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{{Short description|Animal perception of low frequency sounds}}
'''[[Infrasound]]''' is sound at frequencies lower than the low frequency end of human hearing threshold at 20 Hz. It is known, however, that humans can perceive sounds below this frequency at very high pressure levels.<ref>{{cite journal |last=Yeowart |first=N. S. |author2=M. J. Evans |title=Thresholds of audibility for very low-frequency pure tones |journal=[[J. Acoust. Soc. Am.]] |year=1974 |volume=55 |issue=4 |pages=814–818 |doi=10.1121/1.1914605 |pmid=4833076 |bibcode=1974ASAJ...55..814Y|doi-access=free }}</ref> Infrasound can come from many natural as well as man-made sources, including weather patterns, topographic features, ocean wave activity, thunderstorms, [[geomagnetic storms]], earthquakes, [[jet streams]], mountain ranges, and rocket launchings.<ref>{{cite journal |last=Cook |first=R. K. |title=Atmospheric sound propagation |journal=Atmospheric Exploration by Remote Probes |year=1969 |volume=2 |pages=633–669}}</ref><ref>{{cite journal |last=Procunier |first=R. W. |title=Observations of acoustic aurora in the 1-16 Hz range |journal=Geophys. J. R. Astron. Soc. |year=1971 |volume=26 |issue=1–4 |pages=183–189 |bibcode=1971GeoJ...26..183P |doi=10.1111/j.1365-246X.1971.tb03392.x|doi-access=free }}</ref> Infrasounds are also present in the vocalizations of some animals. Low frequency sounds can travel for long distances with very little attenuation and can be detected hundreds of miles away from their sources.<ref name="Kreithen & Quine 1979">{{cite journal |last=Kreithen |first=M. L. |author2=D. B. Quine |title=Infrasound detection by the homing pigeon: A behavioral audiogram |journal=Journal of Comparative Physiology |year=1979 |volume=129 |pages=1–4 |doi=10.1007/bf00679906|s2cid=12127549 }}</ref><ref name="Langbauer et al 1990">{{cite journal |last=Langbauer |first=W. R. |author2=K. B. Payne |author3=R. A. Charif |author4=E. M. Thomas |title=Responses of captive African elephants to playback of low-frequency calls |journal=Canadian Journal of Zoology |year=1990 |volume=67 |issue=10 |pages=2604–2607 |doi=10.1139/z89-368 }}</ref>
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=== Elephants ===
Elephants are the terrestrial animal in which the production of infrasonic calls was first noted by M. Krishnan,<ref name="Krishnan 1972">{{cite journal |last=Krishnan |first=M |title=An Ecological Survey of the Larger Mammals of Peninsular India |year=1972 |journal=The Journal of the Bombay Natural History Society |volume=69 |pages=26–54}}</ref> later discovered by Katy Payne.<ref>{{cite book |last=Payne |first=Katy |title=Silent Thunder: In the presence of Elephants |url=https://archive.org/details/silentthunderinp00payn |url-access=registration |year=1998 |publisher=Simon & Schuster |___location=New York|isbn=9780684801087 }}</ref> The use of low frequency sounds to communicate over long distances may explain certain elephant behaviors that have previously puzzled observers. Elephant groups that are separated by several kilometers have been observed to travel in parallel or to change the direction simultaneously and move directly towards each other in order to meet.<ref name="Langbauer et al 1991">{{cite journal |last=Langbauer |first=W. R. |author2=K. B. Payne |author3=R. A. Charif |author4=L. Rapaport |author5=F. Osborn |title=African elephants respond to distant playbacks of low-frequency conspecific calls |journal=J. Exp. Biol. |year=1991 |volume=157 |pages=35–46|doi=10.1242/jeb.157.1.35 }}</ref> The time of [[estrus]] for females is asynchronous, lasts only for a few days, and occurs only every several years. Nevertheless, males, which usually wander apart from female groups, rapidly gather from many directions to compete for a receptive female.<ref name="Langbauer et al 1991" /> Since infrasound can travel for very long distances, it has been suggested that calls in the infrasonic range might be important for long distance communication for such coordinated behaviors among separated elephants.<ref name="Langbauer et al 1991" /><ref name="Payne et al 1986">{{cite journal |last=Payne |first=K. B. |author2=W. R. Langbauer |author3=E. M. Thomas |title=Infrasonic calls of the Asian elephant (Elephas maximus |journal=Behav. Ecol. Sociobiol. |year=1986 |volume=18 |issue=4 |pages=297–301 |doi=10.1007/bf00300007|bibcode=1986BEcoS..18..297P |s2cid=1480496 }}</ref>
==== Infrasound production and perception ====
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==== Infrasound sensitivity ====
The auditory sensitivity thresholds have been measured behaviorally for one individual young female Indian elephant. The [[Classical conditioning|conditioning]] test for sensitivity requires the elephant to respond to a stimulus by pressing a button with its trunk, which results in a sugar water reward if the elephant correctly identified the appropriate stimulus occurrence.<ref name="Heffner & Heffner 1980">{{cite journal |last=Heffner |first=H. |author2=R. Heffner |title=Hearing in the elephant (Elephas maximus) |journal=Science |year=1980 |volume=208 |pages=518–520 |doi=10.1126/science.7367876 |pmid=7367876 |issue=4443 |bibcode=1980Sci...208..518H}}</ref> To determine auditory sensitivity thresholds, a certain frequency of sound is presented at various intensities to see at which intensity the stimulus ceases to evoke a response. The auditory sensitivity curve of this particular elephant began at 16 Hz with a threshold of 65 dB. A shallow slope decreased to the best response at 1 kHz with a threshold of 8 dB, followed by a steep threshold increase above 4 kHz. According to the 60 dB
The ability to differentiate frequencies of two successive tones was also tested for this elephant using a similar conditioning paradigm. The elephant's responses were somewhat erratic, which is typical for mammals in this test.<ref name="Heffner & Heffner 1980" /> Nevertheless, the ability to discriminate sounds was best at frequencies below 1 kHz particularly at measurements of 500 Hz and 250 Hz.<ref name="Heffner & Heffner 1980" />
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== Birds ==
Although birds do not produce vocalizations in the infrasonic range, reactions to infrasonic stimuli have been observed in several species, such as the [[homing pigeon]], the [[guineafowl]], and the Asian grouse.<ref>{{cite journal |last=Yodlowski |first=M. L. |author2=M. L. Kreithen |author3=W. T. Keeton |title=Detection of atmospheric infrasound by pigeons |journal=Nature |year=1977 |volume=265 |pages=725–726 |doi=10.1038/265725a0 |issue=5596 |pmid=859577 |bibcode=1977Natur.265..725Y|s2cid=4247969 }}</ref><ref>{{cite journal |last=Theurich |first=M. |author2=G. Langner |author3=H. Scheich |title=Infrasound re-sponses in the midbrain of the Guinea Fowl |journal=Neurosci Lett |year=1984 |volume=49 |issue=1–2 |pages=81–86 |doi=10.1016/0304-3940(84)90140-x|pmid=6493602 |s2cid=36335442 }}</ref><ref>{{cite journal |last=Moss |first=R. |author2=I. Lockie |title=Infrasonic components in the song of the Capercaillie Tetrao urogallus |journal=Ibis |year=1979 |volume=121 |pages=95–97 |doi=10.1111/j.1474-919x.1979.tb05021.x}}</ref> It is postulated that birds might use the detection of naturally occurring infrasound for long-range directional cues from distant landmarks, or for weather detection.<ref name="Quine 1981">{{cite journal | last1 = Quine | first1 = Douglas B. | year = 1981 | title = Frequency shift discrimination: Can homing pigeons locate infrasounds by Doppler shifts? | journal = Journal of Comparative Physiology | volume = 141 | issue = 2 | page = 2 | doi=10.1007/bf01342661| s2cid = 40421698 }}</ref> Since hearing tests at infrasonic frequencies have been conducted on a small number of bird species, the true diversity of this ability among birds is unknown.<ref>{{Cite journal|last1=Zeyl|first1=Jeffrey N.|last2=Ouden|first2=Olivier den|last3=Köppl|first3=Christine|last4=Assink|first4=Jelle|last5=
=== Pigeons ===
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==== Infrasound sensitivity ====
In experiments using heart
In order to use infrasound for navigation, it is necessary to be able to localize the source of the sounds. The known mechanisms for sound localizations make use of the time difference cues at the two ears. However, infrasound has such long wavelengths that these mechanisms would not be effective for an animal the size of a pigeon. An alternative method that has been hypothesized is through the use of the [[Doppler shift]].<ref name="Quine 1981" /> A Doppler shift occurs when there is relative motion between a sound source and a perceiver and slightly shifts the perceived frequency of the sound. When a flying bird is changing direction, the amplitude of the Doppler shift between it and an infrasonic source would change, enabling the bird to locate the source. This kind of mechanism would require the ability to detect very small changes in frequency. A pigeon typically flies at 20 km/h, so a turn could cause up to a 12% modulation of an infrasonic stimulus. According to response measurements, pigeons are able to distinguish frequency changes of
In early experiments with infrasound sensitivity in pigeons, surgical removal of the columella, a bone that links the [[tympanic membrane]] to the [[inner ear]], in each ear severely reduced the ability to respond to infrasound, increasing the sensitivity threshold by about 50 dB. Complete surgical removal of the entire [[cochlea]], lagena, and columellae completely abolished any response to infrasound.<ref name="Kreithen & Quine 1979" /> This shows that the receptors for infrasonic stimuli may be located in the inner ear.
==== Infrasound
Neural fibers that are sensitive to infrasonic stimuli have been identified in the pigeon and their characteristics have been studied. It turns out that, although these fibers also originate in the inner ear, they are quite different from normal acoustic fibers.
Infrasound sensitive fibers have very high rates of spontaneous discharge, with a mean of
By injecting fibers that were identified to be sensitive to infrasound with HRP (Horseradish Peroxidase), the ___location and morphology of the stained fibers can be observed in sections under a microscope. Infrasound sensitive fibers are found to be simple bipolar cells in the [[auditory ganglion]] with a diameter of {{nowrap|1.6
== References ==
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*{{cite journal | last1 = Moss | first1 = R | last2 = Lockie | first2 = I | year = 1979 | title = Infrasonic components in the song of the Capercaillie Tetrao urogallus | journal = Ibis | volume = 121 | pages = 95–97 | doi=10.1111/j.1474-919x.1979.tb05021.x}}
*Payne, Katy. Silent Thunder: In the presence of Elephants. New York: Simon & Schuster, 1998
*{{cite journal | last1 = Payne | first1 = K. B. | last2 = Langbauer | first2 = W. R. Jr. | last3 = Thomas | first3 = E. M. | year = 1986 | title = Infrasonic calls of the Asian elephant (Elephas maximus) | journal = Behav. Ecol. Sociobiol. | volume = 18 | issue = 4 | pages = 297–301 | doi=10.1007/bf00300007| bibcode = 1986BEcoS..18..297P | s2cid = 1480496 }}
*{{cite journal | last1 = Quine | first1 = Douglas B. | year = 1981 | title = Frequency shift discrimination: Can homing pigeons locate infrasounds by Doppler shifts? | journal = Journal of Comparative Physiology | volume = 141 | issue = 2 | page = 2 | doi=10.1007/bf01342661| s2cid = 40421698 }}
*{{cite journal | last1 = Schermuly | first1 = L. | last2 = Klinke | first2 = R. | year = 1990 | title = Infrasound sensitive neurons in the pigeon cochlear ganglion | journal = Journal of Comparative Physiology A | volume = 166 | issue = 3 | pages = 355–363 | doi=10.1007/bf00204808| pmid = 2324994 | s2cid = 12962156 }}
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[[Category:Behavioral neuroscience]]
[[Category:Ethology]]
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