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Music history of the United States in the 1960s
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Popular music of the United States in the 1960s became innately tied up into causes, opposing certain ideas, influenced by the sexual revolution, feminism, Black Power and environmentalism. This trend took place in a tumultuous period of massive public unrest in the United States which consisted of the Cold War, Vietnam War, and Civil Rights Movement.
Central to this trend was a folk roots revival that inspired a wave of similar trends across Europe and the rest of the world. This stemmed from a revival of hillbilly music early in the decade, and drew on Appalachian folk-pop pioneers The Weavers. Singer-songwriters like Bob Dylan and Joan Baez broke new ground in lyrical approach and personal style in composition, setting the stage for the next wave of lighter, country music and rhythm and blues-influenced singer-songwriters like James Taylor, Elton John, Carole King, and Cat Stevens, who began topping the charts in the very early 1970s.
The 1960s began with soul music topping the charts, including pure soul divas and singers specializing in the new, rhythm and blues-gospel music fusion with a secular approach. Later specialties in soul cropped up, including girl groups, blue-eyed soul, brown-eyed soul, Memphis soul, Philly soul and, most popular, Motown. The last part of the decade saw soul singer-songwriters like Marvin Gaye invent album-oriented soul, and James Brown and his ever-evolving backing band invent funk.
Modern music in the 1960s was dominated by the Nashville sound until Merle Haggard changed the national country sound to the Bakersfield sound. For a time, the Bakersfield sound was the only homegrown music that could compete in sales against an influx of British bands; this was called the British Invasion, and it sparked a new wave of music and social activism. Psychedelic rock arose from this subculture, which opposed the Vietnam War and supported civil rights and other generally leftist causes. While the energy in this scene remained strong for some time, it soon splintered into competing heavy metal, early art-punk rock and progressive rock.
Contents
[hide]
1 Folk music
2 Rock
2.1 Folk rock
2.2 Psychedelic rock
2.3 Surf rock
2.4 Garage rock
2.5 Blues rock
2.6 Roots rock
2.7 Progressive rock
3 Pop
4 R&B and Soul
5 Country music
6 Other trends
7 References
8 See also
Folk music[edit]
Main article: American folk music revival
These performers and several others were instrumental in launching the folk music revival of the 1950s and 1960s.
The Kingston Trio[1]
The Weavers[1]
Pete Seeger[1]
Woody Guthrie[1]
Odetta[2]
Peter, Paul and Mary[2]
Joan Baez[2]
Bob Dylan[3]
Judy Collins
Leonard Cohen
Joni Mitchell
Carolyn Hester
Phil Ochs
Tom Paxton
Buffy Sainte-Marie[2]
Dave Van Ronk
Tom Rush
Fred Neil
Gordon Lightfoot
Ian and Sylvia
Arlo Guthrie
The Brothers Four[2]
The Limeliters[2]
Rock[edit]
Folk rock[edit]
Bob Dylan and Joan Baez during the civil rights "March on Washington", 28 August 1963
By the 1960s, the scene that had developed out of the American folk music revival had grown to a major movement, utilizing traditional music and new compositions in a traditional style, usually on acoustic instruments.[1][4] In America the genre was pioneered by figures such as Woody Guthrie and Pete Seeger and often identified with progressive or labor politics.[4] In the early sixties figures such as Joan Baez and Bob Dylan had come to the fore in this movement as singer-songwriters.[5] Dylan had begun to reach a mainstream audience with hits including "Blowin' in the Wind" (1963) and "Masters of War" (1963), which brought "protest songs" to a wider public,[6] but, although beginning to influence each other, rock and folk music had remained largely separate genres, often with mutually exclusive audiences.[7]
Early attempts to combine elements of folk and rock included the Animals "House of the Rising Sun" (1964), which was the first commercially successful folk song to be recorded with rock and roll instrumentation.[8] The folk rock movement is usually thought to have taken off with The Byrds' recording of Dylan's "Mr. Tambourine Man" which topped the charts in 1965.[7] With members who had been part of the cafe-based folk scene in Los Angeles, the Byrds adopted rock instrumentation, including drums and 12-string Rickenbacker guitars, which became a major element in the sound of the genre.[7] Later that year Dylan adopted electric instruments, much to the outrage of many folk purists, with his "Like a Rolling Stone" becoming a US hit single.[7] Folk rock particularly took off in California, where it led acts like The Mamas & the Papas and Crosby, Stills and Nash to move to electric instrumentation, and in New York, where it spawned performers including The Lovin' Spoonful and Simon and Garfunkel, with the latter's acoustic "The Sounds of Silence" being remixed with rock instruments to be the first of many hits.[7]
Folk rock reached its peak of commercial popularity in the period 1967-8, before many acts moved off in a variety of directions, including Dylan and the Byrds, who began to develop country rock.[9] However, the hybridization of folk and rock has been seen as having a major influence on the development of rock music, bringing in elements of psychedelia, and helping to develop the ideas of the singer-songwriter, the protest song and concepts of "authenticity".[7][10]
Psychedelic rock[edit]
Jimi Hendrix, 1967
Psychedelic music's LSD-inspired vibe began in the folk scene, with the New York-based Holy Modal Rounders using the term in their 1964 recording of "Hesitation Blues".[11] The first group to advertise themselves as psychedelic rock were the 13th Floor Elevators from Texas, at the end of 1965; producing an album that made their direction clear, with The Psychedelic Sounds of the 13th Floor Elevators the following year.[11]
Psychedelic rock particularly took off in California's emerging music scene as groups followed the Byrds from folk to folk rock from 1965.[12] The psychedelic life style had already developed in San Francisco and particularly prominent products of the scene were The Grateful Dead, Country Joe and the Fish, The Great Society and Jefferson Airplane.[12][13] The Byrds rapidly progressed from purely folk rock in 1966 with their single "Eight Miles High",[14] perceived by many listeners to be a reference to drug use. In actuality, it referred to The Byrds' 1965 plane trip to England.[15]
Psychedelic rock reached its apogee in the last years of the decade. In America the Summer of Love was prefaced by the Human Be-In event and reached its peak at the Monterey Pop Festival,[16] the later helping to make major American stars of Jimi Hendrix and The Who, whose single "I Can See for Miles" delved into psychedelic territory.[17] Key recordings included Jefferson Airplane's Surrealistic Pillow and The Doors' Strange Days.[18] These trends climaxed in the 1969 Woodstock festival,[19] which saw performances by most of the major psychedelic acts, but by the end of the decade psychedelic rock was in retreat. The Jimi Hendrix Experience broke up before the end of the decade and many surviving acts, moved away from psychedelia into more back-to-basics "roots rock", the wider experimentation of progressive rock, or riff laden heavy rock.[12]
Surf rock[edit]
The Beach Boys, 2008
In the early 1960s, one of the most popular forms of rock and roll was Surf Rock, which was characterized by being nearly entirely instrumental and by heavy use of reverb on the guitars. The spring reverb featured in Fender amplifiers of the day, cranked to its maximum volume, produced a guitar tone shimmering with sustain and evoking surf and ocean imagery.
Duane Eddy's "Movin' and Groovin" is thought by many to be the main contender for laying the groundwork as the first surf rock record, while others claim the genre was invented by Dick Dale on "Let's Go Trippin'", which became a hit throughout California. Most early surf bands were formed in during this decade in the Southern California area. By the mid-1960s The Beach Boys, who used complex pop harmonies over a basic surf rock rhythm, had emerged as the dominant surf group and helped popularize the genre.[20] In addition, bands such as The Ventures, The Shadows, The Atlantics, The Surfaris and The Champs were also among the most popular Surf Rock bands of the decade.
Garage rock[edit]
Garage rock was a form of amateurish rock music, particularly prevalent in North America in the mid-1960s and so called because of the perception that it was rehearsed in a suburban family garage.[21][22] Garage rock songs revolved around the traumas of high school life, with songs about "lying girls" being particularly common.[23] The lyrics and delivery were notably more aggressive than was common at the time, often with growled or shouted vocals that dissolved into incoherent screaming.[21] They ranged from crude one-chord music (like the Seeds) to near-studio musician quality (including the Knickerbockers, the Remains, and the Fifth Estate). There were also regional variations in many parts of the country with flourishing scenes particularly in California and Texas.[23] The Pacific Northwest states of Washington and Oregon had perhaps the most defined regional sound.[24]
The style had been evolving from regional scenes as early as 1958. "Louie Louie" by The Kingsmen (1963) is a mainstream example of the genre in its formative stages. By 1963, garage band singles were creeping into the national charts in greater numbers, including Paul Revere and the Raiders (Boise),[25] the Trashmen (Minneapolis)[26] and the Rivieras (South Bend, Indiana).[27] In this early period many bands were heavily influenced by surf rock and there was a cross-pollination between garage rock and frat rock, sometimes viewed as merely a subgenre of garage rock.[28]
The British Invasion of 1964-66 greatly influenced garage bands, providing them with a national audience, leading many (often surf or hot rod groups) to adopt a British Invasion lilt, and encouraging many more groups to form.[23] Thousands of garage bands were extant in the USA and Canada during the era and hundreds produced regional hits.[23] Despite scores of bands being signed to major or large regional labels, most were commercial failures. It is generally agreed that garage rock peaked both commercially and artistically around 1966.[23] By 1968 the style largely disappeared from the national charts and at the local level as amateur musicians faced college, work or the draft.[23] New styles had evolved to replace garage rock (including blues rock, progressive rock and country rock).[23] In Detroit garage rock stayed alive until the early 70s, with bands like the MC5 and The Stooges, who employed a much more aggressive style. These bands began to be labelled punk rock and are now often seen as proto-punk or proto-hard rock.[29]
Blues rock[edit]
The American blues rock had been pioneered in the early 1960s by guitarist Lonnie Mack,[30] but the genre began to take off in the mid-60s as acts followed developed a sound similar to British blues musicians. Key acts included Paul Butterfield (whose band acted like Mayall's Bluesbreakers in Britain as a starting point for many successful musicians), Canned Heat, the early Jefferson Airplane, Janis Joplin, Johnny Winter, The J. Geils Band and Jimi Hendrix with his power trios, The Jimi Hendrix Experience and Band of Gypsys, whose guitar virtuosity and showmanship would be among the most emulated of the decade.[31] Blues rock bands like Allman Brothers Band, Lynyrd Skynyrd and eventually ZZ Top from the southern states, incorporated country elements into their style to produce distinctive Southern rock.[32]
Roots rock[edit]
Creedence Clearwater Revival, 2009
Roots rock is the term now used to describe a move away from the excesses of the psychedelic scene, to a more basic form of rock and roll that incorporated its original influences, particularly country and folk music, leading to the creation of country rock and Southern rock.[33] In 1966 Bob Dylan spearheaded the movement when he went to Nashville to record the album Blonde on Blonde.[34] This, and subsequent more clearly country-influenced albums, have been seen as creating the genre of country folk, a route pursued by a number of, largely acoustic, folk musicians.[34] Other acts that followed the back-to-basics trend were the Canadian group The Band and the Californian-based Creedence Clearwater Revival, both of which mixed basic rock and roll with folk, country and blues, to be among the most successful and influential bands of the late 1960s.[35] The same movement saw the beginning of the recording careers of Californian solo artists like Ry Cooder, Bonnie Raitt and Lowell George,[36] and influenced the work of established performers such as the Rolling Stones' Beggar's Banquet (1968) and the Beatles' Let It Be (1970).[12]
In 1968 Gram Parsons recorded Safe at Home with the International Submarine Band, arguably the first true country rock album.[37] Later that year he joined the Byrds for Sweetheart of the Rodeo (1968), generally considered one of the most influential recordings in the genre.[37] The Byrds continued in the same vein, but Parsons left to be joined by another ex-Byrds member Chris Hillman in forming The Flying Burrito Brothers who helped establish the respectability and parameters of the genre, before Parsons departed to pursue a solo career.[37] Country rock was particularly popular in the Californian music scene, where it was adopted by bands including Hearts and Flowers, Poco and Riders of the Purple Sage,[37] the Beau Brummels[37] and the Nitty Gritty Dirt Band.[38] A number of performers also enjoyed a renaissance by adopting country sounds, including: the Everly Brothers; one-time teen idol Ricky Nelson who became the frontman for the Stone Canyon Band; former Monkee Mike Nesmith who formed the First National Band; and Neil Young.[37] The Dillards were, unusually, a country act, who moved towards rock music.[37] The greatest commercial success for country rock came in the 1970s, with artist including the Doobie Brothers, Emmylou Harris, Linda Ronstadt and the Eagles (made up of members of the Burritos, Poco and Stone Canyon Band), who emerged as one of the most successful rock acts of all time, producing albums that included Hotel California (1976).[39]
The founders of Southern rock are usually thought to be the Allman Brothers Band, who developed a distinctive sound, largely derived from blues rock, but incorporating elements of boogie, soul, and country in the early 1970s.[40] The most successful act to follow them were Lynyrd Skynyrd, who helped establish the "Good ol' boy" image of the subgenre and the general shape of 1970s guitar rock.[40] Their successors included the fusion/progressive instrumentalists Dixie Dregs, the more country-influenced Outlaws, jazz-leaning Wet Willie and (incorporating elements of R&B and gospel) the Ozark Mountain Daredevils.[40] After the loss of original members of the Allmans and Lynyrd Skynyrd, the genre began to fade in popularity in the late 1970s, but was sustained the 1980s with acts like .38 Special, Molly Hatchet and The Marshall Tucker Band.[40]
Progressive rock[edit]
Progressive rock, sometimes used interchangeably with art rock, was an attempt to move beyond established musical formulas by experimenting with different instruments, song types, and forms.[41] From the mid-1960s The Left Banke, The Beatles, The Rolling Stones and The Beach Boys, had pioneered the inclusion of harpsichords, wind and string sections on their recordings to produce a form of Baroque rock and can be heard in singles like Procol Harum's "A Whiter Shade of Pale" (1967), with its Bach inspired introduction.[42] The Moody Blues used a full orchestra on their album Days of Future Passed (1967) and subsequently created orchestral sounds with synthesisers.[41] Classical orchestration, keyboards and synthesisers were a frequent edition to the established rock format of guitars, bass and drums in subsequent progressive rock.[43]
Instrumentals were common, while songs with lyrics were sometimes conceptual, abstract, or based in fantasy and science fiction.[44] The Pretty Things' SF Sorrow (1968) and The Who's Tommy (1969) introduced the format of rock operas and opened the door to "concept albums, usually telling an epic story or tackling a grand overarching theme."[45] King Crimson's 1969 début album, In the Court of the Crimson King, which mixed powerful guitar riffs and mellotron, with jazz and symphonic music, is often taken as the key recording in progressive rock, helping the widespread adoption of the genre in the early 1970s among existing blues rock and psychedelic bands, as well as newly formed acts.[41]
Pop[edit]
Chubby Checker performing in 2005
Chubby Checker during the early 1960s popularizes the enduring dance craze The Twist with his hit cover of Hank Ballard's R&B hit "The Twist".[20]
Gerry Goffin and Carole King become a very influential duo in pop music, writing numerous number one hits including the first song to ever reach number one by a girl group, The Shirelles "Will You Love Me Tomorrow" and the 1962 number one hit, "The Loco-Motion" which was performed by Little Eva.
Sugar Sugar becomes a big hit for The Archies, defining the bubblegum pop genre.
R&B and Soul[edit]
Motown develops as a pop-influenced answer to soul music. The label begins a long run of No. 1 U.S. hit singles in 1961 with "Please Mr. Postman" by The Marvelettes. The label would have numerous No. 1 Billboard hits throughout the decade[46] and into the 1990s.
Soul music develops popularity throughout the decade, led by Ray Charles,[47] Sam Cooke,[48] James Brown, and Otis Redding, among many others.[49]
Funk begins later in the decade with James Brown and Sly & the Family Stone having early hits.
You Keep Me Hanging On uses a fast tempo which would prove innovative in the development of disco music.
Aretha Franklin's 1967 recordings,[50] such as "I Never Loved a Man (The Way I Love You)", "Respect" (originally sung by Otis Redding), and "Do Right Woman-Do Right Man", are considered the apogee of the soul genre, and were among its most commercially successful productions.
Country music[edit]
Triumph and great tragedy marked the 1960s in country music. The genre continued to gain national exposure through network television, with weekly series and awards programs gaining popularity. Sales of records continued to rise as new artists and trends came to the forefront. However, several top stars died under tragic circumstances, including several who were killed in plane crashes.
The predominant musical style during the decade was the Nashville Sound, a style that emphasized string sections, background vocals, crooning lead vocals and production styles seen in country music. The style had first become popular in the late 1950s, in response to the growing encroachment of rock and roll on the country genre, but saw its greatest success in the 1960s. Artists like Jim Reeves, Eddy Arnold, Ray Price, Patsy Cline, Floyd Cramer, Roger Miller,[51] and many others achieved great success through songs such as "He'll Have to Go,"[52] "Danny Boy," "Make the World Go Away", "King of the Road",[51] and "I Fall to Pieces." The country-pop style was also evident on the 1962 album Modern Sounds in Country and Western Music, recorded by rhythm and blues and soul singer Ray Charles. Charles recorded covers of traditional country, folk and classical music standards in pop, R&B and jazz styles. The album was hailed as a critical and commercial success, and would be vastly influential in later country music styles.[47] Songs from the album that were released for commercial airplay and record sales included "I Can't Stop Loving You," "Born to Lose" and "You Don't Know Me."
By the end of the decade, the Nashville Sound became more polished and streamlined, and became known as "countrypolitan." Tammy Wynette, Glen Campbell, Dottie West and Charley Pride were among the top artists adopting this style. While George Jones — by the early 1960s one of country music's most consistent hitmakers — also recorded countrypolitan-styled music, his background remained pure honky tonk, singing of heartbreak and lonlieness in many of his songs. Also, Marty Robbins proved to be one of the genre's most diverse singers,[52] singing everything from straight-ahead country to western to pop to blues ... and even Hawaiian.
Johnny Cash—who became known as "The Man in Black"—became of the most influential musicians of the 1960s (and eventually, 20th century).[52] Although primarily recording country, his songs and sound spanned many other genres including rockabilly, blues, folk, and gospel. His music showed great compassion for minorities and others who were shunned by society, including prison inmates. Two of Cash's most successful albums were recorded live in prison: At Folsom Prison and At San Quentin.
During the latter half of the 1960s, Pride — a native of Sledge, Mississippi — became the first African-American superstar in country music, a genre virtually dominated by white artists. Some of his early hits, sang with a smooth baritone voice and in a style meshing honky-tonk and countrypolitan, included "Just Between You and Me," "The Easy Part's Over," "All I Have to Offer You (Is Me)" and a cover version of Hank Williams' "Kaw-Liga." Pride continued to be successful for more than 20 years, amassing an eventual 29 No. 1 hits on the Billboard Hot Country Singles chart.
A newly emerging style, which had its roots in the 1950s but exploded in the mainstream during the 1960s, was the "Bakersfield sound." Instead of creating a sound similar to mainstream pop music, the Bakersfield sound used honky tonk as its base and added electric instruments and a backbeat, plus stylistic elements borrowed from rock and roll. Buck Owens, Merle Haggard and Wynn Stewart were some of the top artists adopting this sound, and by the late 1960s they were among country music's top selling artists.
Dolly Parton, a native of the Smoky Mountains town of Locust Ridge, Tennessee, gained national exposure on the nationally syndicated program The Porter Wagoner Show. Her mountain-influenced, biographical brand of country and her down-home personality won many fans, and her star power would only begin to rise.
In addition to the syndicated The Porter Wagoner Show, several other television programs were produced to allow country music to reach a wider audience, such as The Jimmy Dean Show in med-decade. At the end of the decade, Hee Haw began a 23-year run, first on CBS and later in syndication; Hee Haw, hosted by Owens and Roy Clark was loosely based on the comedy series Rowan & Martin's Laugh In, and incorporated comedy along with performances by the show's cast or guest performers from the country music field. The Academy of Country Music and Country Music Association awards programs were telecast for the first time in the late 1960s.
The 1960s were marred with tragedy. Johnny Horton, who sang in the saga-song style, was killed in a car accident in 1960. A March 5, 1963, plane crash claimed the lives of Patsy Cline, Cowboy Copas and Hawkshaw Hawkins. Days later, Jack Anglin was killed in a car accident, while Texas Ruby died in a trailer fire in Texas. In July 1964, Jim Reeves lost his life while piloting a plane near Brentwood, Tennessee. Ira Louvin was killed in a car accident in 1965. Success overcame several of those tragic deaths, as both Cline and Reeves had many posthumous hits (with previously recorded songs issued after their deaths) and enjoyed strong followings for many years.
The 1960s began a trend toward a proliferation of No. 1 hits on the Billboard Hot Country Singles chart, thanks to ever-changing data collecting methods. When the 1960s decade opened, there were but four No. 1 songs topping the chart (five, if one counts Marty Robbins' "El Paso"), but by the mid-1960s, there were always at least a dozen songs topping the chart annually. In 1967, there were more than 20 songs reaching the top spot for the first time ever in a single calendar year ... and that number would only continue to rise during the next 20 years.
Other trends[edit]
Woodstock Festival, 1969
Large festivals taking place over several days, 1967's Monterey Pop Festival[16] and 1969's Woodstock Music Festival[19] would feature many popular rock musicians.
Current events become a major influence on popular music. Many songs are written in protest to the Vietnam War. The song "Ohio" was written about the Kent State Massacre, and became a hit for Crosby, Stills, Nash, and Young.
World music sees a huge rise in popularity as many seek interest in other cultures. Ravi Shankar performs at the Monterey and Woodstock festivals. Latin Rock artist Carlos Santana sees popularity throughout the decade. George Harrison develops an interest in the Hare Krishna culture, adding Indian influence to The Beatles' music including the use of a sitar. Reggae begins to popularize at this time.
In 1969, the Rolling Stones organized the ill-fated Altamont Free Concert.
Songs like "Summertime Blues" and "Eve of Destruction" address the issue of the voting age, which at the time was 21. The issue was that soldiers were drafted at 18, but could not vote. The voting age was eventually lowered to eighteen.
A few songs such as Bob Dylan's Blowin' in the Wind address the Civil Rights Movement.
References[edit]
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^ Jump up to: a b c Bogdanov, Woodstra & Erlewine 2002, pp. 1330–1
Jump up ^ J. S. Harrington, Sonic Cool: the Life & Death of Rock 'n' Roll (Milwaukee, WI: Hal Leonard Corporation, 2003), ISBN 0-634-02861-8, p. 191.
Jump up ^ E. Macan, Rocking the Classics: English Progressive Rock and the Counterculture (Oxford: Oxford University Press, 1997), ISBN 0-19-509887-0, pp. 34-5.
Jump up ^ E. Macan, Rocking the Classics: English Progressive Rock and the Counterculture (Oxford: Oxford University Press, 1997), ISBN 0-19-509887-0, p. 64.
Jump up ^ "Progressive rock", Allmusic, retrieved 29 October 2009.
Jump up ^ Gilliland 1969, show 25.
^ Jump up to: a b Gilliland 1969, show 16.
Jump up ^ Gilliland 1969, show 17.
Jump up ^ Gilliland 1969, show 51.
Jump up ^ Gilliland 1969, show 52.
^ Jump up to: a b Gilliland 1969, show 11.
^ Jump up to: a b c Gilliland 1969, show 10.
See also[edit]
1960s in music
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Otolith
From Wikipedia, the free encyclopedia
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Otolith
Otolith organs showing detail of utricle, otoconia, endolymph, cupula, macula, hair cell filaments, and saccular nerve
Juvenile herring. Length 30 mm; 3 months old; still transparent; the otoliths are visible left of the eye.
Details
Identifiers
Latin
statoconium
MeSH
D010037
TA
A15.3.03.086
FMA
77826
Anatomical terminology
[edit on Wikidata]
An otolith (Greek: ὠτο-, ōto- ear + λῐ́θος, líthos, a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The saccule and utricle, in turn, together make the otolith organs. These organs are what allows an organism, including humans, to perceive linear acceleration, both horizontally and vertically (gravity). They have been identified in both extinct and extant vertebrates.[1]
Counting the annual growth rings on the otoliths is a common technique in estimating the age of fish.

Contents
1 Description
2 Mechanism
3 Paleontology
4 Ecology
4.1 Composition
4.2 Growth rate and age
4.3 Diet research
5 See also
6 References
7 External links
Description[edit]
Endolymphatic infillings such as otoliths are structures in the saccule and utricle of the inner ear, specifically in the vestibular labyrinth of all vertebrates (fish, amphibians, reptiles, mammals and birds). In vertebrates, the saccule and utricle together make the otolith organs. Both statoconia and otoliths are used as gravity, balance, movement, and directional indicators in all vertebrates and have a secondary function in sound detection in higher aquatic and terrestrial vertebrates.[2][3] They are sensitive to gravity and linear acceleration. Because of their orientation in the head, the utricle is sensitive to a change in horizontal movement, and the saccule gives information about vertical acceleration (such as when in an elevator).
Similar balance receptors called statocysts can be found in many invertebrate groups but are not contained in the structure of an inner ear. Mollusk statocysts are of a similar morphology to the displacement-sensitive organs of vertebrates;[4] however, the function of the mollusk statocyst is restricted to gravity detection and possibly some detection of angular momentum.[5] These are analogous structures, with similar form and function but not descended from a common structure.
Statoconia (also called otoconia) are numerous grains, often spherical in shape, between 1 and 50 µm; collectively[citation needed]. Statoconia are also sometimes termed a statocyst. Otoliths (also called statoliths) are agglutinated crystals or crystals precipitated around a nucleus, with well defined morphology and together all may be termed endolymphatic infillings.[1][6][7]
Mechanism[edit]
The semicircular canals and sacs in all vertebrates are attached to endolymphatic ducts, which in some groups (such as sharks) end in small openings, called endolymphatic pores, on the dorsal surface of the head.[1] Extrinsic grains may enter through these openings, typically less than a millimeter in diameter. The size of material that enters is limited to sand-sized particles and in the case of sharks is bound together with endogenous an organic matrix that the animal secrets.
In mammals, otoliths are small particles, composed of a combination of a gelatinous matrix and calcium carbonate in the viscous fluid of the saccule and utricle. The inertia of these small particles causes them to stimulate hair cells when the head moves. The hair cells are made up of 40 to 70 stereocilia and one kinocilium, which is connected to an afferent nerve. When the body changes position or begins a movement the weight of the membrane bends the stereocilia and stimulates the hair cells. Hair cells send signals down sensory nerve fibers, which are interpreted by the brain as motion. The brain interprets the orientation of the head by comparing the input from the utricles and saccules from both ears to the input from the eyes, allowing the brain to discriminate a tilted head from the movement of the entire body. When the head is in a normal upright position, the otolith presses on the sensory hair cell receptors. This pushes the hair cell processes down and prevents them from moving side to side. However, when the head is tilted, the pull of gravity on otoliths shift the hair cell processes to the side, distorting them and sending a message to the central nervous system that the head is no longer level but now tilted. (see: BPPV) This theory may have to be reevaluated because of an experiment in which a blindfolded owl in zero gravity was able to keep its head level while a handler was rocking its body back and forth.[8]
There is evidence that the vestibular system of mammals has retained some of its ancestral acoustic sensitivity and that this sensitivity is mediated by the otolithic organs (most likely the sacculus, due to its anatomical location). In mice lacking the otoconia of the utricle and saccule, this retained acoustic sensitivity is lost.[3] In humans vestibular evoked myogenic potentials occur in response to loud, low-frequency acoustic stimulation in patients with the sensorineural hearing loss.[2] Vestibular sensitivity to ultrasonic sounds has also been hypothesized to be involved in the perception of speech presented at artificially high frequencies, above the range of the human cochlea (~18 kHz).[9] In mice sensation of acoustic information via the vestibular system has been demonstrated to have a behaviourally relevant effect; response to an elicited acoustic startle reflex is larger in the presence of loud, low frequency sounds that are below the threshold for the mouse cochlea (~4 Hz), raising the possibility that the acoustic sensitivity of the vestibular system may extend the hearing range of small mammals.[3]
Paleontology[edit]
After the death and decomposition of a fish, otoliths may be preserved within the body of an organism or be dispersed before burial and fossilization. Dispersed otoliths are one of the many microfossils which can be found through a micropalaeontological analysis of a fine sediment. Their stratigraphic significance is minimal, but can still be used to characterize a level or interval. Fossil otoliths are rarely found in situ (on the remains of the animal), likely because they are not recognized separately from the surrounding rock matrix. In some cases, due to differences in colour, grain size, or a distinctive shape, they can be identified. These rare cases are of special significance, since the presence, composition, and morphology of the material can clarify the relationship of species and groups. In the case of primitive fish, various fossil material shows that endolymphatic infillings were similar in elemental composition to the rock matrix but were restricted to coarse grained material, which presumably is better for the detection of gravity, displacement, and sound. The presence of these extrinsic grains, in osteostracans, chondrichthyans, and acanthodians indicates a common inner ear physiology and presence of open endolymphatic ducts.[1]
Ecology[edit]
Composition[edit]
Play media
Animation of the biomineralization of cod otoliths
The composition of fish otoliths is also proving useful to fisheries scientists. The calcium carbonate that the otolith is composed of is primarily derived from the water. As the otolith grows, new calcium carbonate crystals form. As with any crystal structure, lattice vacancies will exist during crystal formation allowing trace elements from the water to bind with the otolith. Studying the trace elemental composition or isotopic signatures of trace elements within a fish otolith gives insight to the water bodies fish have previously occupied.[10] Fish otoliths as old as 172 million years have been used to study the environment in which the fish lived.[11] Robotic micromilling devices have also been used to recover very high resolution records of life history, including diet and temperatures throughout the life of the fish, as well as their natal origin.[12]
The most studied trace and isotopic signatures are strontium due to the same charge and similar ionic radius to calcium; however, scientists can study multiple trace elements within an otolith to discriminate more specific signatures. A common tool used to measure trace elements in an otolith is a laser ablation inductively coupled plasma mass spectrometer. This tool can measure a variety of trace elements simultaneously. A secondary ion mass spectrometer can also be used. This instrument can allow for greater chemical resolution but can only measure one trace element at a time. The hope of this research is to provide scientists with valuable information on where fish have traveled. Combined with otolith annuli, scientists can add how old fish were when they traveled through different water bodies. All this information can be used to determine fish life cycles so that fisheries scientists can make better informed decisions about fish stocks.
Growth rate and age[edit]
A pair of sagittae from a Pacific Cod (Gadus macrocephalus).
Removing an otolith from a red snapper to determine its age
See also: estimating the age of fish
Finfish (class Osteichthyes) have three pairs of otoliths – the sagittae (singular sagitta), lapilli (singular lapillus), and asterisci (singular asteriscus). The sagittae are largest, found just behind the eyes and approximately level with them vertically. The lapilli and asterisci (smallest of the three) are located within the semicircular canals. The sagittae are normally composed of aragonite (although vaterite abnormalities can occur[13]), as are the lapilli, while the asterisci are normally composed of vaterite.
The shapes and proportional sizes of the otoliths vary with fish species. In general, fish from highly structured habitats such as reefs or rocky bottoms (e.g. snappers, groupers, many drums and croakers) will have larger otoliths than fish that spend most of their time swimming at high speed in straight lines in the open ocean (e.g. tuna, mackerel, dolphinfish). Flying fish have unusually large otoliths, possibly due to their need for balance when launching themselves out of the water to "fly" in the air. Often, the fish species can be identified from distinct morphological characteristics of an isolated otolith.
Fish otoliths accrete layers of calcium carbonate and gelatinous matrix throughout their lives. The accretion rate varies with growth of the fish – often less growth in winter and more in summer – which results in the appearance of rings that resemble tree rings. By counting the rings, it is possible to determine the age of the fish in years.[14] Typically the sagitta is used, as it is largest,[15] but sometimes lapilli are used if they have a more convenient shape. The asteriscus, which is smallest of the three, is rarely used in age and growth studies.
In addition, in most species the accretion of calcium carbonate and gelatinous matrix alternates on a daily cycle. It is therefore also possible to determine fish age in days.[16] This latter information is often obtained under a microscope, and provides significant data to early life history studies.
By measuring the thickness of individual rings, it has been assumed (at least in some species) to estimate fish growth because fish growth is directly proportional to otolith growth. However, some studies disprove a direct link between body growth and otolith growth. At times of lower or zero body growth the otolith continues to accrete leading some researchers to believe the direct link is to metabolism, not growth per se. Otoliths, unlike scales, do not reabsorb during times of decreased energy making it even more useful tool to age a fish. Fish never stop growing entirely, though growth rate in mature fish is reduced. Rings corresponding to later parts of the life cycle tend to be closer together as a result.
Age and growth studies of fish are important for understanding such things as timing and magnitude of spawning, recruitment and habitat use, larval and juvenile duration, and population age structure. Such knowledge is in turn important for designing appropriate fisheries management policies.
Diet research[edit]
Since the compounds in fish otoliths are resistant to digestion, they are found in the digestive tracts and scats of piscivorous marine mammals, such as dolphins, seals, sea lions and walruses. Many fish can be identified to genus and species by their sagittal otoliths. Otoliths can therefore, to some extent, be used to reconstruct the prey composition of marine mammal diets.
Sagittal otoliths (sagittae) are bilaterally symmetrical, with each fish having one right and one left. Separating recovered otoliths into right and left, therefore, allows one to infer a minimum number of prey individuals ingested for a given fish species. Otolith size is also proportional to the length and weight of a fish. They can therefore be used to back-calculate prey size and biomass, useful when trying to estimate marine mammal prey consumption, and potential impacts on fish stocks.[17]
Otoliths cannot be used alone to reliably estimate cetacean or pinniped diets, however. They may suffer partial or complete erosion in the digestive tract, skewing measurements of prey number and biomass.[18] Species with fragile, easily digested otoliths may be underestimated in the diet. To address these biases, otolith correction factors have been developed through captive feeding experiments, in which seals are fed fish of known size, and the degree of otolith erosion is quantified for different prey taxa.[19]
The inclusion of fish vertebrae, jaw bones, teeth, and other informative skeletal elements improves prey identification and quantification over otolith analysis alone.[20] This is especially true for fish species with fragile otoliths, but other distinctive bones, such as Atlantic mackerel (Scomber scombrus), and Atlantic herring (Clupea harengus).[21]
See also[edit]
Ossicles
Otolithic membrane
Otolith microchemical analysis
References[edit]
Wikimedia Commons has media related to Otoliths.
^ Jump up to: a b c d Sahney, Sarda; Wilson, Mark V. H. (2001). "Extrinsic labyrinth infillings imply open endolymphatic ducts in Lower Devonian osteostracans, acanthodians, and putative chondrichthyans". Journal of Vertebrate Paleontology. 21 (4): 660–669. doi:10.1671/0272-4634(2001)021[0660:ELIIOE]2.0.CO;2.
^ Jump up to: a b Sheykholeslami, Kianoush; Kaga, Kimitaka (2002). "The otolithic organ as a receptor of vestibular hearing revealed by vestibular-evoked myogenic potentials in patients with inner ear anomalies". Hearing Research. 165 (1–2): 62–67. doi:10.1016/S0378-5955(02)00278-2. ISSN 0378-5955. PMID 12031516.
^ Jump up to: a b c Jones, Gareth P.; Lukashkina, Victoria A.; Russell, Ian J.; Lukashkin, Andrei N. (2010). "The Vestibular System Mediates Sensation of Low-Frequency Sounds in Mice". Journal of the Association for Research in Otolaryngology. 11 (4): 725–732. doi:10.1007/s10162-010-0230-7. ISSN 1525-3961. PMC 2975890. PMID 20821033.
Jump up ^ Gauldie, R. W. (1993). "Polymorphic crystalline structure of fish otoliths". Journal of Morphology. 218 (1): 1–28. doi:10.1002/jmor.1052180102.
Jump up ^ Wolff, Heinz G. (1973). "Multi-directional sensitivity of statocyst receptor cells of the opisthobranch gastropod Aplysia limacina". Marine Behavior and Physiology. 1 (1–4): 361–373. doi:10.1080/10236247209386910.
Jump up ^ Nolf, D. 1985. Otolithi Piscium; in H.-P. Schultze (ed.), Handbook of Paleoichthyology, Vol. 10. Gustav Fischer Verlag, Stuttgart, 145pp.
Jump up ^ Schultze, H.-P. (1990). "A new acanthodian from the Pennsylvanian of Utah, U.S.A. and the distribution of otoliths in gnathostomes". Journal of Vertebrate Paleontology. 10 (1): 49–58. doi:10.1080/02724634.1990.10011789.
Jump up ^ Skylab Archives, 1984.
Jump up ^ Lenhardt, M.; Skellett, R; Wang, P; Clarke, A. (1991). "Human ultrasonic speech perception". Science. 253 (5015): 82–85. Bibcode:1991Sci...253...82L. doi:10.1126/science.2063208. ISSN 0036-8075. PMID 2063208.
Jump up ^ Patterson, William P.; Smith, Gerald R.; Lohmann, Kyger C. (1993). "Continental Paleothermometry and Seasonality Using the Isotopic Composition of Aragonitic Otoliths of Freshwater Fishes". Geophysical Monograph Series: 191–202. Bibcode:1993GMS....78..191P. doi:10.1029/GM078p0191.
Jump up ^ Patterson, William P. (1999). "Oldest isotopically characterized fish otoliths provide insight to Jurassic continental climate of Europe". Geology. 27 (3): 199. doi:10.1130/0091-7613(1999)027<0199:OICFOP>2.3.CO;2.
Jump up ^ Zazzo, A.; Smith, G. R.; Patterson, W. P.; Dufour, E. (2006). "Life history reconstruction of modern and fossil sockeye salmon (Oncorhynchus nerka) by oxygen isotopic analysis of otoliths, vertebrae, and teeth: Implication for paleoenvironmental reconstructions". Earth and Planetary Science Letters. 249 (3/4): 200–215. Bibcode:2006E&PSL.249..200Z. doi:10.1016/j.epsl.2006.07.003.
Jump up ^ Reimer, T.; Dempster, T.; Warren-Myers, F.; Jensen, A. J.; Swearer, S. E. (2016). "High prevalence of vaterite in sagittal otoliths causes hearing impairment in farmed fish". Scientific Reports. 6 (25249). Bibcode:2016NatSR...625249R. doi:10.1038/srep25249.
Jump up ^ Kimura, Daniel K.; Anderl, Delsa M. (2005). "Quality control of age data at the Alaska Fisheries Science Center" (PDF). Marine and Freshwater Research. 56: 783–789. doi:10.1071/mf04141.
Jump up ^ Fish Age and Growth with Otoliths Tennessee Wildlife Resources Agency. Retrieved 2007-04-07.
Jump up ^ Bos, A.R. (1999). "Tidal transport of flounder larvae (Pleuronectes flesus) in the River Elbe, Germany". Archive of Fishery and Marine Research. 47 (1): 47–60.
Jump up ^ Arim, Matías; Naya, Daniel E. (2003). "Pinniped diets inferred from scats: Analysis of biases in prey occurrence". Canadian Journal of Zoology. 81 (1): 67–73. doi:10.1139/z02-221.
Jump up ^ Bowen, W. D. (2000). "Reconstruction of pinniped diets: Accounting for complete digestion of otoliths and cephalopod beaks". Canadian Journal of Fisheries and Aquatic Sciences. 57 (5): 898–905. doi:10.1139/f00-032.
Jump up ^ Grellier, K.; Hammond, P. S. (2005). "Feeding method affects otolith digestion in captive gray seals: Implications for diet composition estimation". Marine Mammal Science. 21 (2): 296–306. doi:10.1111/j.1748-7692.2005.tb01229.x.
Jump up ^ Browne, Patience; Laake, Jeffrey L.; DeLong, Robert L. (2002). "Improving pinniped diet analyses through identification of multiple skeletal structures in fecal samples" (PDF). Fishery Bulletin. 100 (3): 423–433.
Jump up ^ Ouwehand, J.; Leopold, M. F.; Camphuysen, C. J. (2004). "A comparative study of the diet of Guillemots Uria aalge and Razorbills Alca torda killed during the Tricolor oil incident in the south-eastern North Sea in January 2003" (PDF). Atlantic Seabirds. 6 (3): 147–163. [permanent dead link]
External links[edit]
Otolith Research Lab – Bedford Institute of Oceanography.
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