Presentation on theme: "Audition Chris Rorden Deb Hall, MRC Institute of Hearing Research"— Presentation transcript:
1 Audition Chris Rorden Deb Hall, MRC Institute of Hearing Research Anatomy and function of the auditory systemBrainstem disordersWord deafnessAmusiaEnvironmental sound agnosiaAuditory neglect and extinction
2 Anatomy and function source : Ashmore, 2002 The ‘ear’ is a complexphysiological apparatus,not just the visibleouter ear- Reflection of sound in pinna (earlobe) provides spectral cues about elevation of a sound source- Middle ear is a cavity containing an ossicular lever which matches the acoustical impedance of the inner ear so sound energy is effectively transmitted (60%)- Inner ear contains the cochlea where sound is converted into a neural signal
4 Localization and shadowing High frequencies: intensity differences: louder if sound is not in head’s sound shadowLow frequencies: Inter-aural timing differencesElevation: Frequencies influenced by location relative to pinna.
5 Middle Ear - Ossicles 3 of the smallest bones Malleus (hammer)Incus (anvil)Stapes (stirrup)Ossicular chain: Transmits acoustic energy from tympanic membrane to inner earDelivers sound vibrations to inner ear fluidChanges impedance: large, weak movement of ear drum turned to small, forceful movement in cochlear liquid.Muscles can dampen response: Prevents the inner ear from being overwhelmed by excessively strong vibrations
8 Travelling waveAlways starts at the base of the cochlea and moves toward the apexIts amplitude changes as it traverses the length of the cochleaThe position along the basilar membrane atwhich its amplitude is highest depends on the frequency of the stimulus
9 High frequencies have peak influence near base and stapes Traveling waveHigh frequencies have peak influence near base and stapesLow frequencies travel further, have peak near apexA short movie:Green line shows 'envelope' of travelling wave: at this frequency most oscillation occurs 28mm from stapes.
10 Anatomy and function source : Hackney, 2002 Many sound features are encoded before the signal reaches the cortexCochlear nucleus segregates sound informationSignals from each ear converge on the superior olivary complex - important for sound localizationInferior colliculus is sensitive to location, absolute intensity, rates of intensity change, frequency - important for pattern categorizationDescending cortical influences modify the input from the medial geniculate nucleus - important as an adaptive ‘filter’cortexmedial geniculatebodyinferior colliculuscochlear nucleuscomplexcochleasuperior olivary complex
12 Function source : Palmer & Hall, 2002 Numerous bilateral regions are frequency-dependentOverlapping regions are sensitive to intensity and to the temporal changes in soundOne region is sensitive to the spatial properties of sound (R>L)Speech also activates these regions, but neurons are probably responding to the complex acoustic properties in the sound.Perceptual attributes may be importantRighthemisphereSlow-rate temporal pattern in sound
13 Sound intensity and activation Loud sounds (90 dB) activated posterior and medial temporal gyrus (red)Soft (70 dB) sounds activated area (yellow) is found most laterally of TTGMedium intensity (82 dB) sounds activated intermediate area (green). (NeuroImage 2002;17: 710)
14 Auditory neuropsychology Simple modularity of function not clearly apparent- No auditory equivalents of V4 (visual colour area), V5 (visual motion area), fusiform face area etc- Cortical neurons respond to a complex array of stimulus features, and the temporal pattern of those features is importantUnlike visual or somatomotor systems- A lot of auditory processing is supported by the ascending pathway- Studies in several mammalian species have demonstrated that bilateral ablations of the auditory cortex have little effect on simple sound intensity and frequency-based behaviours
15 Brainstem disorders source : Griffiths et al. 1999 Brainstem = cochlear nucleus, superior olivary complex, inferior colliculusLesions rarely compatible with lifeMultiple sclerosis can affect brainstem- Complete deafness is rare- MS patients do not report problems in everyday sound perception- Few systematic studies- Deficit in perceiving frequency changes- Deficit in detecting a gap in noise- Deficit in processing binaural cues for sound localisation
16 A deficit in recognition Auditory agnosiaA deficit in recognitionPerceptionRecognitionAcoustical analysisRepresentationsAuditoryinput“Apperceptiveagnosia”“Associativeagnosia”Auditory agnosia is of this type
17 Auditory agnosia source : Griffiths et al. 1999 Normal brainstem processingMidbrain impairment questionableCortical deficit in perception- Preserved hearing (pure tones)- Disordered perception of certain sounds :Speech - word deafnessMusic - amusiaEnvironmental sounds - environmental sound agnosia
18 A case of word deafness source: Ellis & Young, 1988 Hemphill and Stengel (1940)“I can hear you dead plain, but I cannot get what you say. The noises are not quite natural. I can hear but not understand”- Normal pure tone audiometry- Fluent speech “no errors of grammar beyond what is common for his particular dialect and standard of education”- Normal reading- Normal writing and spelling- Poor spoken word repetition- Gross asymmetry between spoken and written word comprehension
19 Word deafness source : Ellis & Young, 1988 Associated symptoms- Some hearing loss (> 20 dB HL)- Production (Broca’s) aphasia- Perception of melody- Perception of environmental soundsLesion site- Generally large bilateral infarcts- When unilateral, it’s more often the left hemisphere- Involves superior temporal lobe (non-primary auditory cortex)- May or may not involve Heschl’s gyrus (primary auditory cortex)
20 Word deafness 8 Frequency (kHz) Time - filtered harmonic sounds, broad band noise, silent gaps- transitions in amplitude and frequency on three time scales (milliseconds, 10s of milliseconds, seconds)These temporal transitions are rapid and complexTime
21 Word deafness source : Ellis & Young, 1988 Inability to make fine temporal discriminations and track rapidly-changing acoustic signals?“There may be nothing speech specific about the impairment” Ellis & Young, 1988
22 A case of amusia source : Peretz, 1993 Patient CNSymptoms- Unable to recognise even simplest tune- Unable to sing children’s songs that she had known well- No deficit in everyday verbal communication- No deficit in everyday recognition of environmental soundsLesion site- Bilateral temporal lobe damage- When unilateral, it’s more often the right hemisphere
23 Dissociation within musical perception Amusia source : Peretz, 1993Dissociation within musical perception- Right injury - Deficit in melody perception: the variations in pitch- Left injury - Deficit in rhythm perception: the temporal organisation of melody over 100s of milliseconds or seconds time scale
24 AmusiaFrequencyTimeAs in speech, music contains discrete harmonic sounds that vary over timemelody: local variation in features from note to noterhythm: global variations in note duration that relate to a higher order pattern
25 Environmental sound agnosia source : Griffiths et al. 1999Deficit rarely occurs in isolationEnvironmental sounds contain fewer changes in acoustic structure over time than an equivalent length segment of speech or music
26 A common deficit? No! source : Peretz, 1993 Word deafness, amusia and environmental sound agnosia are distinct- speech and music can dissociate after brain damage- music and environmental sounds can dissociate after brain damage- environmental sound perception can be selectively spared- recovery can follow different patterns (e.g. environmental sounds, then music then speech or in the reverse order)
27 A common deficit? Yes! source : Griffiths et al. 1999 Word deafness, amusia and environmental sound agnosia probably co-occur- May not always be report because not all abilities are testedAll 3 types of sound contain a mixture of acoustic featuresDeficit in an intermediate level of analysis, which is rarely tested- Analysing the spectro-temporal pattern in sound
28 Auditory neglect source : Pavani et al., 2003 Symptoms(a) Rightward biases in sound localization(b) Poor relative judgements for sounds on the contralesional side(c) Poor elevation judgements for sounds on the contralesional sideFailure to detect contralesional sound, when presented concurrentlyPoor allocation of attention to sounds separated in time
29 Auditory neglect source : Pavani et al., 2003 Lesion site – usually right hemisphere- inferior parietal lobe- superior temporal gyrus- temporo-parietal junction
30 Many neglect patients exhibit auditory, as well as visual, deficits. Auditory & visual neglect : A common deficit? Yes! source : Pavani et al. 2003Many neglect patients exhibit auditory, as well as visual, deficits.Correlation between severity of clinical visual neglect and experimental auditory neglect measures.“Neglect can often be caused by damage to brain regions containing multisensory representations of space, with the deficit consequently manifesting across multiple sensory modalities, with correlated severity”.visual deficitauditory deficit
31 Visual extinction source : Rorden et al., 1997 Symptom- a chronic bias of spatial attention towards the ipsilesional sideHence, ipsilesional events are perceived earlier than physically synchronous contralesional stimuli. This can be measured using the temporal order judgements test.
32 Visual extinction source : Karnath et al., 2002 The same deficit is also found in audition……and over thesame time scale(~ 200 ms)
33 ...but delay is of the same time scale Auditory & visual extinction : A common deficit? Possibly! source : Karnath et al. 2002Visual and auditory extinction have not been studied in the same patients...but delay is of the same time scale“It seems that the costs for information processing of contralesional events in extinction, induced by the bias of spatial attention towards the ipsilesional side, affect awareness of visual as well as auditory events to a similar degree.”
34 Seifritz et al. 2002Heschl’s Gyrus shows sustained response to sounds, surrounding regions respond to onset.
35 Key references (1) Signals and Perception 2002 Ch1 The mechanisms of hearing by AshmoreCh3 From cochlea to cortex by HackneyCh4 Imaging central auditory function by Palmer & Hall(2) Griffiths et al., Disorders of human complex sound processing Neurocase 5: , 1999(3) Human Cognitive Neuropsychology by Ellis & Young 1988Ch6 Recognising and understanding spoken words(4) Thinking in sound: The cognitive psychology of human audition Editors: McAdams & Bigand 1993Ch7 Auditory agnosia: A functional analysis by Peretz(5) Pavani et al., Auditory and multisensory aspects of visuospatial neglect. Trends in Cognitive Sciences 7: , 2003(6) Karnath et al., Impaired perception of temporal order in auditory extinction. Neuropsychologia 40:
36 Additional references (7) Review of functional organisation of the auditory cortexHall et al., Relationships between human auditory cortical structure and function. Audiology and Neuro-otology 8: 1-18, 2003(8) Case studies of auditory agnosiaReferences to many original papers can be found in (2) Griffiths et al., Disorders of human complex sound processing Neurocase 5: , 1999(9) A case of non-spatial auditory neglectCusack et al., Neglect between but not within auditory objects Journal of Cognitive Neuroscience 12:(10) Temporal order judgement deficits in visual neglectRorden et al., Visual extinction and prior entry: impaired perception of temporal order with intact motion perception after unilateral parietal damage. Neuropsychologia 35: