes contact with an estimated 400–600 granule cells.


title: Cerebellum

text: Purkinje cells also receive input from the inferior olivary nucleus on the contralateral side of the brainstem via climbing fibers. Although the inferior olive lies in the medulla oblongata and receives input from the spinal cord, brainstem and cerebral cortex, its output goes entirely to the cerebellum. A climbing fiber gives off collaterals to the deep cerebellar nuclei before entering the cerebellar cortex, where it splits into about 10 terminal branches, each of which gives input to a single Purkinje cell. In striking contrast to the 100,000-plus inputs from parallel fibers, each Purkinje cell receives input from exactly one climbing fiber; but this single fiber "climbs" the dendrites of the Purkinje cell, winding around them and making a total of up to 300 synapses as it goes. The net input is so strong that a single action potential from a climbing fiber is capable of producing an extended complex spike in the Purkinje cell: a burst of several spikes in a row, with diminishing amplitude, followed by a pause during which activity is suppressed. The climbing fiber synapses cover the cell body and proximal dendrites; this zone is devoid of parallel fiber inputs.


title: Cerebellum

text: Climbing fibers fire at low rates, but a single climbing fiber action potential induces a burst of several action potentials in a target Purkinje cell (a complex spike). The contrast between parallel fiber and climbing fiber inputs to Purkinje cells (over 100,000 of one type versus exactly one of the other type) is perhaps the most provocative feature of cerebellar anatomy, and has motivated much of the theorizing. In fact, the function of climbing fibers is the most controversial topic concerning the cerebellum. There are two schools of thought, one following Marr and Albus in holding that climbing fiber input serves primarily as a teaching signal, the other holding that its function is to shape cerebellar output directly. Both views have been defended in great length in numerous publications. In the words of one review, "In trying to synthesize the various hypotheses on the function of the climbing fibers, one has the sense of looking at a drawing by Escher. Each point of view seems to account for a certain collection of findings, but when one attempts to put the different views together, a coherent picture of what the climbing fibers are doing does not appear. For the majority of researchers, the climbing fibers signal errors in motor performance, either in the usual manner of discharge frequency modulation or as a single announcement of an 'unexpected event'. For other investigators, the message lies in the degree of ensemble synchrony and rhythmicity among a population of climbing fibers."


title: Cerebellum

text: The deep nuclei of the cerebellum are clusters of gray matter lying within the white matter at the core of the cerebellum. They are, with the minor exception of the nearby vestibular nuclei, the sole sources of output from the cerebellum. These nuclei receive collateral projections from mossy fibers and climbing fibers as well as inhibitory input from the Purkinje cells of the cerebellar cortex. The four nuclei (dentate, globose, emboliform, and fastigial) each communicate with different parts of the brain and cerebellar cortex. (The globose and the emboliform nuclei are also referred to as combined in the interposed nucleus). The fastigial and interposed nuclei belong to the spinocerebellum. The dentate nucleus, which in mammals is much larger than the others, is formed as a thin, convoluted layer of gray matter, and communicates exclusively with the lateral parts of the cerebellar cortex. The flocculus of the flocculonodular lobe is the only part of the cerebellar cortex that does not project to the deep nuclei—its output goes to the vestibular nuclei instead.


title: Cerebellum

text: The majority of neurons in the deep nuclei have large cell bodies and spherical dendritic trees with a radius of about 400 μm, and use glutamate as their neurotransmitter. These cells project to a variety of targets outside the cerebellum. Intermixed with them are a lesser number of small cells, which use GABA as a neurotransmitter and project exclusively to the inferior olivary nucleus, the source of climbing fibers. Thus, the nucleo-olivary projection provides an inhibitory feedback to match the excitatory projection of climbing fibers to the nuclei. There is evidence that each small cluster of nuclear cells projects to the same cluster of olivary cells that send climbing fibers to it; there is strong and matching topography in both directions.


title: Cerebellum

text: When a Purkinje cell axon enters one of the deep nuclei, it branches to make contact with both large and small nuclear cells, but the total number of cells contacted is only about 35 (in cats). Conversely, a single deep nuclear cell receives input from approximately 860 Purkinje cells (again in cats).


title: Cerebellum

text: From the viewpoint of gross anatomy, the cerebellar cortex appears to be a homogeneous sheet of tissue, and, from the viewpoint of microanatomy, all parts of this sheet appear to have the same internal structure. There are, however, a number of respects in which the structure of the cerebellum is compartmentalized. There are large compartments that are generally known as "zones"; these can be divided into smaller compartments known as "microzones".


title: Cerebellum

text: The first indications of compartmental structure came from studies of the receptive fields of cells in various parts of the cerebellar cortex. Each body part maps to specific points in the cerebellum, but there are numerous repetitions of the basic map, forming an arrangement that has been called "fractured somatotopy". A clearer indication of compartmentalization is obtained by immunostaining the cerebellum for certain types of protein. The best-known of these markers are called "zebrins", because staining for them gives rise to a complex pattern reminiscent of the stripes on a zebra. The stripes generated by zebrins and other compartmentalization markers are oriented perpendicular to the cerebellar folds—that is, they are narrow in the mediolateral direction, but much more extended in the longitudinal direction. Different markers generate different sets of stripes, the widths and lengths vary as a function of location, but they all have the same general shape.


title: Cerebellum

text: Oscarsson in the late 1970s proposed that these cortical zones can be partitioned into smaller units called microzones. A microzone is defined as a group of Purkinje cells all having the same somatotopic receptive field. Microzones were found to contain on the order of 1000 Purkinje cells each, arranged in a long, narrow strip, oriented perpendicular to the cortical folds. Thus, as the adjoining diagram illustrates, Purkinje cell dendrites are flattened in the same direction as the microzones extend, while parallel fibers cross them at right angles.


title: Cerebellum

text: It is not only receptive fields that define the microzone structure: The climbing fiber input from the inferior olivary nucleus is equally important. The branches of a climbing fiber (usually numbering about 10) usually activate Purkinje cells belonging to the same microzone. Moreover, olivary neurons that send climbing fibers to the same microzone tend to be coupled by gap junctions, which synchronize their activity, causing Purkinje cells within a microzone to show correlated complex spike activity on a millisecond time scale. Also, the Purkinje cells belonging to a microzone all send their axons to the same small cluster of output cells within the deep cerebellar nuclei. Finally, the axons of basket cells are much longer in the longitudinal direction than in the mediolateral direction, causing them to be confined largely to a single microzone. The consequence of all this structure is that cellular interactions within a microzone are much stronger than interactions between different microzones.


title: Cerebellum

text: In 2005, Richard Apps and Martin Garwicz summarized evidence that microzones themselves form part of a larger entity they call a multizonal microcomplex. Such a microcomplex includes several spatially separated cortical microzones, all of which project to the same group of deep cerebellar neurons, plus a group of coupled olivary neurons that project to all of the included microzones as well as to the deep nuclear area.


title: Cerebellum

text: The cerebellum is provided with blood from three paired major arteries: the superior cerebellar artery (SCA), the anterior inferior cerebellar artery (AICA), and the posterior inferior cerebellar artery (PICA). The SCA supplies the upper region of the cerebellum. It divides at the upper surface and branches into the pia mater where the branches anastomose with those of the anterior and posterior inferior cerebellar arteries. The AICA supplies the front part of the undersurface of the cerebellum. The PICA arrives at the undersurface, where it divides into a medial branch and a lateral branch. The medial branch continues backward to the cerebellar notch between the two hemispheres of the cerebellum; while the lateral branch supplies the under surface of the cerebellum, as far as its lateral border, where it anastomoses with the AICA and the SCA.


title: Cerebellum

text: The strongest clues to the function of the cerebellum have come from examining the consequences of damage to it. Animals and humans with cerebellar dysfunction show, above all, problems with motor control, on the same side of the body as the damaged part of the cerebellum. They continue to be able to generate motor activity but lose precision, producing erratic, uncoordinated, or incorrectly timed movements. A standard test of cerebellar function is to reach with the tip of the finger for a target at arm's length: A healthy person will move the fingertip in a rapid straight trajectory, whereas a person with cerebellar damage will reach slowly and erratically, with many mid-course corrections. Deficits in non-motor functions are more difficult to detect. Thus, the general conclusion reached decades ago is that the basic function of the cerebellum is to calibrate the detailed form of a movement, not to initiate movements or to decide which movements to execute.


title: Cerebellum

text: Prior to the 1990s the function of the cerebellum was almost universally believed to be purely motor-related, but newer findings have brought that view into question. Functional imaging studies have shown cerebellar activation in relation to language, attention, and mental imagery; correlation studies have shown interactions between the cerebellum and non-motor areas of the cerebral cortex; and a variety of non-motor symptoms have been recognized in people with damage that appears to be confined to the cerebellum. In particular, the cerebellar cognitive affective syndrome or Schmahmann's syndrome has been described in adults and children. Estimates based on functional mapping of the cerebellum using functional MRI suggest that more than half of the cerebellar cortex is interconnected with association zones of the cerebral cortex.


title: Cerebellum

text: Kenji Doya has argued that the cerebellum's function is best understood not in terms of the behaviors it affects, but the neural computations it performs; the cerebellum consists of a large number of more or less independent modules, all with the same geometrically regular internal structure, and therefore all, it is presumed, performing the same computation. If the input and output connections of a module are with motor areas (as many are), then the module will be involved in motor behavior; but, if the connections are with areas involved in non-motor cognition, the module will show other types of behavioral correlates. Thus the cerebellum has been implicated in the regulation of many differing functional traits such as affection, emotion including emotional body language perception and behavior. The cerebellum, Doya proposes, is best understood as predictive action selection based on "internal models" of the environment or a device for supervised learning, in contrast to the basal ganglia, which perform reinforcement learning, and the cerebral cortex, which performs unsupervised learning. Three decades of brain research have led to the proposal that the cerebellum generates optimized mental models and interacts closely with the cerebral cortex, where updated internal models are experienced as creative intuition ("a ha") in working memory.


title: Cerebellum

text: The comparative simplicity and regularity of the cerebellar anatomy led to an early hope that it might imply a similar simplicity of computational function, as expressed in one of the first books on cerebellar electrophysiology, "The Cerebellum as a Neuronal Machine" by John C. Eccles, Masao Ito, and János Szentágothai. Although a full understanding of cerebellar function has remained elusive, at least four principles have been identified as important: (1) feedforward processing, (2) divergence and convergence, (3) modularity, and (4) plasticity.


title: Cerebellum

text: There is considerable evidence that the cerebellum plays an essential role in some types of motor learning. The tasks where the cerebellum most clearly comes into play are those in which it is necessary to make fine adjustments to the way an action is performed. There has, however, been much dispute about whether learning takes place within the cerebellum itself, or whether it merely serves to provide signals that promote learning in other brain structures. Most theories that assign learning to the circuitry of the cerebellum are derived from the ideas of David Marr and James Albus, who postulated that climbing fibers provide a teaching signal that induces synaptic modification in parallel fiber–Purkinje cell synapses. Marr assumed that climbing fiber input would cause synchronously activated parallel fiber inputs to be strengthened. Most subsequent cerebellar-learning models, however, have followed Albus in assuming that climbing fiber activity would be an error signal, and would cause synchronously activated parallel fiber inputs to be weakened. Some of these later models, such as the "Adaptive Filter" model of Fujita made attempts to understand cerebellar function in terms of optimal control theory.


title: Cerebellum

text: The idea that climbing fiber activity functions as an error signal has been examined in many experimental studies, with some supporting it but others casting doubt. In a pioneering study by Gilbert and Thach from 1977, Purkinje cells from monkeys learning a reaching task showed increased complex spike activity—which is known to reliably indicate activity of the cell's climbing fiber input—during periods when performance was poor. Several studies of motor learning in cats observed complex spike activity when there was a mismatch between an intended movement and the movement that was actually executed. Studies of the vestibulo–ocular reflex (which stabilizes the visual image on the retina when the head turns) found that climbing fiber activity indicated "retinal slip", although not in a very straightforward way.


title: Cerebellum

text: One of the most extensively studied cerebellar learning tasks is the eyeblink conditioning paradigm, in which a neutral conditioned stimulus (CS) such as a tone or a light is repeatedly paired with an unconditioned stimulus (US), such as an air puff, that elicits a blink response. After such repeated presentations of the CS and US, the CS will eventually elicit a blink before the US, a conditioned response or CR. Experiments showed that lesions localized either to a specific part of the interposed nucleus (one of the deep cerebellar nuclei) or to a few specific points in the cerebellar cortex would abolish learning of a conditionally timed blink response. If cerebellar outputs are pharmacologically inactivated while leaving the inputs and intracellular circuits intact, learning takes place even while the animal fails to show any response, whereas, if intracerebellar circuits are disrupted, no learning takes place—these facts taken together make a strong case that the learning, indeed, occurs inside the cerebellum.


title: Cerebellum

text: The large base of knowledge about the anatomical structure and behavioral functions of the cerebellum have made it a fertile ground for theorizing—there are perhaps more theories of the function of the cerebellum than of any other part of the brain. The most basic distinction among them is between "learning theories" and "performance theories"—that is, theories that make use of synaptic plasticity within the cerebellum to account for its role in learning, versus theories that account for aspects of ongoing behavior on the basis of cerebellar signal processing. Several theories of both types have been formulated as mathematical models and simulated using computers.


title: Cerebellum

text: Perhaps the earliest "performance" theory was the "delay line" hypothesis of Valentino Braitenberg. The original theory put forth by Braitenberg and Roger Atwood in 1958 proposed that slow propagation of signals along parallel fibers imposes predictable delays that allow the cerebellum to detect time relationships within a certain window. Experimental data did not support the original form of the theory, but Braitenberg continued to argue for modified versions. The hypothesis that the cerebellum functions essentially as a timing system has also been advocated by Richard Ivry. Another influential "performance" theory is the Tensor network theory of Pellionisz and Llinás, which provided an advanced mathematical formulation of the idea that the fundamental computation performed by the cerebellum is to transform sensory into motor coordinates.


title: Cerebellum

text: Theories in the "learning" category almost all derive from publications by Marr and Albus. Marr's 1969 paper proposed that the cerebellum is a device for learning to associate elemental movements encoded by climbing fibers with mossy fiber inputs that encode the sensory context. Albus proposed in 1971 that a cerebellar Purkinje cell functions as a perceptron, a neurally inspired abstract learning device. The most basic difference between the Marr and Albus theories is that Marr assumed that climbing fiber activity would cause parallel fiber synapses to be strengthened, whereas Albus proposed that they would be weakened. Albus also formulated his version as a software algorithm he called a CMAC (Cerebellar Model Articulation Controller), which has been tested in a number of applications.


title: Cerebellum

text: Damage to the cerebellum often causes motor-related symptoms, the details of which depend on the part of the cerebellum involved and how it is damaged. Damage to the flocculonodular lobe may show up as a loss of equilibrium and in particular an altered, irregular walking gait, with a wide stance caused by difficulty in balancing. Damage to the lateral zone typically causes problems in skilled voluntary and planned movements which can cause errors in the force, direction, speed and amplitude of movements. Other manifestations include hypotonia (decreased muscle tone), dysarthria (problems with speech articulation), dysmetria (problems judging distances or ranges of movement), dysdiadochokinesia (inability to perform rapid alternating movements such as walking), impaired check reflex or rebound phenomenon, and intention tremor (involuntary movement caused by alternating contractions of opposing muscle groups). Damage to the midline portion may disrupt whole-body movements, whereas damage localized more laterally is more likely to disrupt fine movements of the hands or limbs. Damage to the upper part of the cerebellum tends to cause gait impairments and other problems with leg coordination; damage to the lower part is more likely to cause uncoordinated or poorly aimed movements of the arms and hands, as well as difficulties in speed. This complex of motor symptoms is called "ataxia".


title: Cerebellum

text: To identify cerebellar problems, neurological examination includes assessment of gait (a broad-based gait being indicative of ataxia), finger-pointing tests and assessment of posture. If cerebellar dysfunction is indicated, a magnetic resonance imaging scan can be used to obtain a detailed picture of any structural alterations that may exist.


title: Cerebellum

text: The list of medical problems that can produce cerebellar damage is long, including stroke, hemorrhage, swelling of the brain (cerebral edema), tumors, alcoholism, physical trauma such as gunshot wounds or explosives, and chronic degenerative conditions such as olivopontocerebellar atrophy. Some forms of migraine headache may also produce temporary dysfunction of the cerebellum, of variable severity. Infection can result in cerebellar damage in such conditions as the prion diseases and Miller Fisher syndrome, a variant of Guillain–Barré syndrome.


title: Cerebellum

text: The human cerebellum changes with age. These changes may differ from those of other parts of the brain.


title: Cerebellum

text: The cerebellum is the youngest brain region (and body part) in centenarians according to an epigenetic biomarker of tissue age known as epigenetic clock: it is about 15 years younger than expected in a centenarian. Further, gene expression patterns in the human cerebellum show less age-related alteration than that in the cerebral cortex.


title: Cerebellum

text: Some studies have reported reductions in numbers of cells or volume of tissue, but the amount of data relating to this question is not very large.


title: Cerebellum

text: Congenital malformation, hereditary disorders, and acquired conditions can affect cerebellar structure and, consequently, cerebellar function. Unless the causative condition is reversible, the only possible treatment is to help people live with their problems. Visualization of the fetal cerebellum by ultrasound scan at 18 to 20 weeks of pregnancy can be used to screen for fetal neural tube defects with a sensitivity rate of up to 99%.


title: Cerebellum

text: In normal development, endogenous sonic hedgehog signaling stimulates rapid proliferation of cerebellar granule neuron progenitors (CGNPs) in the external granule layer (EGL). Cerebellar development occurs during late embryogenesis and the early postnatal period, with CGNP proliferation in the EGL peaking during early development (postnatal day 7 in the mouse). As CGNPs terminally differentiate into cerebellar granule cells (also called cerebellar granule neurons, CGNs), they migrate to the internal granule layer (IGL), forming the mature cerebellum (by post-natal day 20 in the mouse). Mutations that abnormally activate Sonic hedgehog signaling predispose to cancer of the cerebellum (medulloblastoma) in humans with Gorlin Syndrome and in genetically engineered mouse models.


title: Cerebellum

text: Congenital malformation or underdevelopment (hypoplasia) of the cerebellar vermis is a characteristic of both Dandy–Walker syndrome and Joubert syndrome. In very rare cases, the entire cerebellum may be absent. The inherited neurological disorders Machado–Joseph disease, ataxia telangiectasia, and Friedreich's ataxia cause progressive neurodegeneration linked to cerebellar loss. Congenital brain malformations outside the cerebellum can, in turn, cause herniation of cerebellar tissue, as seen in some forms of Arnold–Chiari malformation.


title: Cerebellum

text: Other conditions that are closely linked to cerebellar degeneration include the idiopathic progressive neurological disorders multiple system atrophy and Ramsay Hunt syndrome type I, and the autoimmune disorder paraneoplastic cerebellar degeneration, in which tumors elsewhere in the body elicit an autoimmune response that causes neuronal loss in the cerebellum. Cerebellar atrophy can result from an acute deficiency of vitamin B1 (thiamine) as seen in beriberi and in Wernicke–Korsakoff syndrome, or vitamin E deficiency.


title: Cerebellum

text: Cerebellar atrophy has been observed in many other neurological disorders including Huntington's disease, multiple sclerosis, essential tremor, progressive myoclonus epilepsy, and Niemann–Pick disease. Cerebellar atrophy can also occur as a result of exposure to toxins including heavy metals or pharmaceutical or recreational drugs.


title: Cerebellum

text: There is a general consensus that the cerebellum is involved in pain processing. The cerebellum receives pain input from both descending cortico-cerebellar pathways and ascending spino-cerebellar pathways, through the pontine nuclei and inferior olives. Some of this information is transferred to the motor system inducing a conscious motor avoidance of pain, graded according to pain intensity.


title: Cerebellum

text: These direct pain inputs, as well as indirect inputs, are thought to induce long-term pain avoidance behavior that results in chronic posture changes and consequently, in functional and anatomical remodeling of vestibular and proprioceptive nuclei. As a result, chronic neuropathic pain can induce macroscopic anatomical remodeling of the hindbrain, including the cerebellum. The magnitude of this remodeling and the induction of neuron progenitor markers suggest the contribution of adult neurogenesis to these changes.


title: Cerebellum

text: The circuits in the cerebellum are similar across all classes of vertebrates, including fish, reptiles, birds, and mammals. There is also an analogous brain structure in cephalopods with well-developed brains, such as octopuses. This has been taken as evidence that the cerebellum performs functions important to all animal species with a brain.


title: Cerebellum

text: There is considerable variation in the size and shape of the cerebellum in different vertebrate species. In amphibians, it is little developed, and in lampreys, and hagfish, the cerebellum is barely distinguishable from the brain-stem. Although the spinocerebellum is present in these groups, the primary structures are small, paired-nuclei corresponding to the vestibulocerebellum. The cerebellum is a bit larger in reptiles, considerably larger in birds, and larger yet in mammals. The large paired and convoluted lobes found in humans are typical of mammals, but the cerebellum is, in general, a single median lobe in other groups, and is either smooth or only slightly grooved. In mammals, the neocerebellum is the major part of the cerebellum by mass, but, in other vertebrates, it is typically the spinocerebellum.


title: Cerebellum

text: The cerebellum of cartilaginous and bony fishes is extraordinarily large and complex. In at least one important respect, it differs in internal structure from the mammalian cerebellum: The fish cerebellum does not contain discrete deep cerebellar nuclei. Instead, the primary targets of Purkinje cells are a distinct type of cell distributed across the cerebellar cortex, a type not seen in mammals. In mormyrid fish (a family of weakly electrosensitive freshwater fish), the cerebellum is considerably larger than the rest of the brain. The largest part of it is a special structure called the "valvula", which has an unusually regular architecture and receives much of its input from the electrosensory system.


title: Cerebellum

text: The hallmark of the mammalian cerebellum is an expansion of the lateral lobes, whose main interactions are with the neocortex. As monkeys evolved into great apes, the expansion of the lateral lobes continued, in tandem with the expansion of the frontal lobes of the neocortex. In ancestral hominids, and in "Homo sapiens" until the middle Pleistocene period, the cerebellum continued to expand, but the frontal lobes expanded more rapidly. The most recent period of human evolution, however, may actually have been associated with an increase in the relative size of the cerebellum, as the neocortex reduced its size somewhat while the cerebellum expanded. The size of the human cerebellum, compared to the rest of the brain, has been increasing in size while the cerebrum decreased in size With both the development and implementation of motor tasks, visual-spatial skills and learning taking place in the cerebellum, the growth of the cerebellum is thought to have some form of correlation to greater human cognitive abilities. The lateral hemispheres of the cerebellum are now 2.7 times greater in both humans and apes than they are in monkeys. These changes in the cerebellum size cannot be explained by greater muscle mass. They show that either the development of the cerebellum is tightly linked to that of the rest of the brain or that neural activities taking place in the cerebellum were important during Hominidae evolution. Due to the cerebellum's role in cognitive functions, the increase in its size may have played a role in cognitive expansion.


title: Cerebellum

text: Most vertebrate species have a cerebellum and one or more cerebellum-like structures, brain areas that resemble the cerebellum in terms of cytoarchitecture and neurochemistry. The only cerebellum-like structure found in mammals is the dorsal cochlear nucleus (DCN), one of the two primary sensory nuclei that receive input directly from the auditory nerve. The DCN is a layered structure, with the bottom layer containing granule cells similar to those of the cerebellum, giving rise to parallel fibers that rise to the superficial layer and travel across it horizontally. The superficial layer contains a set of GABAergic neurons called cartwheel cells that resemble Purkinje cells anatomically and chemically—they receive parallel fiber input, but do not have any inputs that resemble climbing fibers. The output neurons of the DCN are pyramidal cells. They are glutamatergic, but also resemble Purkinje cells in some respects—they have spiny, flattened superficial dendritic trees that receive parallel fiber input, but they also have basal dendrites that receive input from auditory nerve fibers, which travel across the DCN in a direction at right angles to the parallel fibers. The DCN is most highly developed in rodents and other small animals, and is considerably reduced in primates. Its function is not well understood; the most popular speculations relate it to spatial hearing in one way or another.


title: Cerebellum

text: Most species of fish and amphibians possess a lateral line system that senses pressure waves in water. One of the brain areas that receives primary input from the lateral line organ, the medial octavolateral nucleus, has a cerebellum-like structure, with granule cells and parallel fibers. In electrosensitive fish, the input from the electrosensory system goes to the dorsal octavolateral nucleus, which also has a cerebellum-like structure. In ray-finned fishes (by far the largest group), the optic tectum has a layer—the marginal layer—that is cerebellum-like.


title: Cerebellum

text: All of these cerebellum-like structures appear to be primarily sensory-related rather than motor-related. All of them have granule cells that give rise to parallel fibers that connect to Purkinje-like neurons with modifiable synapses, but none have climbing fibers comparable to those of the cerebellum—instead they receive direct input from peripheral sensory organs. None has a demonstrated function, but the most influential speculation is that they serve to transform sensory inputs in some sophisticated way, perhaps to compensate for changes in body posture. In fact, James M. Bower and others have argued, partly on the basis of these structures and partly on the basis of cerebellar studies, that the cerebellum itself is fundamentally a sensory structure, and that it contributes to motor control by moving the body in a way that controls the resulting sensory signals. Despite Bower's viewpoint, there is also strong evidence that the cerebellum directly influences motor output in mammals.


title: Cerebellum

text: Even the earliest anatomists were able to recognize the cerebellum by its distinctive appearance. Aristotle and Herophilus (quoted in Galen) called it the παρεγκεφαλίς ("parenkephalis"), as opposed to the ἐγκέφαλος ("enkephalos") or brain proper. Galen's extensive description is the earliest that survives. He speculated that the cerebellum was the source of motor nerves.


title: Cerebellum

text: Further significant developments did not come until the Renaissance. Vesalius discussed the cerebellum briefly, and the anatomy was described more thoroughly by Thomas Willis in 1664. More anatomical work was done during the 18th century, but it was not until early in the 19th century that the first insights into the function of the cerebellum were obtained. Luigi Rolando in 1809 established the key finding that damage to the cerebellum results in motor disturbances. Jean Pierre Flourens in the first half of the 19th century carried out detailed experimental work, which revealed that animals with cerebellar damage can still move, but with a loss of coordination (strange movements, awkward gait, and muscular weakness), and that recovery after the lesion can be nearly complete unless the lesion is very extensive. By the beginning of the 20th century, it was widely accepted that the primary function of the cerebellum relates to motor control; the first half of the 20th century produced several detailed descriptions of the clinical symptoms associated with cerebellar disease in humans.


title: Cerebellum

text: The name "cerebellum" is a diminutive of "cerebrum" (brain); it can be translated literally as "little brain". The Latin name is a direct translation of the Ancient Greek παρεγκεφαλίς ("parenkephalis"), which was used in the works of Aristotle, the first known writer to describe the structure. No other name is used in the English-language literature, but historically a variety of Greek or Latin-derived names have been used, including "cerebrum parvum", "encephalion", "encranion", "cerebrum posterius", and "parencephalis".


title: Scott Wilson (actor)

text: Scott Wilson (born William Delano Wilson; March 29, 1942 – October 6, 2018) was an American actor. He had more than 50 film credits, including "In the Heat of the Night", "In Cold Blood", "The Great Gatsby", "Dead Man Walking", "Pearl Harbor", and "Junebug". In 1980, Wilson received a Golden Globe nomination for Best Supporting Actor – Motion Picture for his role in William Peter Blatty's "The Ninth Configuration". He played veterinarian Hershel Greene on the AMC television series "The Walking Dead" (2011–2014; 2018). He also had a recurring role on "" as casino mogul Sam Braun, as well as a lead role on the Netflix series "The OA" as Abel Johnson.


title: Scott Wilson (actor)

text: Wilson was born in the small Southern town of Thomasville, Georgia. He made his screen debut portraying characters suspected of murder in his first three films. In his debut film, Wilson played a murder suspect in "In the Heat of the Night" (1967). Wilson appeared in Sparta, Mississippi where ("In the Heat of the Night") takes place, on March 15, 2014, to celebrate the city's 175th anniversary in reference to his debut appearance in the film.


title: Scott Wilson (actor)

text: His follow-up role, in the same year, was in the film version of "In Cold Blood", based on the book of the same name by Truman Capote. Wilson portrayed real-life murderer Richard Hickock, while Robert Blake played his partner, Perry Smith.


title: Scott Wilson (actor)

text: Director Richard Brooks cast Wilson and Blake in the starring roles specifically because they were unknown at the time. The director passed over better-known actors, including Steve McQueen and Paul Newman, for the parts. Wilson later explained Brooks' casting motivations: "Brooks hired two 'unknowns' and he wanted to keep it that way. We were treated like two killers he had somehow run across."


title: Scott Wilson (actor)

text: The film earned Wilson an appearance on the cover of "Life" magazine, published on May 12, 1967. Wilson was just 25 years old at the time. The cover features Truman Capote standing between Wilson and Blake on an empty highway in Kansas. The caption, "Nightmare Revisited", appears with them on the cover. Wilson appeared in "The Great Gatsby" in 1974 opposite Robert Redford. He received a 1980 Golden Globe nomination for Best Supporting Actor for his performance in "The Ninth Configuration" by director and friend William Peter Blatty. He lost the Golden Globe to Timothy Hutton. In 1995 Wilson got attention for his role as a prison chaplain in "Dead Man Walking", starring Susan Sarandon and Sean Penn, based on the book of the same name by Sister Helen Prejean.


title: Scott Wilson (actor)

text: Wilson's long filmography also includes "The Gypsy Moths", "The Right Stuff", "A Year of the Quiet Sun", "Malone", "The Grass Harp", "Junebug", "The Host", "Monster", "Young Guns II", "Pearl Harbor", "Big Stan", "Judge Dredd", the "Shiloh" film series and "". Wilson has filmed on location in South Korea, Japan and Spain. Wilson had a recurring role in several episodes of "" as Sam Braun, father of crime-scene investigator Catherine Willows (portrayed by Marg Helgenberger). Braun was killed off in the episode "Built to Kill, Part 2". In the fall of 2011 he also made an appearance opposite Laura Dern in the HBO series, "Enlightened".


title: Scott Wilson (actor)

text: Wilson was cast as veterinarian, Hershel Greene in the second season of "The Walking Dead" in June 2011. The role has earned him positive reviews, including a "cheer" from "TV Guide", which wrote that he had contributed "subtle shades of humanity to the character of Hershel Greene." Wilson was offered the opportunity to join the show while visiting his 97-year-old mother in Georgia. He has described his mother as "a fan of the show." Wilson left the show in December 2013 after his character was killed off in the season 4 mid-season finale "Too Far Gone". However, he made two guest appearances since his character's death, and made his last onscreen appearance in the fifth episode of the ninth season of "The Walking Dead".


title: Scott Wilson (actor)

text: Wilson reflected on his career in a 2011 interview with "Access Atlanta"'s Rodney Ho: "It's been up and down. It's always been. You have dry spells. At different times, you are starting over. If you love it, you stay with it. That's what I'm doing. I've accomplished more than I would have hoped to have accomplished. I don't want to be a big movie star. I can be someone who walks the streets and not get mobbed. I want to be as fine an actor as I can be. I am still striving to be as good as I can be." Wilson was filming scenes for "The Walking Dead" in Senoia, Georgia, at the time the interview took place. In 2014, Wilson was cast in a recurring role as Dr. Guyot in the Amazon original series "Bosch". In 2016, he appeared in the Netflix series "The OA".


title: Scott Wilson (actor)

text: On October 6, 2018, Wilson died from leukemia at his home in Los Angeles, aged 76. He died just a few hours after NYCC (New York Comic Con). The premiere episode of "The Walking Dead"s ninth season, "A New Beginning", which aired the next day, was dedicated to Wilson's memory which is shown in the final credits of the episode.


title: Susan Collins

text: Susan Margaret Collins (born December 7, 1952) is an American politician serving as the senior United States senator from Maine. A member of the Republican Party, she has held her seat since 1997 and is Maine's longest-serving member of Congress.


title: Susan Collins

text: Born in Caribou, Maine, Collins is a graduate of St. Lawrence University in Canton, New York. Beginning her career as a staff assistant for Senator William Cohen in 1975, she became staff director of the Oversight of Government Management Subcommittee of the Committee on Governmental Affairs (which later became the Committee on Homeland Security and Governmental Affairs) in 1981. Governor John R. McKernan Jr. then appointed her commissioner of the Maine Department of Professional and Financial Regulation in 1987. In 1992 President George H. W. Bush appointed her director of the Small Business Administration's regional office in Boston. Collins became a deputy state treasurer in the office of the Treasurer and Receiver-General of Massachusetts in 1993. After moving back to Maine in 1994, she became the Republican nominee for governor of Maine in the 1994 general election. She was the first female major-party nominee for the post, finishing third in a four-way race with 23% of the vote. After her bid for governor in 1994, she became the founding director of the Center for Family Business at Husson University in Bangor, Maine.


title: Susan Collins

text: Collins was first elected to the Senate in 1996. She was reelected in 2002, in 2008, in 2014, and in 2020. She chaired the Senate Special Committee on Aging from 2015 to 2021 and the Senate Committee on Homeland Security and Governmental Affairs from 2003 to 2007. Collins is a senior Republican woman in the Senate, the dean of Maine's congressional delegation, and the only New England Republican in the 116th, 117th Congress, and 118th Congresses. She has been called a moderate Republican and is often a pivotal vote in the Senate. To date, Collins is the longest-serving Republican woman in the Senate.


title: Susan Collins

text: Collins, who has described herself as a pro-choice Republican, attracted controversy for her vote to confirm Brett Kavanaugh to the Supreme Court. She attributed her vote to her belief that Kavanaugh would not support overturning "Roe v. Wade"; in June 2022, Kavanaugh joined the majority opinion in "Dobbs v. Jackson Women's Health Organization", which overturned "Roe".


title: Susan Collins

text: During the impeachment trial of Bill Clinton, Collins was one of 10 Republican senators to vote to acquit him on the first charge and one of five to vote to acquit on the second, and during the first impeachment trial of Donald Trump, Collins joined all Senate Republicans but one in voting to acquit him of all charges. On February 13, 2021, she was one of seven Republican senators to vote to convict Trump of incitement of insurrection in his second impeachment trial. She was the only Republican senator to vote against confirming Amy Coney Barrett to the Supreme Court. Upon the swearing-in of the 118th Congress, Collins will be the only Republican to represent a Northeastern state in the Senate. Furthermore, since the commencement of the 116th Congress, Collins has been the only Republican from New England in either house of Congress.


title: Susan Collins

text: One of six children, Collins was born in Caribou, Maine, where her family operates a lumber business established by her great-great-great-grandfather, Samuel W. Collins, in 1844. Her parents, Patricia (née McGuigan) and Donald F. Collins (1925–2018), each served as mayor of Caribou. Her father, a decorated veteran of World War II, also served in the Maine Legislature, with one term in the House, and four in the Senate.


title: Susan Collins

text: Collins's mother was born in Barrancabermeja, Colombia, to American parents. Collins is of English and Irish ancestry. Her uncle, Samuel W. Collins Jr., sat on the Maine Supreme Judicial Court from 1988 to 1994 and served in the Maine Senate from 1973 to 1984.


title: Susan Collins

text: Collins attended Caribou High School, where she was president of the student council. During her senior year of high school in 1971, she was chosen to participate in the U.S. Senate Youth Program, through which she visited Washington, D.C., for the first time and had a two-hour conversation with Maine's first female United States Senator, Margaret Chase Smith, also a Republican. Collins is the first program delegate elected to the Senate and holds the seat once held by Smith. After graduating from high school, she continued her education at St. Lawrence University in Canton, New York. Like her father, she was elected to the Phi Beta Kappa national academic honor society. She graduated from St. Lawrence "magna cum laude" with a bachelor's degree in government in 1975.


title: Susan Collins

text: Following graduation, Collins worked as a legislative assistant to U.S. Representative and later U.S. Senator William Cohen from 1975 to 1987. She was also staff director of the Oversight of Government Management Subcommittee on the United States Senate Committee on Homeland Security and Governmental Affairs from 1981 to 1987.


title: Susan Collins

text: In 1987, Collins joined the cabinet of Governor John R. McKernan Jr. as Commissioner of the Department of Professional and Financial Regulation. President George H. W. Bush appointed her the New England regional director for the Small Business Administration in 1992. After briefly serving in this post until the 1992 election of President Bill Clinton, she moved to Massachusetts and became Deputy State Treasurer of Massachusetts under Joe Malone in 1993.


title: Susan Collins

text: Returning to Maine, Collins won an eight-way Republican primary in the 1994 gubernatorial election, becoming the first woman nominated by a major party for governor of Maine. During the campaign, she received little support from Republican leaders and was criticized by conservative groups for her more liberal views on social issues. She lost the general election, receiving 23% of the vote and placing third behind Democrat Joseph E. Brennan and the winner, Independent candidate Angus King, her future Senate colleague.


title: Susan Collins

text: In December 1994, Collins became the founding executive director of the Richard E. Dyke Center for Family Business at Husson College. She served in this post until 1996, when she announced her candidacy for the U.S. Senate seat being vacated by her former boss, William Cohen, who retired to become United States Secretary of Defense under President Clinton. With Cohen's public endorsement, she won a difficult four-way primary and faced Brennan, her opponent in the 1994 gubernatorial election, in the general election. She defeated him, 49% to 44%.


title: Susan Collins

text: Collins was elected to the Senate in 1996. During the campaign she pledged that, if elected, she would serve only two terms.


title: Susan Collins

text: Collins was reelected in 2002 over State Senator Chellie Pingree, 58%–42%, in 2008 over Representative Tom Allen, 61.5%–38.5%, and in 2014 over Shenna Bellows, 68.5%–31.5%. In her first three reelection campaigns, she carried every county in Maine.


title: Susan Collins

text: In 2020, Collins was challenged by Democratic State House Speaker Sara Gideon. The hotly contested race became the most expensive in Maine history, with Collins spending $23 million and Gideon nearly $48 million. The race also had national implications, as defeating Collins was a key part of the Democrats' strategy to achieve a Senate majority. Despite trailing Gideon in every public poll of the race, Collins defeated Gideon by a decisive margin.


title: Susan Collins

text: In 2009, Collins was called one of "the last survivors of a once common species of moderate Northeastern Republican". She is considered a centrist Republican and an influential player in the Senate.


title: Susan Collins

text: Although she shared a centrist ideology with Maine's former senator, Olympia Snowe, Collins is considered a "half-turn more conservative" than Snowe. She was consistently endorsed by the Human Rights Campaign, a major LGBT rights organization, until 2020. She supported John McCain in the 2008 presidential election. She became the state's senior senator in 2013 when Snowe left the Senate and was replaced by independent Angus King, who defeated Collins in the 1994 governor election.


title: Susan Collins

text: In the 1990s, Collins played an important role during the Senate's impeachment trial of Bill Clinton when she and Snowe sponsored a motion that would have allowed the Senate to vote separately on the charges and the remedy. The motion failed, and Snowe and Collins voted to acquit, believing that while Clinton had committed perjury, that was not grounds for removal from office.