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Auditory feedback

From Wikipedia, the free encyclopedia

Auditory feedback (AF) is an aid used by humans to control speech production and singing by helping the individual verify whether the current production of speech or singing is in accordance with his acoustic-auditory intention. This process is possible through what is known as the auditory feedback loop, a three-part cycle that allows individuals to first speak, then listen to what they have said, and lastly, correct it when necessary. From the viewpoint of movement sciences and neurosciences, the acoustic-auditory speech signal can be interpreted as the result of movements (skilled actions) of speech articulators (the lower jaw, lips, tongue, etc.). Auditory feedback can hence be inferred as a feedback mechanism controlling skilled actions in the same way that visual feedback controls limb movements (e.g. reaching movements).

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Transcription

Hello! I’d like you to think about how I’m doing this right now. Not why I’m doing it, because of course, I’m doing it because I like music and I like science and I like to do both those things at the same time. But how can I play music? How can I be hearing it right now? And how can I walk around and play my guitar at the same time without falling on my face? And what is even sound anyway? These are all good questions. Let’s start with the last one, first. The basic answer to “What is sound?” goes like this: Sounds create vibrations in the air that beat against the eardrum, which pushes a series of tiny bones that move internal fluid against a membrane that triggers tiny hair cells -- which aren’t actually hairs -- that stimulate neurons, which in turn send action potentials to the brain, which interprets them as sound. But there’s a lot more to our ears than allowing us to experience the pleasure of birdsong, or the pain of grindcore. The ear’s often overlooked, but even more vital role is maintaining your equilibrium, and without THAT, you wouldn’t be able to dance or strut or even stand up. And you definitely could not do this! At least not without throwing up. In order to really get to the nitty-gritty of how your ears pick up sound, you’ve got to understand how sound works. The key to sound transmission is vibration. When I talk, my vocal folds vibrate. When I slap this table top, or strum a guitar, those vibrations cause air particles to vibrate too, initiating sound waves that carry the vibration through the air. So this, sounds different than this, because different vibrating objects produce differently shaped sound waves. A sound’s frequency is the number of waves that pass a certain point at a given time. A high-pitched noise is the result of shorter waves moving in and out more quickly, while fewer, slower fluctuations result in a lower pitch. How loud a sound registers depends on the wave’s amplitude, or the difference between the high and low pressures created in the air by that sound wave. Now, in order for you to pick up and identify sounds from beeping to barking to Beyonce, sound waves have to reach the part of the ear where those frequencies and air-pressure fluctuations can register and be converted into signals that the brain can understand. So once again, it all boils down to action potentials. But, how does sound get in there? Your ear is divided into three major areas: the external, middle, and inner ear. The external and middle ear are only involved with hearing, while the complex hidden inner is key to both hearing and maintaining your equilibrium. So the pinna, or auricle, is the part that you can see, and wiggle, and grab, or festoon with an earring. It’s made up of elastic cartilage covered in skin, and its main function is to catch sound waves, and pass them along deeper into the ear. Once a sound is caught, it’s funneled down into the external acoustic meatus, or auditory canal, and toward your middle and inner ear. Sound waves traveling down the auditory canal eventually collide with the tympanic membrane, which you probably know as the eardrum. This ultra-sensitive, translucent, and slightly cone-shaped membrane of connective tissue is the boundary between the external and middle ear. When the sweet sound waves of your favorite jam collide with the eardrum, they push it back and forth, making it vibrate so it can pass those vibrations along to the tiny bones in the middle ear. Now, the middle ear, also called the tympanic cavity, is the relay station between the outer and inner ear. Its main job is to amplify those sound waves so that they’re stronger when they enter the inner ear. And it’s gotta amplify them, because the inner ear moves sound through a special fluid, not through air -- and if you’ve ever gone swimming you know that moving through a liquid can be a lot harder than moving through air. The tympanic cavity focuses the pressure of sound waves so that they’re strong enough to move the fluid in the inner ear. And it does this using the auditory ossicles -- a trio of the smallest, and most awesomely named bones in the human body: the malleus, incus, and stapes, commonly known as the hammer, anvil, and stirrup. One end of the malleus connects to the inner eardrum and moves back and forth when the drum vibrates. The other end is attached to the incus, which is also connected to the stapes. Together they form a kind of chain that conducts eardrum vibrations over to another membrane -- the superior oval window -- where they set that fluid in the inner ear into motion. The inner ear is where things get a little complicated, but interesting and also kind of mysterious. With some of the most complicated anatomy in your entire body, it’s no wonder it’s known as the labyrinth. This tiny, complex maze of structures is safely buried deep inside your head, because it’s got two really important jobs to do: One, turn those physical vibrations into electrical impulses the brain can identify as sounds. And two: help maintain your equilibrium so you are continually aware of which way is up and down, which seems like a simple thing, but it is very important. To do this, the labyrinth actually needs two layers -- the bony labyrinth, which is the big fluid-filled system of wavy wormholes -- and the membranous labyrinth, a continuous series of sacs and ducts inside the bony labyrinth that basically follows its shape. Now, the hearing function of the labyrinth is housed in the easy-to-spot structure that’s shaped like a snail’s shell, the cochlea. If you could unspool this little snail shell, and cut it in a cross-section, you’d see that the cochlea consists of three main chambers that run all the way through it, separated by sensitive membranes. The most important one -- at least for our purposes -- is the basilar membrane, a stiff band of tissue that runs alongside that middle, fluid-filled chamber. It’s capable of reading every single sound within the range of human hearing -- and communicating it immediately to the nervous system, because right smack on top of it is another long fixture that’s riddled with special sensory cells and nerve cells, called the organ of corti. So when your cute little ossicle bones start sending pressure waves up the inner fluid, they cause certain sections of basilar membrane to vibrate back and forth. This membrane is covered in more than 20,000 fibers, and they get longer the farther down the membrane you go. Kind of like a harp with many, many strings, the fibers near the base of the cochlea are short and stiff, while those at the end are longer and looser. And, just like harp strings, the fibers resonate at different frequencies. More specifically, different parts of the membrane vibrate, depending on the pitch of the sound coming through. So the part of the membrane with the short fibers vibrates in response to high-frequency pressure. And the areas with the longer fibers resonate with lower-frequency waves. This means that, all of the sounds that you hear -- and how you recognize them -- comes down to precisely what little section of this membrane is vibrating at any given time. If it’s vibrating near the base, then you’re hearing a high-frequency sound. If it’s shakin’ at the end, it’s a low noise. But of course nothing’s getting heard until something tells the brain what’s going on. And the transduction of sound begins when part of the membrane moves, and the fibers there tickle the neighboring organ of corti. This organ is riddled with so-called hair cells, each of which has a tiny hair-like structure sticking out of it. And when one is triggered, it opens up mechanically gated sodium channels. That influx of sodium then generates graded potentials, which might lead to action potentials, and now your nervous system knows what’s going on. Those electrical impulses travel from the organ of corti along the cochlear nerve and up the auditory pathway to the cerebral cortex. But the information that the brain gets is more than just, like, “hey listen up.” The brain can detect the pitch of a sound based solely on the location of the hair cells that are being triggered. And louder sounds move the hair cells more, which generates bigger graded potentials, which in turn generate more frequent action potentials. So the cerebral cortex interprets all those signals, and also plugs them into stored memories and experiences, so it can finally say oh, that’s a chickadee, or a knock at the door, or the slow burn of an 80s saxophone solo, or whatever. So that’s how you hear. But we’re not done with you yet -- we gotta talk about equilibrium. The way we maintain our balance works in a similar way to the way we hear, but instead of using the cochlea, it uses another squiggly structure in the labyrinth that looks like it’s straight out of an Alien movie -- a series of sacs and canals called the vestibular apparatus. This set-up also uses a combination of fluid and sensory hair cells. But this time, the fluid is controlled not by sound waves but by the movement of your head. The most ingenious parts of this structure are three semicircular canals, which all sit in the sagittal, frontal, and transverse planes. Based on the movement of fluid inside of them, each canal can detect a different type of head rotation, like side-to-side, and up-and-down, and tilting, respectively. And every one of the canals widens at its base into sac-like structures, called the utricle and saccule, which are full of hair cells that sense the motion of the fluid. So by reading the fluid’s movement in each of the canals, these cells can give the brain information about the acceleration of the head. So if I move my head like this, because I’m, like, super into my jam, that fluid moves and stimulates hair cells that read up and down head movement, which then send action potentials along the acoustic nerve to my brain, where it processes the fact that I’m bobbing my head. And, just as your brain interprets the pitch and volume of a sound by both where particular hair cells are firing in the cochlea and how frequent those action potentials are coming in, so too does it use the location of hair cells in the vestibular apparatus to detect which direction my head is moving through space, and the frequency of those action potentials to detect how quickly my head is accelerating. But things can get messy. Doing stuff like spinning on a chair, or sitting on a rocky boat, can make you sick because it creates a sensory conflict. In the case of me spinning around on my chair, the hair cells in my vestibular apparatus are firing because of all that inner-ear fluid sloshing around — but the sensory receptors in my spine and joints tell my brain that I’m sitting still. On a rocking boat, my vestibular senses say I’m moving up and down, but if I’m looking at the deck, my eyes are telling my brain that I’m sitting still. The disconnect between these two types of movement, by the way, is why we get motion sickness. It doesn’t take long for my brain to get confused, and then mad enough at me to make me barf. Aaand I’m sorry that we’re ending with barf. But, we are. Today your ears heard me tell you how your cochlea, basilar membrane, and hair cells register and transduct sound into action potentials. You also learned how different parts of your vestibular apparatus respond to specific motions, and how that helps us keep our equilibrium. Special thanks to our Headmaster of Learning Thomas Frank for his support for Crash Course and for free education. Thank you to all of our Patreon patrons who make Crash Course possible through their monthly contributions. If you like Crash Course and want to help us keep making great new videos like this one -- and get some extra special, interesting stuff -- you can check out patreon.com/crashcourse Crash Course is filmed in the Doctor Cheryl C. Kinney Crash Course Studio. This episode was written by Kathleen Yale, edited by Blake de Pastino, and our consultant is Dr. Brandon Jackson. Our director is Nicholas Jenkins, the script supervisor and editor is Nicole Sweeney, our sound designer is Michael Aranda, and the graphics team is Thought Café.

Speech

Auditory feedback allows one to monitor their speech and rectify production errors quickly when they identify one, making it an important component of fluent speech productions.[1] The role of auditory feedback on speech motor control is often investigated by exposing participants to frequency-altered feedback. Inducing brief and unpredictable changes in the frequency of their auditory feedback has consistently been shown to induce a "pitch-shift reflex", which suggests that this reflex aids in stabilizing voice frequency around the desired target.[2][3]

However, due to the fact that auditory feedback needs more than 100 milliseconds before a correction occurs at the production level,[4] it is a slow correction mechanism in comparison with the duration (or production time) of speech sounds (vowels or consonants). Thus, auditory feedback is too slow to correct the production of a speech sound in real-time. Nonetheless, it has been shown that auditory feedback is capable of changing speech-sound production over a series of trials (i.e. adaptation by relearning; see e.g. perturbation experiments done with the DIVA model: neurocomputational speech processing). 10 minutes is typically sufficient for a nearly-full adaptation. Research has also shown that auditory linguistic prompts resulted in greater correction to acoustic perturbations than non-linguistic prompts, reflecting the decrease in accepted variance for intended speech when external linguistic templates are available to the speaker.[5]

Speech Acquisition and Development

Auditory feedback is an important aid during speech acquisition by toddlers, by providing the child with information about speech outcomes that are used to pick-up and eventually hone speech motor planning processes. Auditory inputs are typically produced by a communication partner (e.g. caretaker) and heard by the toddler, who subsequently tries to imitate them.[6][7] Children as young as the age of four have demonstrated the ability to adapt speech motor patterns to perceived changes in vowel auditory feedback, which enables them to maintain the accuracy of their speech output.[8] However, children's speech motor adaption abilities are not fully optimised due to their limited auditory perceptual skills. Thus, improvements in children's ability to perceive relevant acoustic property will usually be followed by an improvement in their speech adaption performance.[9]

Individuals who are born deaf often fail to acquire fluent speech, further reinforcing how auditory feedback plays a crucial role in speech acquisition and development.[10]

Delayed auditory feedback experiments indicate that auditory feedback is important during speech production, even in adults. It has been shown that severe disfluencies in speech occur when the timing of voice feedback is delayed for a normal speaker.[11][12] Individuals who become deaf post-lingually and are unable to receive vocal feedback anymore also typically experience a deterioration in speech quality,[13][14] highlighting the importance of auditory feedback in speech formation throughout one's lifetime.

Impacts on speech disorders

Stuttering

Stuttering is said to be due to ineffective monitoring of auditory feedback, mainly caused by a deficit in the cortical auditory system modulation during speech planning.[15] When fluent speakers detect a sudden irregularity in a specific acoustic parameter of their auditory feedback, they are able to instantly correct the error in their speech production. Individuals who stutter, on the other hand, are found to have weaker-than-normal abilities to correct such errors.[16] Individuals that stutter hence demonstrate ineffective auditory comparisons of desired speech movements, as compared to fluent speakers.[17]

Delayed auditory feedback has been found to be an effective treatment for some individuals who stutter,[18] since extending the time between speech and auditory perception allows for more time to process and correct errors.

Apraxia of speech

It is posited that individuals with apraxia of speech have weak feedforward programs, which results in the disfluencies of their speech.[19] These individuals hence develop a heavy reliance on auditory feedback to minimize and repair speech errors[20] even in later stages of their lives, whilst fluent speakers easily transitions from feedback dependent to feedforward-dominant.[21] This is not ideal since heavy reliance on mostly auditory feedback is said to be inefficient for the production of rapid and accurate speech.[22]

Auditory masking has been found to decrease disfluency duration and increase vocal intensity as well as syllable rate in some individuals with apraxia of speech.[23] Since apraxia of speech is said to be due to weak feedforward programs and high dependence on auditory feedback, auditory masking can be reasoned to increase fluency by decreasing the frequency of a speaker attending auditorily to speech errors, and hence reducing the likelihood of disfluency-generating corrections.

Impacts on Visually Impaired individuals

Enhanced auditory processing can be observed in individuals with visual impairment, who partially compensate for their lack of vision with greater sensitivity in their other sensories.[24] Their increased sensitivity to auditory feedback allows them to demonstrate impressive spatial awareness despite their lack of sight.[25][26][27]

Desktop Assistance

Studies have shown that when vision is no longer the primary source for obtaining information, focus shifts from vision to hearing in the desktop environment.[28] Currently, there are assistive technologies such as screen readers, which aids visually impaired individuals in obtaining information on their desktop screens via auditory feedback (E.g. JAWS[29]). The assistance can come in the form of either speech based auditory feedback or non-speech based auditory feedback. Speech based interfaces are based on human speech, whilst non-speech based interfaces are based on environmental sounds such as music or artificial sound effects.

For the visually impaired, sole reliance on speech based auditory feedback imposes a heavier cognitive load which is irritating for users.[30] In contrast, non-speech auditory feedback is pleasant and conveys information more quickly, but lacks detailed information in their conveyance and training is required to understand the cues. Hence, the most ideal interface currently is adaptive auditory feedback, which automatically transitions between speech and non-speech cues based on the user state. Such an interface has been found to be more comfortable and generates higher satisfaction amongst visually impaired users.[31]

Impacts on other disorders

Graphomotor learning in writing disorders

A trial was conducted to explore whether auditory feedback had an influence on learning how to write. It was found that in adults, auditory feedback enabled the writer to better discern their writing motions. This resulted in an increase in flow and quickness of writing when using sounds to learn the writing of new characters.[32] Subsequent studies then tested the use of auditory feedback as an aid for children with dysgraphia to learn how to write. It was found that after multiple sittings of using auditory feedback while writing, children could write more smoothly, rapidly and clearly.[33]

Products based on auditory feedback principles have been invented to aid individuals with such writing disorders. Children with speech disorders can also benefit from such products. For example, a headphone called Forbrain[34] uses a bone conductor and a series of dynamic filters to correct the perception of one's own voice. This improves concentration, attention, speech, coordination, and other sensory functions. It was awarded by the BETT Show[35] in 2015 in the category "ICT Special Educational Needs Solutions".

Motor learning in movement disorders

Patients with cerebral palsy have little walking capability, due to limitations of their nervous system.[36] Auditory feedback in the form of periodic audio signals was found to have a significant improvement on the gait of patients, with several explanations proposed. One model argues that auditory feedback acts as an additional information channel for the motor systems, thereby decreasing the onset of motor faults and refining the gait of patients.[37] Another model posits that audio signals influence the gait of patients by directing motion patterns, such as heel strike timings. By wearing a device that provides immediate auditory feedback on the quality of one's gait, children with cerebral palsy learned to set down their feet in proper ways that avoided the sounds created when negative gaits were detected.[36]

Social interaction and motor coordination learning in behavioural disorders

The use of an auditory feedback-based treatment is found to have improved on the social interaction, mimicking and coordination skills of children with autism spectrum disorder.[38] This is achieved through a software which uses sensors to track the body motions of children. Each gesture made will activate a voice recording articulating pieces of sentences.[38] Children then have to reorder these sentence pieces to form a storyline. Different indicators of coordination such as motion quantity and speed were also recorded to keep track of the child's improvement through these auditory cues.[38]

Impacts on Music Performance

Instrument Performance

Auditory feedback is important in the picking up of a new musical piece. By exposing beginner piano players to irregular auditory feedback, they make more mistakes as compared to those who are given logical and anticipatable auditory feedback.[39] Learning in the presence of auditory feedback also improved one's recollection of the musical piece.[40]

However, multiple studies have shown that even without auditory feedback, there is not much disturbance to the performance of seasoned musicians.[41][42] In the absence or delay of auditory feedback, musicians turn to auditory imagery to direct their performance.[42] Other forms of feedback can also be used in compensation instead, such as visual feedback where musicians look at their hands to lead their performance.[43] Major disturbances were only seen in the area of pedaling, where results have shown that pianists were prone to stepping the pedal less often in the absence of auditory feedback.[42]

Singing

The importance of auditory feedback in the case of human singing is reviewed by Howell.[44] In the context of singing, it is important for singers to maintain pitch accuracy, even when they are drowned out by orchestral accompaniment or by fellow singers. Many studies have looked into the effects of both external auditory feedback and proprioception (also known as internal feedback) on pitch control. It has been found that external auditory feedback is crucial in maintaining pitch accuracy, especially for adults without voice training.[45][46] This is further supported by recent research which revealed how non-professional singers show lower pitch accuracy when they receive lesser auditory feedback. However, the research also highlighted how the pitch of professional singers remains almost unaffected by auditory feedback since they are able to rely on their internal feedback after years of training.[47]

Bird songs

The role of auditory feedback in the learning and production of bird-song has been studied in several research papers. It has been found that songbirds rely on auditory feedback to compare the sounds that they make with inborn tunes or songs that they memorize from others.[48] Numerous studies have shown that without the ability to hear themselves, songbirds develop erratic songs or show a deterioration in the songs that they sing after experiencing hearing loss.[49][50] Several scientific models have been put forward to explain the worsening of birdsongs after the loss of hearing. (E.g. see Brainard and Doupe's (2000) error adjustment channel in the anterior forebrain: auditory feedback in birdsong learning).[49]

However, the decline of birdsong quality can vary greatly between different demographics. For example, other studies have found the songs of older songbirds remained consistent, or had a slower rate of deterioration after going deaf.[51] Some researchers have attributed to songbirds learning how to use other forms of non-auditory feedback such as sensory information to maintain the quality of their songs.[50] This process is called sensory-motor coupling. Others have argued that older songbirds have a longer access to auditory feedback to learn their songs, which results in more practice and thus more stable production of songs even after deafening.[51]

See also

References

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