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Olive Branch Petition
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The Olive Branch Petition
The Olive Branch Petition was adopted by the Second Continental Congress on July 5, 1775 in a final attempt to avoid a full-on war between Great Britain and the thirteen colonies represented in that Congress. The Congress had already authorized the invasion of Canada more than a week earlier, but the petition affirmed American loyalty to Great Britain and beseeched King George III to prevent further conflict. That the petition was followed by the July 6 Declaration of the Causes and Necessity of Taking Up Arms made its success in London improbable.[1] In August 1775, the colonies were formally declared to be in rebellion by the Proclamation of Rebellion, and the petition was rejected by Great Britain—even though King George had refused to read the Olive Branch Petition before declaring the colonists traitors.[2]
Contents
[hide]
1 Drafting
2 Reception and rejection
3 Consequences
4 References
5 External links
Drafting[edit]
The Second Continental Congress convened in May 1775, and most delegates followed John Dickinson in his quest to reconcile with King George III of Great Britain. However, a rather small group of delegates led by John Adams believed that war was inevitable. During the course of the Second Continental Congress, Adams and his allies decided that the wisest course of action was to remain quiet and wait for the opportune time to rally the people.
This decision allowed Dickinson and his followers to pursue their own course for reconciliation.[3] Dickinson was the primary author of the petition, though Benjamin Franklin, John Jay, John Rutledge and Thomas Johnson also served on the drafting committee.[4] Dickinson claimed that the colonies did not want independence but wanted to more equitable trade and tax regulations. He suggested that the King devise a plan to settle trade disputes and give the colonists either free trade and taxes equal to those levied on the people of Great Britain or strict trade regulation in lieu of taxes. The introductory paragraph of the letter named twelve of the thirteen colonies, all except Georgia. The letter was approved on July 5 and signed by John Hancock, President of the Second Congress, and by representatives of the named twelve colonies. It was sent to London on July 8, 1775, in the care of Richard Penn and Arthur Lee.[5] Dickinson hoped that news of the Battles of Lexington and Concord combined with the "humble petition" would persuade the King to respond with a counter proposal or to open negotiations.[3]
Reception and rejection[edit]
In a letter to a friend, Adams wrote that the petition served no purpose, that war was inevitable and that the colonies should have already raised a navy and taken British officials prisoner. His letter was intercepted by British officials and news of its contents reached Great Britain at about the same time as the petition itself. British advocates of a military response to the colonists used Adams' letter to claim that the petition itself was insincere.[5]
On August 21, Penn and Lee provided a copy of the petition to Lord Dartmouth, the colonial secretary, followed with the original on September 1. Penn and Lee reported back on September 2 that "...we were told that as his Majesty did not receive it on the throne, no answer would be given."[6] In response to the news of the Battle of Bunker Hill, the King had already issued the Proclamation for Suppressing Rebellion and Sedition on August 23, declaring the North American colonies in a state of rebellion and ordering "all Our officers...and all Our obedient and loyal subjects, to use their utmost endeavours to withstand and suppress such rebellion."[7] The hostilities Adams had foreseen undercut the petition, which the King had answered even before it reached him.[8]
Consequences[edit]
The King’s refusal to consider the petition gave Adams and others the opportunity to push for independence and to characterize the King as intransigent and uninterested in addressing the colonists' grievances. It polarized the issue in the minds of many colonists, who realized that, from that point forward, the choice was between complete independence or complete submission to British rule,[5] a realization crystallized a few months later in Thomas Paine's widely read pamphlet Common Sense.
References[edit]
Jump up ^ "Declaration of taking up arms: resolutions of the Second Continental Congress". Constitution Society. Retrieved September 23, 2013.
Jump up ^ Bailey, Thomas; Kennedy, David; Cohen, Lizabeth (1998). The American Pageant (11 ed.). New York: Houghton Mifflin Company.
^ Jump up to: a b Ferling, John E (2003). A leap in the dark: the struggle to create the American republic. Oxford, England; New York: Oxford University Press.
Jump up ^ Beeman, Richard (2013). Our lives, our fortune, our sacred honor: the forging of American independence, 1774–1776. New York: Basic Books. ISBN 9780465026296.
^ Jump up to: a b c Brown, Weldon A. (1941). Empire or independence; a study in the failure of reconciliation, 1774–1783. Port Washington, New York: Kennikat Press (published 1966). OCLC 341868.
Jump up ^ Richard Penn; Arthur Lee. "Petition to George III, King of Great Britain, 1775". nypl.org. Image 5208532. Retrieved 3 October 2017.
Jump up ^ Axelrod, Alan (2008). Profiles in folly: history's worst decisions and why they went wrong. New York: Sterling. p. 150. ISBN 1402747683. [unreliable source?]
Jump up ^ Maier, Pauline (1997). American scripture: making the Declaration of Independence. New York: Knopf. pp. 24–25, 249–250. ISBN 0679454926.
External links[edit]
Works related to Olive Branch Petition at Wikisource
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Rules of cribbage
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Over the 400 or so years that the game of cribbage has been around, variations and rules have evolved. House rules, preferences and even misunderstandings have been passed from one player to another. However, the core of the rules remains constant, and should be familiar to most players. The rules here apply to the standard two-, three- or four-player games, with details of variations being listed below.
Contents
[hide]
1 The deal
2 The crib
2.1 Example cribs
2.1.1 Two players
2.1.2 Three players
3 The starter
4 The play
4.1 Example plays
4.1.1 Two players
4.1.2 Three players
5 The show
5.1 Example scores
5.1.1 Two players
5.1.2 Three players
6 The end
7 Match
7.1 Two player game
7.2 Three player game
7.2.1 Winner takes all
7.2.2 Continued play
8 Variations
9 References
10 External links
The deal[edit]
Cribbage uses a standard 52-card deck of cards. The jokers are removed; the suits are equal in status. The players cut for first deal, with the player cutting the lowest card (the ace counts as one, and is the lowest card) dealing first. If the cutters tie, the cards are re-shuffled and re-cut. The dealer shuffles, offers the deck to the player on his or her right to cut (required in tournament play), and deals cards singly to each player, starting with the player on the dealer's left. During the deal, if any card is exposed by the dealer or found face-up in the deck, cards must be redealt.[1]
Cards must be dealt so that each player ends up with four cards after the crib is formed, and the crib should also have four cards. For two players, each is dealt six cards (though some play with five cards dealt to each player and two to the crib). For three or four players, each is dealt five cards. In the case of three players, a single card is dealt face down in front of the dealer to start the crib.
# of Players
# cards dealt to each player
# cards dealt to crib
# cards discarded by each player to crib
2 (standard) 6 0 2
2 (alternate) 5 2 1
3 5 1 1
4 5 0 1
The crib[edit]
Once the cards have been dealt, each player chooses four cards to retain, discarding the other one or two face-down to form the "crib" that will be used later by the dealer.[2] At this point, each player's hand and the crib will contain exactly four cards.
Example cribs[edit]
Two players[edit]
Player
Cards dealt
Discarded
Hand
Alice
(dealer)
Bob
Crib
Three players[edit]
Player
Cards dealt
Discarded
Hand
Claire
(dealer)
David
Eve
Crib
The starter[edit]
The player on the dealer's left cuts the undealt portion of the deck (leaving at least 4 cards), and the dealer reveals the top card, called the "starter" or the "cut", placing it on top of the deck face up. (It is illegal to peek at any other cards in the deck during this process.)[3] If this card is a Jack, the dealer scores two points for "his heels", also known as "his nibs", or simply "nibs". The game can end on a cut of a Jack for the dealer.
The play[edit]
The play (often called pegging) starts with the player on the dealer's left and continues clockwise. Each player lays one card in turn onto the table so that it is visible, stating the cumulative value of the cards played. (For example, the first player lays a 4 and says "four", the next lays a 7 and says "eleven", and so on). Face cards are worth ten; Aces are worth one. Play must not exceed 31, so a player who cannot lay a card without bringing the total above 31 passes by saying "Go". The other players continue to lay cards in turn without exceeding 31 until no cards can be played. If a player can lay a card they must do so. The last player to lay a card scores two points if 31 is reached exactly ("31 for two"); otherwise one point is scored, e.g., "29 for one", or "30 for one", etc. The one-point score is known as "One for the go",[4] or simply the "Go". The count is then reset to zero and play resumes, starting with the player to the left of the last card played. Players with cards remaining repeat this process until all players' cards have been played.
In addition to scoring one or two points for the last card, players score points according to the following rules:
fifteen-twos
two points for a cumulative total of exactly fifteen ("fifteen two")
runs
three points for completing a run of three cards, regardless of the order in which they are laid (a 6, then a 4, then a 5 is a run of three even though they were not laid in order)
four points for completing a run of four
five points for completing a run of five
six points for completing a run of six
seven points for completing the run of seven; e.g. playing 2, 4, 6, A, 3, 5 and 7
pairs
two points for laying a card of the same rank as the previous card, thus completing a pair
six points for laying a third card of the same rank (a "pair royal" or "trips")
twelve points for laying a fourth card of the same rank (a "double pair royal" or "quad")
If a card completes more than one scoring combination, then all combinations are scored. For example, if the first three cards played are 5s, the third one scores eight points: two for making 15, and six for a pair royal. Card combinations cannot span a reset; once the total reaches 31 (or a Go has been scored) and counting has restarted at zero, cards already played cannot contribute to runs or pairs. During this phase of play run combinations cannot span a pair; in a play of 2, 3, 3, 4 the pair interrupts the run so only the pair is counted for points.
Players choose when to lay each card in order to maximise their score according to the scheme shown below. The first player to reach 121 wins the game.
Example plays[edit]
Two players[edit]
Alice
(dealer)
Bob
Player
Card
Cumulative
Score
Announced
Bob 10 "ten"
Alice 15 2 points (fifteen exactly) "fifteen for two"
Bob 22 "twenty-two"
Alice 28 3 points (run: 5, 6, 7) "twenty-eight for three"
Bob "go"
Alice 30 1 point "thirty for one"
Bob 6 "six"
Alice 10 "ten"
Bob 14 3 points (pair, last card) "fourteen for three"
Three players[edit]
Claire
(dealer)
David
Eve
Player
Card
Cumulative
Score
Announced
David 7 "seven"
Eve 14 2 points (pair) "fourteen for two"
Claire 21 6 points (three-of-a-kind) "twenty-one for six"
David 26 "twenty-six"
Eve 31 4 points (pair, 31 exactly) "thirty-one for four"
Claire 8 "eight"
David 18 "eighteen"
Eve 28 2 points (pair) "twenty-eight for two"
Claire "go"
David "go"
Eve 31 2 points (31 exactly) "thirty-one for two"
Claire 10 "ten"
David 14 "fourteen"
Claire 23 1 point (last card) "twenty-three for one"
The show[edit]
Once the play is complete, each player in turn receives points based on the content of their hand. Starting with the player on the dealer's left, players spread out their cards on the playing surface and calculate their score based on these four cards and the starter card:
fifteen-twos
two points for each separate combination of two or more cards totalling exactly fifteen
runs
three points for a run of three consecutive cards (regardless of suit)
four points for completing a run of four
five points for completing a run of five
pairs
two points for a pair of cards of a kind
six points for three cards of a kind (known as a "pair royal", comprising three distinct pairs)
twelve points for four cards of a kind (a "double pair royal", comprising six distinct pairs)
flush
four points for a flush, where all four cards in the hand are of the same suit, with an additional point if the starter card is also of that suit.
one point for holding the Jack of the same suit as the starter card ("one for his nob"[5] or "... his nobs"[6])
Common combinations are often scored as a group. For example, a run of three cards with an additional card matching one of the three in value, e.g., 2–2–3–4, is termed a "double run", and scores eight according to the above rules, three for each of the runs and two for the pair. Even more valuable are "triple runs" (three three-card straights including a three-of-a-kind, e.g., 2–2–2–3–4, that score fifteen) and "double-double runs" (four three-card straights including pairs, e.g., 2–3–3–4–4, that score sixteen). Combined runs may also include fifteen-twos. A 24 hand, the largest commonly seen, comprises a double-double run and four fifteen-twos. Two examples are 4–4–5–5–6 and 6–7–7–8–8.
The dealer scores his hand last and then turns the cards in the crib face up. These cards, in conjunction with the starter card, are scored by the dealer as an additional hand. The rules for scoring the crib are the same as scoring a hand, with the exception of the flush; a four-card flush in the crib is scored only if it is the same suit as the starter card (for a total of five points).
The highest possible score for a hand is 29 points: a starter card of a 5, and a hand of 5, 5, 5, J with the Jack being the same suit as the starter card. The score might be announced thus:
Fifteen two, fifteen four, fifteen six, fifteen eight, [four J-5 combinations]
fifteen ten, fifteen twelve, fifteen fourteen, fifteen sixteen, [four 5–5–5 combinations]
twelve for pairs [four 5s] is twenty-eight,
and nobs makes twenty-nine.
Scores between 0 and 29 are all possible, with the exception of 19, 25, 26 and 27.[7] Players may colloquially refer to a blank hand (one scoring no points) as a "nineteen hand".
Example scores[edit]
Two players[edit]
Starter
card
Alice
(dealer)
Bob
Crib
Player
Cards
Score
Announced
Bob
2 "fifteen two"
2 "fifteen four"
4 "four for a run"
1 "and one for his nob"
Total 9
Alice
2 "fifteen two"
2 "fifteen four"
8 "and eight for a double run"
Total 12
Alice
(crib)
2 "fifteen two"
2 "fifteen four"
2 "and two for a pair"
Total 6
Three players[edit]
Starter
card
Claire
(dealer)
David
Eve
Crib
Player
Cards
Score
Announced
David
2 "fifteen two"
2 "fifteen four"
3 "and three for a run"
Total 7
Eve
2 "fifteen two"
2 "fifteen four"
2 "fifteen six"
2 "and two for a pair"
Total 8
Claire
2 "fifteen two"
3 "and three for a run"
Total 5
Claire
(crib)
2 "fifteen two"
3 "and three for a run"
Total 5
The end[edit]
After the dealer has scored the crib, all cards are collected and the deal passes to the player on the dealer's left. The next round starts with the deal.
Although the rules of cribbage do not require it (except in tournament play), the traditional method of keeping score is to use a cribbage board. This is a flat board, usually made of wood, with separate series of holes that record each player's score. It is usually arranged in five hole sections for easier scoring. Players each have two pegs that mark their current and previous scores, and all scoring is done by moving the back peg ahead of the front peg.
When a player reaches the target score for the game (usually 121), the game ends with that player the winner.
Match[edit]
A match (much like tennis) consists of more than one game, often an odd number (3 games, 5 games, 7 games etc.). The match points are scored on the cribbage board using the holes reserved for match points. On a spiral board, these are often at the bottom of the board in a line with 5 or 7 holes. On a traditional board, they are often placed in the middle of the board or at the top/bottom.
Two player game[edit]
In a two player game of cribbage a player scores one match point for each game won. Their opponent will begin the next game as first dealer. If a player skunks their opponent (reaches 121 points before their opponent scores 91 points) then that player scores one extra match point for that game (two match points in total). If a player double skunks their opponent (reaches 121 points before their opponent reaches 61), then they score two extra match points for the game (four match points in total). If a player triple skunks their opponent (reaches 121 points before their opponent reaches 31 points), they automatically win the match regardless of how many match points are needed to win. Double and triple skunks are not included in the official rules of cribbage play and are optional. There are several different formats for scoring match points.
Match point scoring
Scoring Variation
Points for a normal win
Points for skunking opponent
Points for double skunking opponent
Points for triple skunking opponent
Official Tournament rules 2 points 3 points no extra points no extra points
Variation of Tournament rules 2 points 3 points 4 points no extra points
Free play rules 1 point 2 points 4 points no extra points
Free play rules with triple skunk 1 points 2 points 4 points Automatic win of match
Three player game[edit]
Winner takes all[edit]
When playing a three player match in a winner takes all format, the winner scores two match points (just like in two-way cribbage) for each game won. If he/she skunks just the third opponent, they score an additional match point (3 total) with second place receiving one point. If he/she double skunks both opponents, he/she still scores three match points but second place would not receive any points at all.
Continued play[edit]
In continued play format, the winner of the match earns two match points for three player cribbage and four match points for five player cribbage (plus applicable match points if the player has skunked/double skunked their opponents). The remaining players play until there is a second winner, who scores one match point for three player cribbage and two match points for five player cribbage (with no extra points for skunking opponents). In five player cribbage, the remaining three players play until there is a third winner, who scores one match point (with no extra points for skunking opponents).
Variations[edit]
Three players: Five cards are dealt to each player and one card directly to the crib, and each player then discards one card to the crib, as shown in the examples above. Three players can score individually, with the winner the first to reach 121, or in a "two against one" team format, where the two-player team must score 121 to win before the lone player reaches 61. Another variation of the "two against one" team format, is that prior to the cut, the lone player picks up the crib, examines all 8 cards, and then discards 4 cards to the crib. Both the team and lone player need to reach 121 to win.
Four players: Five cards are dealt to each player, each of whom discards one to the crib. The players can play as individuals or as two sets of partners.
Five-card cribbage (called the "old game"): Each of two players is dealt five cards, two of which are discarded into the crib. The crib thus consists of four cards but each hand only three. The non-dealer is given a three-point start and play is to 61.
Five players: Five cards are dealt to each player except the dealer, who has only four cards. The four non-dealers each discard one card to the crib.
Muggins: This is a scoring variant in which a player who fails to count all the points to which he is entitled in the play or the show loses the unclaimed points to an opponent who calls "muggins" or "cut throat".
Lowball (or "Loser's Crib"): This is a misère variant in which the normal rules apply but the aim is to avoid scoring. The loser is the first to 121.
Jokers: Jokers are fully wild, with their rank and suit decided only at the moment of play. The choice of card may even replicate a card already in play, allowing for 5 of a kind (20 points), 6 of a kind (30 points), etc. When a joker is cut as the starter, the dealer scores 2 for heels and each player may choose a different rank and suit for the joker when hands are scored.
Toss Fives: This is a variant in which players must discard any 5s they may have into the crib (even an opponent's crib).
Auction Cribbage:[8] In Auction Cribbage, any player may bid for the points in the box after the cards are dealt. Bidding continues in turn until no further bids are offered; the winning bidder then immediately deducts that number of points from their hand; the crib is scored at the usual time and its points awarded to the winning bidder for that round. If no bid is placed, the dealer retains the crib.
Null point penalty:[citation needed] When a player scores zero points during "the show", their opponent scores one point. This applies to both players hands as well as in the crib.
Back 10 (Backup Ten): The hand and the crib must contain points. If either hand does not, the owner of the hand must go back ten points.[9]
References[edit]
Jump up ^ "Cribbage Official Tournament Rules" (PDF). 4.3 Exposed Cards. American Cribbage Congress. August 2010. p. 13. Retrieved April 11, 2012.
Jump up ^ Parlett, David (October 1987). The Penguin Book of Card Games. Treasure Press. ISBN 1-85051-221-3.
Jump up ^ "The Mechanics of Playing Cribbage". The American Cribbage Congress. 2004-10-01. Retrieved 2008-03-01.
Jump up ^ "Cribbage Rules". Cribbage Corner. 2008-05-01. Retrieved 2008-05-05.
Jump up ^ Block, Mike; et al. "Six Card Cribbage". Card Games. Pagat.com. Retrieved 9 July 2010.
Jump up ^ "Rule 1. The Mechanics of Playing Cribbage: Definitions". Cribbage Official Tournament Rules. American Cribbage Congress. Retrieved 9 July 2010.
Jump up ^ Steven S. Lumetta (2007-05-15). "Amusing Cribbage Facts". Archived from the original on 2008-05-16. Retrieved 2008-03-03.
Jump up ^ http://www.poker.tj/games-encyclopedia/strategy-cribbage-poker.html. Missing or empty |title= (help)
Jump up ^ http://www.ecribbage.com
External links[edit]
Cribbage rules from Cribbage Corner
Official tournament rules of the American Cribbage Congress
Cribbage Scoring Chart
Retrieved from "https://en.wikipedia.org/w/index.php?title=Rules_of_cribbage&oldid=838561282"
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Vestibular system
From Wikipedia, the free encyclopedia
Jump to: navigation, search
The vestibular system, in most mammals, is the sensory system that provides the leading contribution to the sense of balance and spatial orientation for the purpose of coordinating movement with balance. Together with the cochlea, a part of the auditory system, it constitutes the labyrinth of the inner ear in most mammals. As movements consist of rotations and translations, the vestibular system comprises two components: the semicircular canals, which indicate rotational movements; and the otoliths, which indicate linear accelerations. The vestibular system sends signals primarily to the neural structures that control eye movements, and to the muscles that keep an animal upright. The projections to the former provide the anatomical basis of the vestibulo-ocular reflex, which is required for clear vision; and the projections to the muscles that control posture are necessary to keep an animal upright.
The brain uses information from the vestibular system in the head and from proprioception throughout the body to understand the body's dynamics and kinematics (including its position and acceleration) from moment to moment.
Contents
[hide]
1 Semicircular canal system
1.1 Structure
1.2 Push-pull systems
1.3 Vestibulo-ocular reflex (VOR)
1.4 Mechanics
1.5 Central processing
1.6 Projection pathways
2 Otolithic organs
3 Experience from the vestibular system
3.1 Vestibular/somatogyral illusions
4 Pathologies
5 See also
6 References
7 Further reading
8 External links
Semicircular canal system[edit]
Cochlea and vestibular system
The semicircular canal system detects rotational movements. The semicircular canals are its main tools to achieve this detection.
Structure[edit]
Main article: Semicircular canal
Since the world is three-dimensional, the vestibular system contains three semicircular canals in each labyrinth. They are approximately orthogonal (right angles) to each other, and are called the horizontal (or lateral), the anterior semicircular canal (or superior) and the posterior (or inferior) semicircular canal. Anterior and posterior canals may be collectively called vertical semicircular canals.
Movement of fluid within the horizontal semicircular canal corresponds to rotation of the head around a vertical axis (i.e. the neck), as when doing a pirouette.
The anterior and posterior semicircular canals detect rotations of the head in the sagittal plane (as when nodding), and in the frontal plane, as when cartwheeling. Both anterior and posterior canals are orientated at approximately 45° between frontal and sagittal planes.
The movement of fluid pushes on a structure called the cupula, which contains hair cells that transduce the mechanical movement to electrical signals[1]
Push-pull systems[edit]
Push-pull system of the semicircular canals, for a horizontal head movement to the right.
The canals are arranged in such a way that each canal on the left side has an almost parallel counterpart on the right side. Each of these three pairs works in a push-pull fashion: when one canal is stimulated, its corresponding partner on the other side is inhibited, and vice versa.
This push-pull system makes it possible to sense all directions of rotation: while the right horizontal canal gets stimulated during head rotations to the right (Fig 2), the left horizontal canal gets stimulated (and thus predominantly signals) by head rotations to the left.
Vertical canals are coupled in a crossed fashion, i.e. stimulations that are excitatory for an anterior canal are also inhibitory for the contralateral posterior, and vice versa.
Vestibulo-ocular reflex (VOR)[edit]
The vestibulo-ocular reflex. A rotation of the head is detected, which triggers an inhibitory signal to the extraocular muscles on one side and an excitatory signal to the muscles on the other side. The result is a compensatory movement of the eyes.
Main article: Vestibulo-ocular reflex
The vestibulo-ocular reflex (VOR) is a reflex eye movement that stabilizes images on the retina during head movement by producing an eye movement in the direction opposite to head movement, thus preserving the image on the center of the visual field. For example, when the head moves to the right, the eyes move to the left, and vice versa. Since slight head movements are present all the time, the VOR is very important for stabilizing vision: patients whose VOR is impaired find it difficult to read, because they cannot stabilize the eyes during small head tremors. The VOR reflex does not depend on visual input and works even in total darkness or when the eyes are closed.
This reflex, combined with the push-pull principle described above, forms the physiological basis of the Rapid head impulse test or Halmagyi-Curthoys-test, in which the head is rapidly and forcefully moved to the side, while observing whether the eyes keep looking in the same direction.
Mechanics[edit]
The mechanics of the semicircular canals can be described by a damped oscillator.[citation needed] If we designate the deflection of the cupula with θ {\displaystyle \theta } , and the head velocity with q ˙ {\displaystyle {\dot {q}}} , the cupula deflection is approximately[citation needed]
θ ( s ) = α s ( T 1 s + 1 ) ( T 2 s + 1 ) q ˙ ( s ) {\displaystyle \theta (s)={\frac {\alpha s}{(T_{1}s+1)(T_{2}s+1)}}{\dot {q}}(s)}
α is a proportionality factor, and s corresponds to the frequency. For humans, the time constants T1 and T2 are approximately 3 ms and 5 s, respectively[citation needed]. As a result, for typical head movements, which cover the frequency range of 0.1 Hz and 10 Hz, the deflection of the cupula is approximately proportional to the head-velocity. This is very useful, since the velocity of the eyes must be opposite to the velocity of the head in order to have clear vision.
Central processing[edit]
Signals from the vestibular system also project to the cerebellum (where they are used to keep the VOR effective, a task usually referred to as learning or adaptation) and to different areas in the cortex. The projections to the cortex are spread out over different areas, and their implications are currently not clearly understood.
Projection pathways[edit]
The vestibular nuclei on either sides of the brain stem exchange signals regarding movement and body position. These signals are sent down the following projection pathways.
To the cerebellum. Signals sent to the cerebellum are relayed back as muscle movements of the head, eyes, and posture.
To nuclei of cranial nerves III, IV, and VI. Signals sent to these nerves cause the vestibulo-ocular reflex. They allow for the eyes to fix on a moving object while staying in focus.
To the reticular formation. Signals sent to the reticular formation signal the new posture the body has taken on, and how to adjust circulation and breathing due to body position.
To the spinal cord. Signals sent to the spinal cord allow quick reflex reactions to both the limbs and trunk to regain balance.
To the thalamus. Signals sent to the thalamus allow for head and body motor control as well as being conscious of body position.[2]
Via the Ventral Pathway, which contributes to vertical orientation and perception of the direction of gravity.[3]
Otolithic organs[edit]
While the semicircular canals respond to rotations, the otolithic organs sense linear accelerations. Humans have two otolithic organs on each side, one called the utricle, the other called the saccule. The utricle contains a patch of hair cells and supporting cells called a macula. Similarly, the saccule contains a patch of hair cells and a macula. Each hair cell of a macula has 40-70 stereocilia and one true cilium called a kinocilium. The tips of these cilia are embedded in an otolithic membrane. This membrane is weighted down with protein-calcium carbonate granules called otoliths. These otoliths add to the weight and inertia of the membrane and enhance the sense of gravity and motion. With the head erect, the otolithic membrane bears directly down on the hair cells and stimulation is minimal. When the head is tilted, however, the otolithic membrane sags and bends the stereocilia, stimulating the hair cells. Any orientation of the head causes a combination of stimulation to the utricles and saccules of the two ears. The brain interprets head orientation by comparing these inputs to each other and to other input from the eyes and stretch receptors in the neck, thereby detecting whether the head is tilted or the entire body is tipping.[2] Essentially, these otolithic organs sense how quickly you are accelerating forward or backward, left or right, or up or down.[4] Most of the utricular signals elicit eye movements, while the majority of the saccular signals projects to muscles that control our posture.
While the interpretation of the rotation signals from the semicircular canals is straightforward, the interpretation of otolith signals is more difficult: since gravity is equivalent to a constant linear acceleration, one somehow has to distinguish otolith signals that are caused by linear movements from those caused by gravity. Humans can do that quite well, but the neural mechanisms underlying this separation are not yet fully understood. Humans can sense head tilting and linear acceleration even in dark environments because of the orientation of two groups of hair cell bundles on either side of the striola. Hair cells on opposite sides move with mirror symmetry, so when one side is moved, the other is inhibited. The opposing effects caused by a tilt of the head cause differential sensory inputs from the hair cell bundles allow humans to tell which way the head is tilting,[5] Sensory information is then sent to the brain, which can respond with appropriate corrective actions to the nervous and muscular systems to ensure that balance and awareness are maintained.[6]
Experience from the vestibular system[edit]
Experience from the vestibular system is called equilibrioception. It is mainly used for the sense of balance and for spatial orientation. When the vestibular system is stimulated without any other inputs, one experiences a sense of self-motion. For example, a person in complete darkness and sitting in a chair will feel that he or she has turned to the left if the chair is turned to the left. A person in an elevator, with essentially constant visual input, will feel she is descending as the elevator starts to descend. There are a variety of direct and indirect vestibular stimuli which can make people feel they are moving when they are not, not moving when they are, tilted when they are not, or not tilted when they are.[7] Although the vestibular system is a very fast sense used to generate reflexes, including the righting reflex, to maintain perceptual and postural stability, compared to the other senses of vision, touch and audition, vestibular input is perceived with delay.[8][9]
Vestibular/somatogyral illusions[edit]
Further information: Sensory illusions in aviation
Pathologies[edit]
Diseases of the vestibular system can take different forms, and usually induce vertigo[citation needed][10] and instability or loss of balance, often accompanied by nausea. The most common vestibular diseases in humans are vestibular neuritis, a related condition called labyrinthitis, Ménière's disease, and BPPV. In addition, the function of the vestibular system can be affected by tumors on the vestibulocochlear nerve, an infarct in the brain stem or in cortical regions related to the processing of vestibular signals, and cerebellar atrophy.
Alcohol can also cause alterations in the vestibular system for short periods and will result in vertigo and possibly nystagmus due to the variable viscosity of the blood and the endolymph during the consumption of alcohol. The common term for this type of sensation is the bed spins.[citation needed]
PAN I - The alcohol concentration is higher in the blood than in the vestibular system, hence the endolymph is relatively dense.
PAN II - The alcohol concentration is lower in the blood than in the vestibular system, hence the endolymph is relatively dilute.
PAN I will result in subjective vertigo in one direction and typically occurs shortly after ingestion of alcohol when blood alcohol levels are highest. PAN II will eventually cause subjective vertigo in the opposite direction. This occurs several hours after ingestion and after a relative reduction in blood alcohol levels.[citation needed]
Benign paroxysmal positional vertigo, or BPPV for short, is a condition resulting in acute symptoms of vertigo in people. It is probably caused when pieces that have broken off otoliths have slipped into one of the semicircular canals. In most cases it is the posterior canal that is affected. In certain head positions, these particles shift and create a fluid wave which displaces the cupula of the canal affected, which leads to dizziness, vertigo and nystagmus.
A similar condition to BPPV may occur in dogs and other mammals, but the term vertigo cannot be applied because it refers to subjective perception. Terminology is not standardized for this condition.
A common vestibular pathology of dogs and cats is colloquially known as "old dog vestibular disease", or more formally idiopathic peripheral vestibular disease, which causes sudden episode of loss of balance, circling, head tilt, and other signs. This condition is very rare in young dogs but fairly common in geriatric animals, and may affect cats of any age.[11]
Vestibular dysfunction has also been found to correlate with cognitive and emotional disorders, including depersonalization and derealization.[12]
See also[edit]
Dark cell
Migraine-associated vertigo
Statocyst
References[edit]
Jump up ^ Boulpaep, Emile L.; Boron, Walter F. (2005). Medical physiology: a cellular and molecular approach. St. Louis, Mo: Elsevier Saunders. ISBN 1-4160-2328-3. OCLC 56963726. CS1 maint: Multiple names: authors list (link)
^ Jump up to: a b Saladin, Kenneth S. (2011). Anatomy & Physiology: The Unity of Form and Function. New York: McGraw-Hill. ISBN 0-07-337825-9. OCLC 799004854.
Jump up ^ Vaziri, Siavash; Connor, Charles E (2016). "Representation of Gravity-Aligned Scene Structure in Ventral Pathway Visual Cortex.". Current Biology. 26: 766–74. PMID 26923785. doi:10.1016/j.cub.2016.01.022. Lay summary. ... the ventral pathway, ...could support perceiving and/or predicting physical events involving objects subject to gravity ...
Jump up ^ "The Physiology of the Senses: Balance" (PDF).
Jump up ^ "The Otolith Organs: The Utricle and Sacculus". NCBI Bookshelf - Neuroscience.
Jump up ^ Angelaki DE, Cullen KE (2008). "Vestibular system: the many facets of a multimodal sense". Annu. Rev. Neurosci. 31: 125–50. PMID 18338968. doi:10.1146/annurev.neuro.31.060407.125555.
Jump up ^ Lawson, B. D., & Riecke, B. E. (2014). The Perception of Body Motion. Handbook of Virtual Environments, CRC Press, 163-196.
Jump up ^ Barnett-Cowan, M., and Harris, L. R. (2009), Perceived timing of vestibular stimulation relative to touch, light and sound. Experimental Brain Research, 198: 221-231. "doi: 10.1007/s00221-009-1779-4".
Jump up ^ Barnett-Cowan, M. (2013), Vestibular perception is slow: a review. Multisensory Research, 26: 387-403. "doi: 10.1163/22134808-00002421".
Jump up ^ "Vertigo". University of Maryland Medical Center. Retrieved 13 November 2015.
Jump up ^ Rossmeisl, John (2010). "Vestibular Disease in Dogs and Cats". Veterinary Clinics of North America: Small Animal Practice. 40 (1): 80–100. doi:10.1016/j.cvsm.2009.09.007. Retrieved 12 March 2017.
Jump up ^ Smith, Paul F; Darlington, Cynthia L (2013). "Personality changes in patients with vestibular dysfunction.". Frontiers in human neuroscience. 7: 678. PMC 3810789  . PMID 24194706. doi:10.3389/fnhum.2013.00678. Lay summary. patients with vestibular disorders have been reported to experience other personality changes that suggest that vestibular sensation is implicated in the sense of self. These are depersonalization and derealization symptoms such as feeling "spaced out", "body feeling strange" and "not feeling in control of self". We propose in this review that these symptoms suggest that the vestibular system may make a unique contribution to the concept of self through information regarding self-motion and self-location that it transmits, albeit indirectly, to areas of the brain such as the temporo-parietal junction
Further reading[edit]
S. M. Highstein, R. R. Fay, A. N. Popper, editors (2004). The vestibular system. Berlin: Springer. ISBN 0-387-98314-7. OCLC 56068617. CS1 maint: Multiple names: authors list (link)CS1 maint: Extra text: authors list (link) (Comment: A book for experts, summarizing the state of the art in our understanding of the balance system)
Thomas Brandt (2003). Vertigo : Its Multisensory Syndromes. Berlin: Springer. ISBN 0-387-40500-3. OCLC 52472049. (Comment: For clinicians, and other professionals working with dizzy patients.)
"Driver Fatigue: Is Something Missing?" (pdf). Christopher Brill, Peter A. Hancock, Richard D. Gilson - University of Central Florida - 2003. (Comment: Research on driver or motion-induced sleepiness aka 'sopite syndrome' links it to the vestibular labyrinths.)
Vestibular system Kathleen Cullen and Soroush Sadeghi, Scholarpedia, 3(1):3013. doi:10.4249/scholarpedia.3013
[1] A preview of an article on how vestibular disorders can cause symptoms that look like mental disorders.
External links[edit]
Vestibular Disorders Association For more information about vestibular (inner ear balance) disorders.
(Video) Head Impulse Testing site (vHIT) Site with thorough information about vHIT
SensesWeb, which contains animations of all sensory systems, and additional links.
Dizzytimes.com Online Community for Sufferers of Vertigo and Dizziness.
Vestibular System, Neuroscience Online (electronic neuroscience textbook)
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