Hearing Loss Advanced Topic
Hearing loss is commonly categorized by why hearing no longer works, either as tudiesensorineural or conductive. Sensorineural hearing loss means that hearing was lost by damaging the nerves or inner ear structures. Conductive hearing loss means transmission of sound to the inner ear is impaired. Either of these types of hearing loss can come from multiple causes, such as auditory fatigue, trauma, disease, drugs, or genetics. In many cases, the cause is simply unknown. Sound exposure is an important cause of hearing loss and is the focus of the discussion below.
Hair cells are sensory receptor cells for hearing and are the structures most easily damaged by sound. They are part of the organ of Corti, which is housed in the cochlear duct inside the cochlea (see Hearing in Mammals for an introduction to how mammals hear).
The hair cells rest on top of the basilar membrane. Small hair-like extensions called stereocilia at the top of the hair cells are embedded in the tectorial membrane. Sound waves cause the basilar membrane to vibrate. This creates a differential motion between the basilar and tectorial membranes, causing the hair cell stereocilia to bend. This leads to internal changes within the hair cells that release chemicals that create electrical signals. Auditory nerve fibers synapse (connect) with the hair cells and pass these signals on to the brain.
The physical properties of the basilar membrane (its thickness and width at each end) make it particularly good at detecting frequencies. Since the width and thickness of the basilar membrane varies from the top to the bottom, it does not vibrate evenly all over from a sound wave. The apical end of the basilar membrane vibrates most at low frequency tones, and the basal end vibrates most at high frequency tones. The basilar membrane is, therefore, tonotopic, or responds in order from high (base) to low (apex) frequencies. Therefore, based on which hair cells are responding along the basilar membrane, a specific group of ganglion cells becomes active, and the brain is able to perceive what tones have been received.
The minimum level of sound that is detectable is defined as the threshold of hearing and will vary with each frequency. Thresholds are typically measured over many frequencies with repeated trials and may vary by individual over time (Hearing Sensitivity Studies). They are usually displayed as a “hearing curve” which is a graph of thresholds for the frequencies tested. An animal’s hearing can be impaired temporarily or permanently by overexposure to certain characteristics of sound.
Hearing loss from sound exposure actually depends on the sensitivity of the animal to a sound and the interaction of three characteristics of the sound: the frequency of the sound (for more information on frequency, see How do you characterize sounds? Frequency), the intensity of the sound (for more information on intensity, see How do you characterize sounds? Intensity or Loudness), and the duration or how long the animal is exposed to that sound. Exposure to a short, very loud sound or to some lower level sound over a long period of time can damage the hair cells by over-extending the stereocilia, shrinking the cell, or breaking connections between cells and cilia. If hair cell damage is minor, hair cells can recover and hearing will return to normal. If the hair cells are severely damaged, they will not recover. These effects range from minor auditory fatigue to major cell death.
If the hair cells and stereocilia do not have sufficient time to recover between sounds, the ear experiences auditory fatigue, which is also known as noise-induced hearing loss (NIHL). An animal can accommodate sounds that are above its hearing threshold until a certain combination of intensity and duration is reached. Above this limit, the animal’s hearing threshold will be elevated. If the hearing threshold returns to baseline levels then it is known as a temporary threshold shift (TTS). Temporary threshold shift (TTS) studies have been conducted with several species of marine mammals.
The degree of any threshold shift depends on the three characteristics of the sound that the animal is exposed to along with the sensitivity of each individual to the sound. These factors may vary by individual and by species (see Hearing Loss). Currently there are some generalities about the effects of sound on animal hearing thresholds:
- the degree of hearing loss from any sound is likely to differ among species and among individuals
- for pure tones, hearing loss centers around the frequency of the tone
- even for sounds that cause TTS, there may be a point at which the interaction of frequency, intensity and duration of exposure will cause an irreversible hearing loss.
If the hearing threshold does not return to baseline levels, the effect is called a permanent threshold shift (PTS). PTS can occur as a result of repeated occurrences of TTS, or it can occur catastrophically as a result of a single exposure to a very intense sound.
Acoustic trauma is severe traumatic injury from sound. Ears are especially subject to these injuries which result from the inability of a tissue to tolerate very high, sudden pressures like impulse noise from gunshots. Damage due to acoustic trauma is distinct, and scientists studying the ears of whales can identify its characteristics. Injury to the auditory system from extreme events like explosions, blows to the head, or other concussive forces are sometimes mistakenly called acoustic trauma. These injuries are not caused by sound.
- Au, W.W.L., Popper, A.N., and Fay R.R. (eds). 2000, “Hearing in Whales and Dolphins.” Springer-Verlag, Inc., New York, NY.
- Gelfand, S.A. 1998, “Hearing: An Introduction to Psychological and Physiological Acoustics.” 3rd ed. Marcel Dekker publishing, New York.
- Ketten, D.R. 2000, “Cetacean Ears.” In: Hearing by Whales and Dolphins (W.W.L. Au, A.N. Popper, and R.R. Fay, eds.) Springer-Verlag, Inc., New York, NY. p. 43-108.
- Wursig, B. and Richardson, W.J. 2002, “Effects of Noise.” In: The Encyclopedia of Marine Mammals (W.F. Perrin, B. Wursig, and J.G.M. Thewissen, eds). Academic Press, San Diego, CA.
- Yost, W.A. 1994, “Fundamentals of Hearing: An Introduction.” 3rd ed. Academic Press, New York, NY.
Hearing Curve Notes and References
- Thresholds shown for Odontocetes and Pinnipeds are a composite of measured lowest thresholds for multiple species. (Richardson, et. al., 1995)
- Range for Mysticete thresholds is estimated from mathematical models based on ear anatomy or inferred from emitted sounds, with the dashed line suggesting the region of best sensitivity (Ketten, 1994, 1998; Frankel, et. al., 1995)
- Ambient noise and sea state noise curves from Urick (1983) are spectral levels in 1 Hertz wide frequency bands, with units of dB re 1 µPa2/Hz
- Frankel, A.S., Mobley, J.R., Jr., and Herman, L.M. 1995. Estimation of auditory response thresholds in humpback whales using biologically meaningful sounds. In: Kastelein, R.A., Thomas, J.A., and Nachtigall, P.E. (eds.). Sensory Systems of Aquatic Mammals. De Spil Publishers, Woerden, Netherlands.
- Ketten, D.R. 1994. Functional analyses of whale ears: Adaptations for underwater hearing. IEEE Proceedings of Underwater Acoustics 1: 264-270.
- Ketten, D.R. 1998. Marine mammal auditory systems: A summary of audiometric and anatomical data and its implications for underwater acoustic impacts. NOAA Technical Memorandum NOAA-TM-NMFS-SWFSC-256.
- Office of Naval Research. 2001. Final Environmental Impact Statement for the North Pacific Acoustic Laboratory, May 2001. Office of Naval Research, Arlington, VA.
- Richardson, W.J., Greene, C.R., Jr., Malme, C.I., and Thomson, D.H. 1995. Marine Mammals and Noise. Academic Press. San Diego, CA.
- Urick, R.J. 1983. Principles of Underwater Sound. New York: McGraw-Hill.