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John McCain
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John McCain
on December 23, 2017, the State Department announced that the United States would provide Ukraine with "enhanced defensive capabilities".
## Sixth and final Senate term.
McCain chaired the January 5, 2017, hearing of the Senate Armed Services Committee where Republican and Democratic senators and intelligence officers, including James R. Clapper Jr., the Director of National Intelligence, Michael S. Rogers, the head of the National Security Agency and United States Cyber Command presented a "united front" that "forcefully reaffirmed the conclusion that the Russian government used hacking and leaks to try to influence the presidential election."
In June 2017, McCain voted to support President
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Trump's controversial arms deal with Saudi Arabia.
Repeal and replacement of Obamacare (the Patient Protection and Affordable Care Act) was a centerpiece of McCain's 2016 re-election campaign, and in July 2017 he said, "Have no doubt: Congress must replace Obamacare, which has hit Arizonans with some of the highest premium increases in the nation and left 14 of Arizona's 15 counties with only one provider option on the exchanges this year." He added that he supports affordable and quality health care, but objected that the pending Senate bill did not do enough to shield the Medicaid system in Arizona.
In response to the death of Chinese Nobel Peace Prize laureate Liu Xiaobo, who died of organ
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failure while in government custody, McCain said that "this is only the latest example of Communist China's assault on human rights, democracy, and freedom."
In September 2017, as the Rohingya crisis in Myanmar became ethnic cleansing of the Rohingya Muslim minority, McCain announced moves to scrap planned future military cooperation with Myanmar.
In October 2017, McCain praised President Trump's decision to decertify Iran's compliance with the Iran nuclear deal (JCPOA) while not yet withdrawing the U.S. from the agreement, saying that the Obama-era policy failed "to meet the multifaceted threat Iran poses. The goals President Trump presented in his speech today are a welcomed long overdue
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change."
### Brain tumor diagnosis and surgery.
On July 14, 2017, McCain underwent a minimally invasive craniotomy at Mayo Clinic Hospital in Phoenix, Arizona, in order to remove a blood clot above his left eye. His absence prompted Senate Majority Leader Mitch McConnell to delay a vote on the Better Care Reconciliation Act. Five days later, Mayo Clinic doctors announced that the laboratory results from the surgery confirmed the presence of a glioblastoma, which is a very aggressive brain tumor. Standard treatment options for this tumor include chemotherapy and radiation, although even with treatment, average survival time is approximately 14 months. McCain was a survivor of previous cancers,
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including melanoma.
President Trump made a public statement wishing Senator McCain well, as did many others, including former President Obama. On July 19, McCain's senatorial office issued a statement that he "appreciates the outpouring of support he has received over the last few days. He is in good spirits as he continues to recover at home with his family in Arizona. He is grateful to the doctors and staff at Mayo Clinic for their outstanding care, and is confident that any future treatment will be effective." On July 24, McCain announced via Twitter that he would return to the United States Senate the following day.
### Return to the Senate.
McCain returned to the Senate on July 25, less
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than two weeks after brain surgery. He cast a deciding vote allowing the Senate to begin consideration of bills to replace the Affordable Care Act. Along with that vote, he delivered a speech criticizing the party-line voting process used by the Republicans, as well as by the Democrats in passing the Affordable Care Act to begin with, and McCain also urged a "return to regular order" utilizing the usual committee hearings and deliberations. On July 28, he cast the decisive vote against the Republicans' final proposal that month, the so-called "skinny repeal" option, which failed 49–51.
McCain did not vote in the Senate after December 2017, remaining instead in Arizona to undergo cancer treatment.
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On April 15, 2018, he underwent surgery for an infection relating to diverticulitis and the following day was reported to be in stable condition.
## Committee assignments.
- Committee on Armed Services (Chair)
- as chair of the full committee may serve as an ex-officio member of any subcommittee
- Committee on Homeland Security and Governmental Affairs
- Permanent Subcommittee on Investigations
- Subcommittee on Financial and Contracting Oversight
- Committee on Indian Affairs
- Committee on Intelligence (ex-officio)
## Caucus memberships.
- International Conservation Caucus
- Senate Diabetes Caucus
- Senate National Security Caucus (Co-Chair)
- Sportsmen's Caucus
- Senate Wilderness
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and Public Lands Caucus
- Senate Ukraine Caucus
- Republican Main Street Partnership.
# Death and funeral.
On August 24, 2018, McCain's family announced that he would no longer receive treatment for his cancer. He died the following day at 4:28 p.m. MST (11:28 p.m. UTC), with his wife and family beside him, at his home in Cornville, Arizona.
McCain lay in state in the Arizona State Capitol in Phoenix on August 29, which would have been his 82nd birthday. This was followed by a service at North Phoenix Baptist Church on August 30. His remains were then moved to Washington, D.C. to lie in state in the rotunda of the United States Capitol on August 31, which was followed by a service at the
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Washington National Cathedral on September 1. He was a "lifelong Episcopalian" who attended, but did not join, a Southern Baptist church for at least 17 years; memorial services were scheduled in both denominations. Prior to his death, McCain requested that former Presidents George W. Bush and Barack Obama deliver eulogies at his funeral, and asked that both President Donald Trump and former Alaska Governor and 2008 vice presidential nominee Sarah Palin not attend any of the services. McCain himself planned the funeral arrangements and selected his pallbearers for the service in Washington; the pallbearers included former Vice President Joe Biden, former Wisconsin Senator Russ Feingold, former
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Secretary of Defense William Cohen, actor Warren Beatty, and Russian dissident Vladimir Vladimirovich Kara-Murza.
Multiple foreign leaders attended McCain's service: Secretary General of NATO Jens Stoltenberg, President of Ukraine Petro Poroshenko, Speaker of Taiwan's Congress Su Jia-chyuan, National Defense Minister of Canada Harjit Sajjan, Defense Minister Jüri Luik and Foreign Minister Sven Mikser of Estonia, Foreign Minister of Latvia Edgars Rinkēvičs, Foreign Minister of Lithuania Linas Antanas Linkevičius, and Foreign Affairs Minister of Saudi Arabia Adel al-Jubeir.
Dignitaries who gave eulogies at the Memorial Service in Washington National Cathedral included Barack Obama, George W.
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Bush, Henry Kissinger, Joe Lieberman, and his daughter Meghan McCain. The New Yorker magazine hailed the service as the biggest resistance meeting of the Trump era.
Many American political figures paid tribute at the funeral. Those who attended included former United States Presidents Obama, Bush, Clinton, Carter; First Ladies Michelle, Laura, Hillary, Rosalyn; and former Vice Presidents Biden, Cheney, Gore, and Quayle. Former President George H.W. Bush (who died three months after McCain) was too ill to attend the service and President Trump was not invited. Many figures from political life both current and former from both political parties attended. Figures included, John F. Kelly, John
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Mattis, Bob Dole, Madeleine Albright, John Kerry, Mitch McConnell, Paul Ryan, Nancy Pelosi, Chuck Schumer, Mitt Romney, Lindsey Graham, Jeff Flake, Elizabeth Warren, Jon Huntsman. Ivanka Trump and Jared Kushner attended to the displeasure of Meghan McCain. Journalists Carl Bernstein, Tom Brokaw, and Charlie Rose, as well as actors Warren Beatty, Annette Benning, and comedians Jay Leno, and Joy Behar also attended the funeral.
On September 2, the funeral cortege traveled from Washington, D.C. through Annapolis, Maryland, where the streets were lined with crowds of onlookers, to the Naval Academy. A private service was held at the Naval Academy Chapel, attended by the brigade of midshipmen and
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McCain's classmates. After the chapel service, McCain was buried at the United States Naval Academy Cemetery, next to his Naval Academy classmate and lifelong friend Admiral Charles R. Larson.
Many celebrities paid tribute to the late Senator on Twitter. Those included, Tom Hanks who tweeted "Duty. Honor. Country. Our nation thanks you, John McCain. There has been no finer son of America". Whoopi Goldberg, Ellen DeGeneres, Reese Witherspoon, Jimmy Kimmel, and Khloe Kardashian also tweeted out remembrances of the late Senator.
Arizona Governor Doug Ducey was empowered to appoint McCain's interim replacement until a special election is held in 2020 to determine who is to serve out the remainder
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of McCain's term, which ends in January 2023 and thus appointed the then former Arizona U.S. Senator Jon Kyl to fill the vacancy. Under Arizona law, the appointed replacement must be of the same party as McCain, a Republican. Newspaper speculation about potential appointees has included McCain's widow Cindy, former Senator Jon Kyl, and former Representatives Matt Salmon and John Shadegg. Ducey said that he would not make a formal appointment until after McCain's final funeral and burial; on September 4, two days after McCain was buried, Ducey appointed Kyl to fill McCain's seat.
## Tributes.
McCain received many tributes and condolences, including from Congressional colleagues, all living
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former Presidents – Jimmy Carter, George H. W. Bush, Bill Clinton, George W. Bush, Barack Obama – and former Vice President Joe Biden, as well as Vice President Mike Pence and President Richard Nixon's daughters Tricia Nixon Cox and Julie Nixon Eisenhower. French President Emmanuel Macron, Taiwanese President Tsai Ing-wen, Ukrainian President Petro Poroshenko and Prime Minister Volodymyr Groysman, Australian Prime Minister Scott Morrison, who had just taken office the previous day, and former Prime Minister Malcolm Turnbull, British Prime Minister Theresa May and former Prime Minister David Cameron, Canadian Prime Minister Justin Trudeau and former Prime Minister Stephen Harper, German Chancellor
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Angela Merkel and foreign minister Heiko Maas, Israeli Prime Minister Benjamin Netanyahu, Indian Prime Minister Narendra Modi, Afghanistan chief executive Abdullah Abdullah, Pakistani foreign minister Shah Mehmood Qureshi, the 14th Dalai Lama, and former Vietnamese ambassador to Washington Nguyễn Quốc Cường also sent condolences.
Colonel Trần Trọng Duyệt, who ran the Hỏa Lò Prison when McCain was held there, remarked, "At that time I liked him personally for his toughness and strong stance. Later on, when he became a US Senator, he and Senator John Kerry greatly contributed to promote Vietnam-US relations so I was very fond of him. When I learnt about his death early this morning, I feel very
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sad. I would like to send condolences to his family." In a TV interview, Senator Lindsey Graham said McCain's last words to him were "I love you, I have not been cheated." His daughter, Meghan McCain, shared her grief, stating that she was present at the moment he died.
At the 70th Primetime Emmy Awards, McCain was recognized in the "In Memoriam" segment, right before Aretha Franklin. Many fans questioned the inclusion of McCain in the segment because he wasn't known for television. He had, however, appeared in various television projects, including hosting and several cameo appearances on "Saturday Night Live". He also made appearances on "Parks and Recreation" and "24". McCain was also a
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good friend of "SNL" creator Lorne Michaels, who produced that year's Emmy Awards ceremony.
Senate Minority Leader Chuck Schumer (D-NY) announced that he would introduce a resolution to rename the Russell Senate Office Building after McCain. A quarter peal of Grandsire Caters in memory of McCain was rung by the bellringers of Washington National Cathedral the day following his death. Another memorial quarter peal was rung on September 6 on the Bells of Congress at the Old Post Office in Washington.
## Negative reaction from the White House.
Trump reportedly rejected the White House's plans to release a statement praising McCain's life, and he initially said nothing about McCain himself in
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a tweet that extended condolences to McCain's family. In addition, the flag at the White House, which had been lowered to half-staff the day of McCain's death (August 25), was raised back to full-staff at 12:01 a.m. on August 27. Trump reportedly felt that media coverage of McCain's death was excessive given that McCain was never president. In contrast with the White House's initial decision, many governors, both Democratic and Republican, had ordered flags in their states to fly at half-staff until McCain's interment, and Senate leaders Mitch McConnell and Chuck Schumer requested support from the Defense Department so that flags would be flown at half-staff on all government buildings. Following
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public backlash from the American Legion and AMVETS, Trump relented and ordered the White House flag back to half-staff later in the day on August 27. Trump belatedly issued a statement praising McCain's service to the country, and he signed a proclamation ordering flags to be flown at half-staff until McCain's interment at the Naval Academy Cemetery.
In March 2019—seven months after McCain's death—Trump issued a series of public statements that criticized McCain at least four times in five days. Trump also claimed that he approved McCain's funeral but was not thanked for it. However, the Washington National Cathedral responded that no governmental or presidential approval was needed for McCain's
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funeral because he was not a former president. McCain's lying in state was approved by the Senate, while Trump did approve the transport for McCain's body. Trump also described himself as having "got the job done" on the Veterans Choice Act while claiming McCain failed on the same issue. However, McCain was actually one of the two main authors of the bill, which President Barack Obama signed into law in 2014. Trump simply signed the VA MISSION Act of 2018 (), an expansion of that law, which was worked on by McCain and others, and includes McCain's name in its full title. Trump also falsely claimed that McCain graduated "last in his class" when McCain was actually fifth from last.
# Political
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positions.
Various advocacy groups have given McCain scores or grades as to how well his votes align with the positions of each group. The American Conservative Union awarded McCain a lifetime rating of 82 percent through 2015, while McCain registered an average lifetime 12 percent "Liberal Quotient" from Americans for Democratic Action through 2015. CrowdPac, which rates politicians based on donations made and received, gave Senator McCain a score of 4.3C with 10C being the most conservative and 10L being the most liberal.
The non-partisan "National Journal" rates a Senator's votes by what percentage of the Senate voted more liberally than he or she, and what percentage more conservatively,
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in three policy areas: economic, social, and foreign. For 2005–2006 (as reported in the 2008 "Almanac of American Politics"), McCain's average ratings were as follows: economic policy: 59 percent conservative and 41 percent liberal; social policy: 54 percent conservative and 38 percent liberal; and foreign policy: 56 percent conservative and 43 percent liberal. In 2012, the "National Journal" gave McCain a composite score of 73 percent conservative and 27 percent liberal, while in 2013 he received a composite score of 60 percent conservative and 40 percent liberal.
Columnists such as Robert Robb and Matthew Continetti used a formulation devised by William F. Buckley Jr. to describe McCain as
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"conservative" but not "a conservative", meaning that while McCain usually tended towards conservative positions, he was not "anchored by the philosophical tenets of modern American conservatism". Following his 2008 presidential election loss, McCain began adopting more orthodox conservative views; the magazine "National Journal" rated McCain along with seven of his colleagues as the "most conservative" Senators for 2010 and he achieved his first 100 percent rating from the American Conservative Union for that year. During Barack Obama's presidency, McCain was one of the top five Republicans most likely to vote with Obama's position on significant votes; McCain voted with Obama's position on
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such votes more than half the time in 2013 and was "censured by the Arizona Republican party for a so-called 'liberal' voting record".
From the late 1990s until 2008, McCain was a board member of Project Vote Smart which was set up by Richard Kimball, his 1986 Senate opponent. The project provides non-partisan information about the political positions of McCain and other candidates for political office. Additionally, McCain used his Senate website to describe his political positions.
# Cultural and political image.
McCain's personal character was a dominant feature of his public image. This image includes the military service of both himself and his family, the circumstances and tensions
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surrounding the end of his first marriage and beginning of second, his maverick political persona, his temper, his admitted problem of occasional ill-considered remarks, and his close ties to his children from both his marriages.
McCain's political appeal was more nonpartisan and less ideological compared to many other national politicians. His stature and reputation stemmed partly from his service in the Vietnam War. He also carried physical vestiges of his war wounds, as well as his melanoma surgery. When campaigning, he quipped: "I am older than dirt and have more scars than Frankenstein."
Writers often extolled McCain for his courage not just in war but in politics, and wrote sympathetically
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about him. McCain's shift of political stances and attitudes during and especially after the 2008 presidential campaign, including his self-repudiation of the maverick label, left many writers expressing sadness and wondering what had happened to the McCain they thought they had known. By 2013, some aspects of the older McCain had returned, and his image became that of a kaleidoscope of contradictory tendencies, including, as one writer listed, "the maverick, the former maverick, the curmudgeon, the bridge builder, the war hero bent on transcending the call of self-interest to serve a cause greater than himself, the sore loser, old bull, last lion, loose cannon, happy warrior, elder statesman,
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lion in winter."
In his own estimation, McCain was straightforward and direct, but impatient. His other traits included a penchant for lucky charms, a fondness for hiking, and a sense of humor that sometimes backfired spectacularly, as when he made a joke in 1998 about the Clintons that was widely deemed not fit to print in newspapers: "Do you know why Chelsea Clinton is so ugly? – Because Janet Reno is her father." McCain subsequently apologized profusely, and the Clinton White House accepted his apology. McCain did not shy away from addressing his shortcomings, and he apologized for them. He was known for sometimes being prickly and hot-tempered with Senate colleagues, but his relations with
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his own Senate staff were more cordial, and inspired loyalty towards him. He formed a strong bond with two senators, Joe Lieberman and Lindsey Graham, over hawkish foreign policy and overseas travel, and they became dubbed the "Three Amigos".
McCain acknowledged having said intemperate things in years past, though he also said that many stories have been exaggerated. One psychoanalytic comparison suggested that McCain was not the first presidential candidate to have a temper, and cultural critic Julia Keller argued that voters want leaders who are passionate, engaged, fiery, and feisty. McCain employed both profanity and shouting on occasion, although such incidents became less frequent over
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the years. Lieberman made this observation: "It is not the kind of anger that is a loss of control. He is a very controlled person." Senator Thad Cochran, who knew McCain for decades and had battled him over earmarks, expressed concern about a McCain presidency: "He is erratic. He is hotheaded. He loses his temper and he worries me." Yet Cochran supported McCain for president when it was clear he would win the nomination. The "Chicago Tribune" editorial board called McCain a patriot, who although sometimes wrong was fearless, and that he deserves to be thought of among the few US senators in history, whose names are more recognizable than some presidents.
All of McCain's family members were
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on good terms with him, and he defended them against some of the negative consequences of his high-profile political lifestyle. His family's military tradition extends to the latest generation: son John Sidney IV ("Jack") graduated from the U.S. Naval Academy in 2009, becoming the fourth generation John S. McCain to do so, and is a helicopter pilot; son James served two tours with the Marines in the Iraq War; and son Doug flew jets in the navy. His daughter Meghan became a blogging and Twittering presence in the debate about the future of the Republican Party following the 2008 elections, and showed some of his maverick tendencies. In 2017 Meghan joined the cast of the popular ABC talk show
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"The View" as a co-host. Senator McCain himself also appeared as a guest on the program.
McCain appeared in several television shows and films while he was a sitting senator. He made uncredited cameo appearances in "Wedding Crashers" and "24" and had two uncredited cameos in "Parks and Recreation". McCain also hosted "Saturday Night Live" in 2002 and appeared in two episodes in 2008.
# Awards and honors.
In addition to his military honors and decorations, McCain was granted a number of civilian awards and honors.
In 1997, "Time" magazine named McCain as one of the "25 Most Influential People in America".
In 1999, McCain shared the Profile in Courage Award with Senator Russ Feingold for
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their work towards campaign finance reform. The following year, the same pair shared the Paul H. Douglas Award for Ethics in Government. In 2005, The Eisenhower Institute awarded McCain the Eisenhower Leadership Prize. The prize recognizes individuals whose lifetime accomplishments reflect Dwight D. Eisenhower's legacy of integrity and leadership. In 2006, the Bruce F. Vento Public Service Award was bestowed upon McCain by the National Park Trust. The same year, McCain was awarded the Henry M. Jackson Distinguished Service Award by the Jewish Institute for National Security Affairs, in honor of Senator Henry M. "Scoop" Jackson. In 2007, the World Leadership Forum presented McCain with the Policymaker
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of the Year Award; it is given internationally to someone who has "created, inspired or strongly influenced important policy or legislation". In 2010, President Mikheil Saakashvili of Georgia awarded McCain the Order of National Hero, an award never previously given to a non-Georgian. In 2015, the Kiev Patriarchate awarded McCain its own version of the Order of St. Vladimir. In 2016, Allegheny College awarded McCain, along with Vice President Joe Biden, its Prize for Civility in Public Life. In August 2016, Petro Poroshenko, the President of Ukraine, awarded McCain with the highest award for foreigners, the Order of Liberty. In 2017, Hashim Thaçi, the President of Kosovo, awarded McCain the
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"Urdhër i Lirisë" (Order of Freedom) medal for his contribution to the freedom and independence of Kosovo, and its partnership with the U.S. McCain also received the Liberty Medal from the National Constitution Center in 2017. In the spring of 2018 McCain was decorated with the Grand Cordon of the Order of the Rising Sun from the Japanese Emperor for 'strengthening bilateral relations and promoting friendship between Japan and the United States'.
McCain received several honorary degrees from colleges and universities in the United States and internationally. These include ones from Colgate University (LL.D 2000), The Citadel (DPA 2002), Wake Forest University (LL.D May 20, 2002), the University
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of Southern California (DHL May 2004), Northwestern University (LL.D June 17, 2005), Liberty University (2006), The New School (2006), and the Royal Military College of Canada (D.MSc June 27, 2013). He was also made an Honorary Patron of the University Philosophical Society at Trinity College Dublin in 2005.
On 11 July 2018, , originally named in honor of the Senator's father and grandfather, was rededicated in the Senator's name also.
# Bibliography.
## Books.
- "Faith of My Fathers" by John McCain, Mark Salter (Random House, August 1999) (later made into the 2005 television film "Faith of My Fathers")
- "Worth the Fighting For" by John McCain, Mark Salter (Random House, September 2002)
-
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"Why Courage Matters: The Way to a Braver Life" by John McCain, Mark Salter (Random House, April 2004)
- "Character Is Destiny: Inspiring Stories Every Young Person Should Know and Every Adult Should Remember" by John McCain, Mark Salter (Random House, October 2005)
- "Hard Call: Great Decisions and the Extraordinary People Who Made Them" by John McCain, Mark Salter (Hachette, August 2007)
- "Thirteen Soldiers: A Personal History of Americans at War" by John McCain, Mark Salter (Simon & Schuster, November 2014)
- "The Restless Wave: Good Times, Just Causes, Great Fights, and Other Appreciations" by John McCain, Mark Salter (Simon & Schuster, May 2018)
## Articles and forewords.
- "How
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the POW's Fought Back", by John S. McCain III, Lieut. Commander, U.S. Navy, "U.S. News & World Report", May 14, 1973 (reprinted for web under different title in 2008). Reprinted in "Reporting Vietnam, Part Two: American Journalism 1969–1975" (The Library of America, 1998)
- "The Code of Conduct and the Vietnam Prisoners of War", by John S. McCain, Commander USN, National War College, April 8, 1974 (actual paper)
- Foreword by John McCain to "A Code to Keep: The True Story of America's Longest-Held Civilian POW in Vietnam" by Ernest C. Brace (St. Martin's Press, 1988)
- Speeches of John McCain, 1988–2000
- Foreword by John McCain to "Glory Denied: The Saga of Jim Thompson, America's Longest-held
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Prisoner" by Tom Philpott (W. W. Norton, 2001)
- Foreword by John McCain to "The Best and the Brightest" by David Halberstam (Random House, 2001 edition)
- Foreword by John S. McCain to "Unfinished Business: Afghanistan, the Middle East and Beyond – Defusing the Dangers That Threaten America's Security" by Harlan Ullman (Citadel Press, June 2002)
- Foreword by John McCain and Max Cleland to "Odysseus in America: Combat Trauma and the Trials of Homecoming" by Jonathan Shay (Scribner, November 2002)
- Foreword by John McCain to "Debunking 9/11 Myths: Why Conspiracy Theories Can't Stand Up to the Facts" by the Editors of "Popular Mechanics" (Hearst, August 2006)
- Introduction by John McCain
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to "Pearl Harbor, the Day of Infamy, an Illustrated History" by Dan van der Vat (Black Walnut Books, 2007)
- "An Enduring Peace Built on Freedom: Securing America's Future" by John McCain "Foreign Affairs", November/December 2007
# See also.
- Electoral history of John McCain
- List of United States Senators born outside the United States
# Bibliography.
- Alexander, Paul. "Man of the People: The Life of John McCain" (John Wiley & Sons, Hoboken, New Jersey 2002). .
- Brock, David and Waldman, Paul. "Free Ride: John McCain and the Media" (Anchor Books, New York 2008). .
- Drew, Elizabeth. "Citizen McCain" (Simon & Schuster, New York 2002). .
- Feinberg, Barbara Silberdick. "John McCain:
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Serving His Country" (Millbrook Press, Brookfield, Connecticut 2000). .
- Hubbell, John G. "P.O.W.: A Definitive History of the American Prisoner-Of-War Experience in Vietnam, 1964–1973" (Reader's Digest Press, New York 1976). .
- Karaagac, John. "John McCain: An Essay in Military and Political History" (Lexington Books, Lanham, Maryland 2000). .
- McCain, John and Salter, Mark, "Faith of My Fathers" (Random House, New York 1999). .
- McCain, John and Salter, Mark. "Worth the Fighting For" (Random House, New York 2002). .
- Rochester, Stuart I. and Kiley, Frederick. "Honor Bound: American Prisoners of War in Southeast Asia, 1961–1973" (Naval Institute Press, Annapolis, Maryland 1999). .
-
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Schecter, Cliff. "The Real McCain: Why Conservatives Don't Trust Him and Why Independents Shouldn't" (PoliPoint Press, Sausalito, California 2008). .
- Timberg, Robert. "John McCain: An American Odyssey" (Touchstone Books, New York 1999). . Chapter 1 available online.
- Timberg, Robert. "The Nightingale's Song" (Simon & Schuster, New York 1996). . Chapter 1 available online.
- Welch, Matt. "McCain: The Myth of a Maverick" (Palgrave Macmillan, New York 2007). .
# External links.
- Senator John McCain official U.S. Senate site
- John McCain for Senate
- Sean Wilentz: "John McCain." In: "Encyclopædia Britannica", February 15, 2018
- Gates, H.L. John McCain's Interactive Family Tree. PBS.
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tives Don't Trust Him and Why Independents Shouldn't" (PoliPoint Press, Sausalito, California 2008). .
- Timberg, Robert. "John McCain: An American Odyssey" (Touchstone Books, New York 1999). . Chapter 1 available online.
- Timberg, Robert. "The Nightingale's Song" (Simon & Schuster, New York 1996). . Chapter 1 available online.
- Welch, Matt. "McCain: The Myth of a Maverick" (Palgrave Macmillan, New York 2007). .
# External links.
- Senator John McCain official U.S. Senate site
- John McCain for Senate
- Sean Wilentz: "John McCain." In: "Encyclopædia Britannica", February 15, 2018
- Gates, H.L. John McCain's Interactive Family Tree. PBS. February 11, 2016. Accessed February 17, 2017
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SOB
Sob is a verb meaning cry.
Sob and SoB may refer to:
- Souls on Board (sometimes POB for People on Board), used in Aviation communication
- Seventeen or Bust, a distributed computing project
- Special Operations Battalion (SOB), an elite unit of the Croatian army
- Society of Old Brooklynites
- Son of a bitch, an insult or curse phrase
- Sons of Ben (MLS supporters association)
- Suggested Opening Bid, a term in auctioneering
- Super Oralloy Bomb, a nuclear bomb
- Senate Office Building
- Dirksen Senate Office Building
- Russell Senate Office Building
- Hart Senate Office Building
- S.O.B. Hill, a mountain in Utah, United States
- Start of business, the start of the business
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day (as opposed to EOB)
# Biomedicine.
- Shortness of breath or Dyspnea, a medical symptom relating to breathing difficulties
- "Sobralia", a genus of orchids
- Super Optimal Broth, a bacterial growth media
# Music and media.
- "S.O.B." (film), a 1981 film by Blake Edwards
- "The Sob", a 1953 Turkish film
- S.O.B. (band), a Japanese grindcore band
- "S.O.B." (song), by Nathaniel Rateliff and the Night Sweats, 2015
- Scars on Broadway, an American rock band
- Sons of Batman, a group of characters in a "Batman" comics miniseries
- Sons of Butcher (band), a Canadian rock band
- "Sons of Butcher" (TV show), a cartoon inspired by the band
- Sounds of Blackness, an American vocal and
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an American vocal and instrumental ensemble
- SOB's, Sounds of Brazil, a live music venue in New York City
- Souls on Board, an Australian indie rock band
- Styles of Beyond, an underground rap group
- "Switched-On Bach", an album by Walter Carlos
- "S.O.B.s", an episode of the television series "Arrested Development"
# Transport.
- Seaford-Oyster Bay Expressway
- Balaton Airport, SOB in IATA code
- Southbourne railway station, SOB in Network Rail code
- Stadt-Omnibus Bern, a former public transport operator in Bern, Switzerland
- Südostbahn, a railway company in Switzerland
- SüdostBayernBahn, a railway company in southern Germany
- Spirit of Britain, a Dover - Calais P&O Ferry
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Galerius
Galerius (; ; "circa" 250 – April or May 311) was Roman emperor from 305 to 311. During his reign, he campaigned, aided by Diocletian, against the Sassanid Empire, sacking their capital Ctesiphon in 299. He also campaigned across the Danube against the Carpi, defeating them in 297 and 300. Although he was a staunch opponent of Christianity, Galerius ended the Diocletianic Persecution when he issued an Edict of Toleration in Serdica in 311.
# Early life.
Galerius was born near Serdica, in Dacia Ripensis, later named Dacia Mediterranea, though some modern scholars consider the strategic site where he later built his palace named after his mother – Felix Romuliana (Gamzigrad) – his birth
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and funeral place. His father was a Thracian and his mother Romula was a Dacian woman, who left Dacia because of the Carpians' attacks. He originally followed his father's occupation, that of a herdsman, where he got his surname of Armentarius (Latin: "armentum", herd).
He served with distinction as a soldier under Emperors Aurelian and Probus, and in 293 at the establishment of the Tetrarchy, was designated "Caesar" along with Constantius Chlorus, receiving in marriage Diocletian's daughter Valeria (later known as Galeria Valeria), and at the same time being entrusted with the care of the Illyrian provinces. After a few years campaigning against Sarmatians and Goths on the Danube, he received
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command of the legions on the eastern imperial limits. Soon after his appointment, Galerius was dispatched to Egypt to fight the rebellious cities Busiris and Coptos.
# War with Persia.
## Invasion, counterinvasion.
In 294, Narseh, a son of Shapur I, who had been passed over for the Sassanid succession, came into power in Persia. Narseh probably moved to eliminate Bahram III, a young man installed by a noble named Vahunam in the wake of Bahram II's death in 293. In early 294, Narseh sent Diocletian the customary package of gifts, but within Persia, he was destroying every trace of his immediate predecessors, erasing their names from public monuments. He sought to identify himself with the
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warlike reigns of Ardashir (r. 226–41) and Shapur (r. 241–72), who had sacked Roman Antioch and captured Emperor Valerian.
In 295 or 296, Narseh declared war on Rome. He appears to have first invaded western Armenia, retaking the lands delivered to Tiridates in the peace of 287. He occupied the lands there until the following year. The historian Ammianus Marcellinus, circa 320-395, is the only source detailing the initial invasion of Armenia. Southern (1999, 149) dates the invasion to 295; Barnes (1982, 17, 293) mentions an earlier, unsuccessful invasion by Narseh based on the fact that the title "Persici Maximi" was given to all four emperors; Odahl (2004, 59) concurs with Barnes and suggests
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that Saracen princes in the Syrian desert collaborated with Narseh's invasion. Narseh then moved south into Roman Mesopotamia, where he inflicted a severe defeat on Galerius, then commander of the eastern forces, in the region between Carrhae (Harran, Turkey) and Callinicum (Raqqa, Syria). Diocletian may or may not have been present at the battle, but presented himself soon afterwards at Antioch, where the official version of events was made clear: Galerius was to take all the blame for the affair. In Antioch, Diocletian forced Galerius to walk a mile in advance of his imperial cart while still clad in the purple robes of an emperor. The message conveyed was clear; the loss at Carrhae was not
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due to the failings of the empire's soldiers, but due to the failings of their commander, and Galerius' failures would not be accepted. Galerius' position at the head of the caravan possibly was merely the conventional organization of an imperial progression, designed to show a Caesar's deference to his Augustus.
Galerius's army was reinforced probably in the spring of 298 by new contingents collected from the empire's Danubian holdings. Narseh did not advance from Armenia and Mesopotamia leaving Galerius to lead the offensive in 298 with an attack on northern Mesopotamia via Armenia. Diocletian may or may not have been present to assist the campaign. Narseh retreated to Armenia to fight Galerius'
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force, to Narseh's disadvantage; the rugged Armenian terrain was favorable to Roman infantry, but not to Sassanid cavalry. Local aid gave Galerius the advantage of surprise over the Persian forces, and, in two successive battles, Galerius secured victories over Narseh.
During the second encounter, the Battle of Satala in 298, Roman forces seized Narseh's camp, his treasury, his harem, and his wife. Narseh's wife would live out the remainder of the war in Daphne, a suburb of Antioch, serving as a constant reminder to the Persians of the Roman victory. Galerius advanced into Media and Adiabene, winning continuous victories, most prominently near Theodosiopolis (Erzurum), and securing Nisibis
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(Nusaybin) before 1 October 298. He moved down the Tigris, taking Ctesiphon, and gazing onwards to the ruins of Babylon before returning to Roman territory via the Euphrates. No source ever specifically claims that Ctesiphon was sacked, but it is assumed to have been, primarily due to the seizure of Narseh's wife and harem.
## Peace negotiations.
Narseh had previously sent an ambassador to Galerius to plead for the return of his wife and children, but Galerius had dismissed this ambassador, reminding him of how Shapur had treated Valerian. The Romans, in any case, treated Narseh's captured family well perhaps seeking to evoke comparisons to Alexander and his beneficent conduct towards the
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family of Darius III. Peace negotiations began in the spring of 299, with both Diocletian and Galerius presiding. Their "magister memoriae" (secretary) Sicorius Probus was sent to Narseh to present terms.
The conditions of the Peace of Nisibis were heavy: Persia would give up territory to Rome, making the Tigris the boundary between the two empires. Further terms specified that Armenia was returned to Roman domination with the fort of Ziatha as its border; Caucasian Iberia would pay allegiance to Rome under a Roman appointee; Nisibis, now under Roman rule, would become the sole conduit for trade between Persia and Rome; and Rome would exercise control over the five satrapies between the Tigris
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and Armenia: Ingilene, Sophanene (Sophene), Arzanene (Aghdznik), Corduene, and Zabdicene (near modern Hakkâri, Turkey). These regions included the passage of the Tigris through the Anti-Taurus range; the Bitlis pass, the quickest southerly route into Persian Armenia; and access to the Tur Abdin plateau. With these territories, Rome would have an advance station north of Ctesiphon, and would be able to slow any future advance of Persian forces through the region. Under the terms of the peace, Tiridates would regain both his throne and the entirety of his ancestral claim, and Rome would secure a wide zone of cultural influence in the region. Because the empire was able to sustain such constant
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warfare on so many fronts, it has been taken as a sign of the essential efficacy of the Diocletianic system and the goodwill of the army towards the tetrarchic enterprise.
# Rule as Augustus.
After the elevation of Constantius I and Galerius to the rank of "Augustus", two new Caesars were required to take their place. The two persons whom Galerius promoted to the rank of Caesar were very much his creatures, and he hoped to enhance his authority throughout the empire with their elevation.
First was Maximinus Daia, whose mother was Galerius' sister. An inexperienced youth with little formal education, he was invested with the purple, exalted to the dignity of Caesar, and assigned the command
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of Egypt and Syria. Second was Severus, Galerius' comrade in arms; he was sent to Milan to receive the possession of Italy and Africa. According to the forms of the constitution, Severus acknowledged the supremacy of the western emperor, but he was absolutely devoted to the commands of his benefactor Galerius, who, reserving to himself the intermediate countries from the confines of Italy to those of Syria, firmly established his power over three-quarters of the empire.
His hopes were dashed when his colleague Constantius died at York in 306 and the legions elevated his son Constantine to the position of Augustus. Galerius only discovered this when he received a letter from Constantine, who
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informed him of his father's death, modestly asserted his natural claim to the succession, and respectfully lamented that the enthusiastic violence of his troops had not allowed him to obtain the imperial purple in the regular and constitutional manner. The first emotions of Galerius were those of surprise, disappointment, and rage, and as he could seldom restrain his passions, he threatened to burn both the letter and the messenger.
Later, however, when he had time to reconsider his position, he inevitably saw that his chances of winning a war against Constantine were doubtful at best. Galerius was well aware of Constantine’s strengths – Constantine had been his guest for some time at Nicomedia
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– and knew that Constantius' legions were wildly devoted to his son. Therefore, without either condemning or ratifying the choice of the Roman army, Galerius accepted the son of his deceased colleague as the ruler of the provinces beyond the Alps; but he gave him only the title of Caesar, and the fourth rank among the Roman princes, whilst he conferred the vacant place of Augustus on his favourite Severus.
The ambitious spirit of Galerius was only just over this disappointment when he beheld the unexpected loss of Italy to Maxentius, who was married to his daughter Valeria Maximilla. Galerius’ need for additional revenue had persuaded him to make a very strict and rigorous examination of the
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property of his subjects for the purpose of a general taxation. A very minute survey was taken of their real estates and, wherever there was the slightest suspicion of concealment, torture was used to obtain a sincere declaration of their personal wealth. Italy had traditionally been exempt from any form of taxation, but Galerius ignored this precedent, and the officers of the revenue already began to number the Roman people, and to settle the proportion of the new taxes. Italy began to murmur against this indignity and Maxentius used this sentiment to declare himself emperor in Italy, to the fury of Galerius. Therefore, Galerius ordered his colleague Severus to immediately march to Rome, in
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the full confidence that, by his unexpected arrival, he would easily suppress the rebellion. Severus was captured after his troops deserted to their old commander Maximian, who had once again been elevated to the rank of co-emperor, this time by his son Maxentius. Severus was later executed.
The importance of the occasion needed the presence and abilities of Galerius. At the head of a powerful army collected from Illyricum and the East, he entered Italy, determined to avenge Severus and to punish the rebellious Romans. But due to the skill of Maximian, Galerius found every place hostile, fortified, and inaccessible; and though he forced his way as far as Narni, within sixty miles of Rome, his
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control in Italy was confined to the narrow limits of his camp.
Seeing that he was facing ever-greater difficulties, Galerius made the first advances towards reconciliation, and dispatched two officers to tempt the Romans by the offer of a conference, and the declaration of his paternal regard for Maxentius, reminding them that they would obtain much more from his willing generosity than anything that might have been obtained through a military campaign. The offers of Galerius were rejected with firmness, his friendship refused, and it was not long before he discovered that, unless he retreated, he would share Severus' fate. It was not a moment too soon; large monetary gifts from Maxentius
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to his soldiers had corrupted the fidelity of the Illyrian legions. When Galerius finally began his withdrawal from Italy, it was only with great difficulty that he managed to stop his veterans deserting him.
In frustration, Galerius allowed his legions to ravage the countryside as they passed northwards. Maxentius declined to make a general engagement.
With so many emperors now in existence, in 308 Galerius, together with the retired emperor Diocletian and the now active Maximian, called an imperial 'conference' at Carnuntum on the River Danube to rectify the situation and bring some order back into the imperial government. Here it was agreed that Galerius’ long-time friend and military companion
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Licinius, who had been entrusted by Galerius with the defense of the Danube while Galerius was in Italy, would become Augustus in the West, with Constantine as his Caesar. In the East, Galerius remained Augustus and Maximinus remained his Caesar. Maximian was to retire, and Maxentius was declared a usurper.
Galerius’ plan soon failed. The news of Licinius’ promotion was no sooner carried into the East, than Maximinus, who governed the provinces of Egypt and Syria, rejected his position as Caesar, and, notwithstanding the prayers as well as arguments of Galerius, exacted the equal title of Augustus. For the first, and indeed for the last time, six emperors administered the Roman world. Though
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the opposition of interest, and the memory of a recent war, divided the empire into two great hostile powers, their mutual fears and the fading authority of Galerius produced an apparent tranquility in the imperial government.
The last years of Galerius saw him relinquishing his aspirations towards being the supreme ruler of the empire, though he managed to retain the position of first among equals. He spent the remainder of his years enjoying himself and ordering some important public works, such as discharging into the Danube the superfluous waters of Lake Pelso, and cutting down the immense forests that encompassed it.
# Persecution of Christians.
Christians had lived in peace during most
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of the rule of Diocletian. The persecutions that began with an edict of February 24, 303, were credited by Christians to Galerius' work, as he was a fierce advocate of the old ways and old gods. Christian houses of assembly were destroyed, for fear of sedition in secret gatherings.
Diocletian was not anti-Christian during the first part of his reign, and historians have claimed that Galerius decided to prod him into persecuting them by secretly burning the Imperial Palace and blaming it on Christian saboteurs. Regardless of who was at fault for the fire, Diocletian's rage was aroused and he began one of the last and greatest Christian persecutions in the history of the Roman Empire.
It was
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at the insistence of Galerius that the last edicts of persecution against the Christians were published, beginning on February 24, 303, and this policy of repression was maintained by him until the appearance of the general edict of toleration, issued from Nicomedia in April 311, apparently during his last bout of illness (see "Edict of Toleration by Galerius"). Galerius's last request, that Christians should pray for him as he suffered with a painful and fatal illness, was in vain for he died six days later.
Initially one of the leading figures in the persecutions, Galerius later admitted that the policy of trying to eradicate Christianity had failed, saying: "wherefore, for this our indulgence,
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they ought to pray to their God for our safety, for that of the republic, and for their own, that the republic may continue uninjured on every side, and that they may be able to live securely in their homes." Lactantius gives the text of the edict in his moralized chronicle of the bad ends to which all the persecutors came, "De Mortibus Persecutorum". This marked the end of official persecution of Christians. Christianity was officially legalized in the Roman Empire two years later in 313 by Constantine and Licinius in the Edict of Milan.
# Death.
Galerius died in late April or early May 311 from a horribly gruesome disease described by Eusebius and Lactantius, possibly some form of bowel
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cancer, gangrene or Fournier gangrene.
Galerius was buried in his mausoleum at Gamzigrad-Romuliana, which was part of the palace he built at his birthplace, today's Zaječar in Serbia. Several lumps composed of corroded iron ring mail (lorica hamata) have been found at the site. This mail armour may have been worn by the wax figure of the emperor that was burned during the imperial funeral and apotheosis ceremony. The entire site has been inscribed into the World Heritage List in June 2007.
# Anti-Roman accusations.
According to Lactantius, Galerius affirmed his Dacian identity and avowed himself the enemy of the Roman name once made emperor, even proposing that the empire should be called,
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not the Roman, but the Dacian Empire, much to the horror of the patricians and senators. He exhibited anti-Roman attitude as soon as he had attained the highest power, treating the Roman citizens with ruthless cruelty, like the conquerors treated the conquered, all in the name of the same treatment that the victorious Trajan had applied to the conquered Dacians, forefathers of Galerius, two centuries before.
# Honours.
Galerius Peak in Antarctica is named after Emperor Galerius.
# See also.
- Arch and Tomb of Galerius
- Civil Wars of the Tetrarchy (306–324)
# References.
## Ancient sources.
- "Codex Theodosianus".
- "Epitome de Caesaribus".
- Eusebius of Caesarea.
- Festus. "Breviarium".
-
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Lactantius. "De Mortibus Persecutorum" ("On the Deaths of the Persecutors").
- "XII Panegyrici Latini" ("Twelve Latin Panegyrics").
- Zosimus. "Historia Nova" ("New History").
## Modern sources.
- Banchich, Thomas M. "Iulianus (c. 286–293 AD)." "De Imperatoribus Romanis" (1997). Accessed March 8, 2008.
- Barnes, Timothy D. "Lactantius and Constantine." "The Journal of Roman Studies" 63 (1973): 29–46.
- Barnes, Timothy D. "Constantine and Eusebius". Cambridge, MA: Harvard University Press, 1981.
- Barnes, Timothy D. "The New Empire of Diocletian and Constantine". Cambridge, MA: Harvard University Press, 1982.
- Bleckmann, Bruno. "Diocletianus." In "Brill's New Pauly, Volume 4", edited
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by Hubert Cancik and Helmut Schneider, 429–38. Leiden: Brill, 2002.
- Bowman, Alan K., Peter Garnsey, and Averil Cameron. "The Cambridge Ancient History, Volume XII: The Crisis of Empire". Cambridge University Press, 2005.
- Brown, Peter. "The Rise of Western Christendom". Oxford: Blackwell Publishing, 2003.
- Burgess, R.W. "The Date of the Persecution of Christians in the Army". "Journal of Theological Studies" 47:1 (1996): 157–158.
- Corcoran, Simon. "The Empire of the Tetrarchs: Imperial Pronouncements and Government, AD 284–324". Oxford: Clarendon Press, 1996.
- Corcoran, "Before Constantine", Simon. "Before Constantine." In "The Cambridge Companion to the Age of Constantine", edited
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by Noel Lenski, 35–58. New York: Cambridge University Press, 2006. Hardcover Paperback
- DiMaio, Jr., Michael. "Constantius I Chlorus (305–306 AD)." "De Imperatoribus Romanis" (1996a). Accessed March 8, 2008.
- DiMaio, Jr., Michael. "Galerius (305–311 AD)." "De Imperatoribus Romanis" (1996b). Accessed March 8, 2008.
- DiMaio, Jr., Michael. "Maximianus Herculius (286–305 AD)." "De Imperatoribus Romanis" (1997). Accessed March 8, 2008.
- Elliott, T. G. "The Christianity of Constantine the Great". Scranton, PA: University of Scranton Press, 1996.
- Gibbon, Edward, ‘Decline and Fall of the Roman Empire’, Chapter 14
- Harries, Jill. "Law and Empire in Late Antiquity". Cambridge: Cambridge University
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Press, 1999. Hardcover Paperback
- Helgeland, John. "Christians and the Roman Army A.D. 173–337." "Church History" 43:2 (1974): 149–163, 200.
- Jones, A.H.M. "The Later Roman Empire, 284–602: A Social, Economic and Administrative Survey". Baltimore: Johns Hopkins University, 1986.
- Leadbetter, Bill. "Galerius and the Will of Diocletian", London and New York, Routledge, 2010. Hardcover Paperback
- Lenski, Noel. "The Reign of Constantine." In "The Cambridge Companion to the Age of Constantine", edited by Noel Lenski, 59–90. New York: Cambridge University Press, 2006b. Hardcover Paperback
- Mackay, Christopher S. "Lactantius and the Succession to Diocletian." "Classical Philology" 94:2 (1999):
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198–209.
- Mathisen, Ralph W. "Diocletian (284–305 AD.)." "De Imperatoribus Romanis" (1997). Accessed February 16, 2008.
- Odahl, Charles Matson. "Constantine and the Christian Empire". New York: Routledge, 2004. Hardcover Paperback
- Pohlsander, Hans. "The Emperor Constantine". London & New York: Routledge, 2004a. Hardcover Paperback
- Pohlsander, Hans. "Constantine I (306 – 337 AD)." "De Imperatoribus Romanis" (2004b). Accessed December 16, 2007.
- Potter, David S. "The Roman Empire at Bay: AD 180–395". New York: Routledge, 2005. Hardcover Paperback
- Southern, Pat. "The Roman Empire from Severus to Constantine". New York: Routledge, 2001.
- Rostovtzeff, Michael. "The Social and Economic
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D 180–395". New York: Routledge, 2005. Hardcover Paperback
- Southern, Pat. "The Roman Empire from Severus to Constantine". New York: Routledge, 2001.
- Rostovtzeff, Michael. "The Social and Economic History of the Roman Empire". Oxford: Oxford University Press, 1966.
- Treadgold, Warren. "A History of the Byzantine State and Society". Stanford: Stanford University Press, 1997.
- Williams, Stephen. "Diocletian and the Roman Recovery". New York: Routledge, 1997.
# External links.
- Medieval Sourcebook: Edict of Toleration by Galerius, 311.
- Gaius Galerius Valerius Maximianus
- Lactantius about Galerius in his ""De Mortibus Persecutorum"" chapter XXIII & XXVII
- Catholic Encyclopedia
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Dendritic spine
A dendritic spine (or spine) is a small membranous protrusion from a neuron's dendrite that typically receives input from a single axon at the synapse. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical signals to the neuron's cell body. Most spines have a bulbous head (the spine head), and a thin neck that connects the head of the spine to the shaft of the dendrite. The dendrites of a single neuron can contain hundreds to thousands of spines. In addition to spines providing an anatomical substrate for memory storage and synaptic transmission, they may also serve to increase the number of possible contacts between neurons.
# Structure.
Dendritic
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spines are small with spine head volumes ranging 0.01 µm to 0.8 µm. Spines with strong synaptic contacts typically have a large spine head, which connects to the dendrite via a membranous neck. The most notable classes of spine shape are "thin", "stubby", "mushroom", and "branched". Electron microscopy studies have shown that there is a continuum of shapes between these categories. The variable spine shape and volume is thought to be correlated with the strength and maturity of each spine-synapse.
## Distribution.
Dendritic spines usually receive excitatory input from axons, although sometimes both inhibitory and excitatory connections are made onto the same spine head. Excitatory axon proximity
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to dendritic spines is not sufficient to predict the presence of a synapse, as demonstrated by the Lichtman lab in 2015.
Spines are found on the dendrites of most principal neurons in the brain, including the pyramidal neurons of the neocortex, the medium spiny neurons of the striatum, and the Purkinje cells of the cerebellum. Dendritic spines occur at a density of up to 5 spines/1 µm stretch of dendrite. Hippocampal and cortical pyramidal neurons may receive tens of thousands of mostly excitatory inputs from other neurons onto their equally numerous spines, whereas the number of spines on Purkinje neuron dendrites is an order of magnitude larger.
## Cytoskeleton and organelles.
The cytoskeleton
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of dendritic spines is particularly important in their synaptic plasticity; without a dynamic cytoskeleton, spines would be unable to rapidly change their volumes or shapes in responses to stimuli. These changes in shape might affect the electrical properties of the spine. The cytoskeleton of dendritic spines is primarily made of filamentous actin (F-actin). tubulin Monomers and microtubule-associated proteins (MAPs) are present, and organized microtubules are present. Because spines have a cytoskeleton of primarily actin, this allows them to be highly dynamic in shape and size. The actin cytoskeleton directly determines the morphology of the spine, and actin regulators, small GTPases such as
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Rac, RhoA, and CDC42, rapidly modify this cytoskeleton. Overactive Rac1 results in consistently smaller dendritic spines.
In addition to their electrophysiological activity and their receptor-mediated activity, spines appear to be vesicularly active and may even translate proteins. Stacked discs of the smooth endoplasmic reticulum (SERs) have been identified in dendritic spines. Formation of this "spine apparatus" depends on the protein synaptopodin and is believed to play an important role in calcium handling. "Smooth" vesicles have also been identified in spines, supporting the vesicular activity in dendritic spines. The presence of polyribosomes in spines also suggests protein translational
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activity in the spine itself, not just in the dendrite.
# Physiology.
## Receptor activity.
Dendritic spines express glutamate receptors (e.g. AMPA receptor and NMDA receptor) on their surface. The TrkB receptor for BDNF is also expressed on the spine surface, and is believed to play a role in spine survival. The tip of the spine contains an electron-dense region referred to as the "postsynaptic density" (PSD). The PSD directly apposes the active zone of its synapsing axon and comprises ~10% of the spine's membrane surface area; neurotransmitters released from the active zone bind receptors in the postsynaptic density of the spine. Half of the synapsing axons and dendritic spines are physically
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tethered by calcium-dependent cadherin, which forms cell-to-cell adherent junctions between two neurons.
Glutamate receptors (GluRs) are localized to the postsynaptic density, and are anchored by cytoskeletal elements to the membrane. They are positioned directly above their signalling machinery, which is typically tethered to the underside of the plasma membrane, allowing signals transmitted by the GluRs into the cytosol to be further propagated by their nearby signalling elements to activate signal transduction cascades. The localization of signalling elements to their GluRs is particularly important in ensuring signal cascade activation, as GluRs would be unable to affect particular downstream
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effects without nearby signallers.
Signalling from GluRs is mediated by the presence of an abundance of proteins, especially kinases, that are localized to the postsynaptic density. These include calcium-dependent calmodulin, CaMKII (calmodulin-dependent protein kinase II), PKC (Protein Kinase C), PKA (Protein Kinase A), Protein Phosphatase-1 (PP-1), and Fyn tyrosine kinase. Certain signallers, such as CaMKII, are upregulated in response to activity.
Spines are particularly advantageous to neurons by compartmentalizing biochemical signals. This can help to encode changes in the state of an individual synapse without necessarily affecting the state of other synapses of the same neuron. The
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length and width of the spine neck has a large effect on the degree of compartmentalization, with thin spines being the most biochemically isolated spines.
## Plasticity.
Dendritic spines are very "plastic", that is, spines change significantly in shape, volume, and number in small time courses. Because spines have a primarily actin cytoskeleton, they are dynamic, and the majority of spines change their shape within seconds to minutes because of the dynamicity of actin remodeling. Furthermore, spine number is very variable and spines come and go; in a matter of hours, 10-20% of spines can spontaneously appear or disappear on the pyramidal cells of the cerebral cortex, although the larger "mushroom"-shaped
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spines are the most stable.
Spine maintenance and plasticity is activity-dependent and activity-independent. BDNF partially determines spine levels, and low levels of AMPA receptor activity is necessary to maintain spine survival, and synaptic activity involving NMDA receptors encourages spine growth. Furthermore, two-photon laser scanning microscopy and confocal microscopy have shown that spine volume changes depending on the types of stimuli that are presented to a synapse.
## Importance to learning and memory.
### Evidence of importance.
Spine plasticity is implicated in motivation, learning, and memory. In particular, long-term memory is mediated in part by the growth of new dendritic
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spines (or the enlargement of pre-existing spines) to reinforce a particular neural pathway. Because dendritic spines are plastic structures whose lifespan is influenced by input activity, spine dynamics may play an important role in the maintenance of memory over a lifetime.
Age-dependent changes in the rate of spine turnover suggest that spine stability impacts developmental learning. In youth, dendritic spine turnover is relatively high and produces a net loss of spines. This high rate of spine turnover may characterize critical periods of development and reflect learning capacity in adolescence—different cortical areas exhibit differing levels of synaptic turnover during development, possibly
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reflecting varying critical periods for specific brain regions. In adulthood, however, most spines remain persistent, and the half-life of spines increases. This stabilization occurs due to a developmentally regulated slow-down of spine elimination, a process which may underlie the stabilization of memories in maturity.
Experience-induced changes in dendritic spine stability also point to spine turnover as a mechanism involved in the maintenance of long-term memories, though it is unclear how sensory experience affects neural circuitry. Two general models might describe the impact of experience on structural plasticity. On the one hand, experience and activity may drive the discrete formation
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of relevant synaptic connections that store meaningful information in order to allow for learning. On the other hand, synaptic connections may be formed in excess, and experience and activity may lead to the pruning of extraneous synaptic connections.
In lab animals of all ages, environmental enrichment has been related to dendritic branching, spine density, and overall number of synapses. In addition, skill training has been shown to lead to the formation and stabilization of new spines while destabilizing old spines, suggesting that the learning of a new skill involves a rewiring process of neural circuits. Since the extent of spine remodeling correlates with success of learning, this suggests
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a crucial role of synaptic structural plasticity in memory formation. In addition, changes in spine stability and strengthening occur rapidly and have been observed within hours after training.
Conversely, while enrichment and training are related to increases in spine formation and stability, long-term sensory deprivation leads to an increase in the rate of spine elimination and therefore impacts long-term neural circuitry. Upon restoring sensory experience after deprivation in adolescence, spine elimination is accelerated, suggesting that experience plays an important role in the net loss of spines during development. In addition, other sensory deprivation paradigms—such as whisker trimming—have
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been shown to increase the stability of new spines.
Research in neurological diseases and injuries shed further light on the nature and importance of spine turnover. After stroke, a marked increase in structural plasticity occurs near the trauma site, and a five- to eightfold increase from control rates in spine turnover has been observed. Dendrites disintegrate and reassemble rapidly during ischemia—as with stroke, survivors showed an increase in dendritic spine turnover. While a net loss of spines is observed in Alzheimer's disease and cases of intellectual disability, cocaine and amphetamine use have been linked to increases in dendritic branching and spine density in the prefrontal cortex
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and the nucleus accumbens. Because significant changes in spine density occur in various brain diseases, this suggests a balanced state of spine dynamics in normal circumstances, which may be susceptible to disequilibrium under varying pathological conditions.
There is also some evidence for loss of dendritic spines as a consequence of aging. One study using mice has noted a correlation between age-related reductions in spine densities in the hippocampus that and age-dependent declines in hippocampal learning and memory.
### Importance contested.
Despite experimental findings that suggest a role for dendritic spine dynamics in mediating learning and memory, the degree of structural plasticity’s
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importance remains debatable. For instance, studies estimate that only a small portion of spines formed during training actually contribute to lifelong learning. In addition, the formation of new spines may not significantly contribute to the connectivity of the brain, and spine formation may not bear as much of an influence on memory retention as other properties of structural plasticity, such as the increase in size of spine heads.
# Modeling.
Theoreticians have for decades hypothesized about the potential electrical function of spines, yet our inability to examine their electrical properties has until recently stopped theoretical work from progressing too far. Recent advances in imaging
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techniques along with increased use of two-photon glutamate uncaging have led to a wealth of new discoveries; we now suspect that there are voltage-dependent sodium, potassium, and calcium channels in the spine heads.
Cable theory provides the theoretical framework behind the most "simple" method for modelling the flow of electrical currents along passive neural fibres. Each spine can be treated as two compartments, one representing the neck, the other representing the spine head. The compartment representing the spine head alone should carry the active properties.
## Baer and Rinzel's continuum model.
To facilitate the analysis of interactions between many spines, Baer & Rinzel formulated
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a new cable theory for which the distribution of spines is treated as a continuum. In this representation, spine head voltage is the local spatial average of membrane potential in adjacent spines. The formulation maintains the feature that there is no direct electrical coupling between neighboring spines; voltage spread along dendrites is the only way for spines to interact.
## Spike-diffuse-spike model.
The SDS model was intended as a computationally simple version of the full Baer and Rinzel model. It was designed to be analytically tractable and have as few free parameters as possible while retaining those of greatest significance, such as spine neck resistance. The model drops the continuum
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approximation and instead uses a passive dendrite coupled to excitable spines at discrete points. Membrane dynamics in the spines are modelled using integrate and fire processes. The spike events are modelled in a discrete fashion with the wave form conventionally represented as a rectangular function.
## Modeling spine calcium transients.
Calcium transients in spines are a key trigger for synaptic plasticity. NMDA receptors, which have a high permeability for calcium, only conduct ions if the membrane potential is suffiently depolarized. The amount of calcium entering a spine during synaptic activity therefore depends on the depolarization of the spine head. Evidence from calcium imaging
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experiments (two-photon microscopy) and from compartmental modelling indicates that spines with high resistance necks experience larger calcium transients during synaptic activity.
# Development.
Dendritic spines can develop directly from dendritic shafts or from dendritic filopodia. During synaptogenesis, dendrites rapidly sprout and retract filopodia, small membrane organelle-lacking membranous protrusions. Recently, I-BAR protein MIM was found to contribute to the initiation process. During the first week of birth, the brain is predominated by filopodia, which eventually develop synapses. However, after this first week, filopodia are replaced by spiny dendrites but also small, stubby spines
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that protrude from spiny dendrites. In the development of certain filopodia into spines, filopodia recruit presynaptic contact to the dendrite, which encourages the production of spines to handle specialized postsynaptic contact with the presynaptic protrusions.
Spines, however, require maturation after formation. Immature spines have impaired signaling capabilities, and typically lack "heads" (or have very small heads), only necks, while matured spines maintain both heads and necks.
# Clinical significance.
Cognitive disorders such as ADHD, autism, intellectual disability, and fragile X syndrome, may be resultant from abnormalities in dendritic spines, especially the number of spines and
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their maturity. The ratio of matured to immature spines is important in their signaling, as immature spines have impaired synaptic signaling. Fragile X syndrome is characterized by an overabundance of immature spines that have multiple filopodia in cortical dendrites.
# History.
Dendritic spines were first described at the end of the 19th century by Santiago Ramón y Cajal on cerebellar neurons. Ramón y Cajal then proposed that dendritic spines could serve as contacting sites between neurons. This was demonstrated more than 50 years later thanks to the emergence of electron microscopy. Until the development of confocal microscopy on living tissues, it was commonly admitted that spines were
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