NEUROFISIOLOGIA E PATOLOGIA DEI PROCESSI COGNITIVI: Il Lobo Temporale What are we doing with our brains at this moment? • • • • • • • • • • • ’ Feeling your chair Squirming (moving) Watching Listening Remembering Paying attention Writing Feeling anxious Feeling hungry What happens when you ask a question? Learning What are we doing with our brains at this moment? • • • • • • • • • • • ’ Feeling your chair Squirming (moving) Watching (me, slides, mobile, the lady on the right) Listening Remembering Paying attention Writing Feeling anxious Feeling hungry What happens when you ask a question? Learning Lobi e solchi degli emisferi cerebrali Localizzazione dell’insula , visibile aprendo la scissura di Silvio ( solco laterale) Major lobes – hidden and visible insula and Sylvian fissure Insula is responsible for: ‘gut feelings’ like sense of nausea and disgust, interoception (feeling internal organs), emotional awarness. Sylvian fissure runs between parietal and temporal lobes horizontaly towards junction with occipital lobe. It contains supratemporal plane that hosts primary and secondary auditory cortex and part of Wernicke’s area for speech comprehension. 6 AREE CORTICALI MONOMODALI PRIMARIE: sensitive primarie, motoria primaria MONOMODALI SECONDARIE : sensitive secondarie motorie secondarie AREE ETEROMODALI : aree associative frontali, parietali, temporali Aree motorie, uditive, somatosensitive, visive , olfattive e associative negli emisferi di 3 diversi mammiferi. Si può notare l’enorme aumento nelll’uomo delle aree associative Le aree corticali motorie, uditive, somatosensitive, visive , olfattive sono aree primarie strettamente correlate con il mondo esterno ( sensitivo-motorie) Nell’uomo le aree associative consentono un elaborazione avanzata degli stimoli interni/esterni Vengono divise in unimodali ( es aree 18-19 corteccia visiva) e eteromodali o multimodali che sono correlate a funzione di integrazione AREE ASSOCIATIVE ETEROMODALI • CORTECCIA PREFRONTALE • CORTECCIA PARIETALE POSTEROINFERIORE • CORTECCIA TEMPORALE LATERALE • GIRO PARAIPPOCAMPALE • convergenza di tutti gli stimoli dalle aree unimodali • controllo sistema limbico e centri sottocorticali LOBO TEMPORALE giro temporale superiore giro temporale medio giro temporale inferiore giro fusiforme ( occipito-temporale) Funzioni: integrazione uditiva, linguaggio (porzione posteriore del giro superiore, area di Wernicke) funzioni visuospaziali, apprendimento e memoria External part of the temporal lobe on a dissected brain Central sulcus limit between frontal and parietal lobe and posterior limit of primary motor area Post central sulcus posterior limit of primary sensory area in parietal lobe With a blue arrow: gyrus With a red arrow: sulcus Terminal branch sylvian fissure Heschl’s gyrus external part of primary auditive area Gyrus supra marginalis ( parietal lobe) Wernicke’s area belongs to superior temporal gyrus Sylvian fissure- Superior temporal gyrus Superior temporal sulcus Middle temporal gyrus Inferior temporal gyrus Inferior temporal sulcus 6 MRI on horizontal plane of the temporal lobe Midbrain Temporal pole Amygdaloid nucleus Parahippocampal gyrus Arbitrary delineation between parahippocampal gyrus and lingual gyrus Lingual gyrus Ento rhinal sulcus Collateral sulcus The course of the collateral sulcus may be interrupted in some part of its course Collateral sulcus Fusiform gyrus Lingual gyrus and fusiform gyrus belong to the temporo occipital area 20 I sistemi di connessione Fasci di associazione tra le diverse aree corticali (DTI) A, arcuate (superior longitudinal) fasciculus; C, cingulum; EC, extreme capsule; F, fornix and stria terminalis ; H, hippocampus; IL, inferior longitudinal fasciculus; IO, inferior occipitofrontal fasciculus; P, putamen; SO, superior occipitofrontal fasciculus (combined with the subcallosal fasciculus adjacent to it); Th, thalamus; U, uncinate fasciculus. SINDROMI DA DISCONNESSIONE: INTERRUZIONE DELLE CONNESSIONI TRA AREE UNIMODALI/ ETEROMODAI DIVERSE ( es.: anomia per i colori) http://www.nfos.org/degree/opt41/3 Dalle connessioni tra le diverse aree cerebrali nascono reti specifiche. • Linguaggio • Attenzione spaziale • Funzioni esecutive e comportamento • Emozioni e memoria • Riconoscimento di facce e oggetti Localization of function in the nervous system: Functional networks 5 major brain systems subserving cognition and behavior Left perisylvian language network Parieto-frontal network for spatial attention Occipitotemporal network for object/face recognition Medial temporal/limbic network for learning & memory Prefrontal network for attention & comportment Lateralization of functions (approximate) • Left-hemisphere: – Sequential analysis • Analytical • Problem solving – Language and communication – Emotional functions • Recognizing emotions • Expressing emotions – Music Competence • Right-hemisphere: – Simultaneous analysis • Synthetic – Visual-Spatial skills • • • • Cognitive maps Personal space Facial recognition Drawing – Emotional functions • Perceiving emotions • Showing emotions – Music perception SUONI E LINGUAGGIO Onward to the Brain! • Organ of Corti transmits signals to the cochlear nerve • Medulla – Cochlear Nucleus to Superior Olivary Complex • Lateral Lemniscus fiber bundle carries information to the inferior colliculus • Proceeds to the Medial Geniculate Nucleus of the thalamus and on to the auditory cortex http://www.neuroreille.com/promenade/english/ptw/zoom1.htm Auditory cortex • In humans primary auditory cortex is located within Heschl’s gyrus. – Heschl’s gyrus corresponds to Brodmann’s area 41. • Another important region in auditory cortex is planum temporale located posterior to Heschl’s gyrus. – Planum temporale is much larger in the left hemisphere (up to 10 times) in right handed individuals. – It plays important role in language understanding. • Posterior to planum temporale is Broadmann area 22 that Carl Wernicke associated with speech comprehension (Wernicke area). 22 Speech perception Brain response to: Words Pseudowords Reversed speech Binder and colleagues (1997) studied activation of brain areas to words, reverse speech and pseudowords and found that Heschl’s gyrus and the planum temporale were activated similarly for all stimuli. This supports the notion of hierarchical processing of sounds with Heschl’s gyrus representing early sensory analysis. Speech signals activated larger portion of auditory cortex than non-speech sounds in posterior superior temporal gyrus and superior temporal sulcus, but there was no difference in activation between words, pseudowords and reversed speech. The conclusion is that these regions do not reflect semantic processing of the words but reflect phonological processing of the speech sounds. 23 Functional mapping of auditory processing The planum temporale (PT) location close to Wernicke’s area for speech comprehension, points towards its role as the site for auditory speech and language processing. However neuroimaging studies of PT provide evidence that functional role of PT is not limited to speech. PT is a hub for auditory scene analysis, decoding sensory inputs and comparing them to memories and past experiences. PT further directs cortical processing to decode spatial location and auditory object identification. Planum temporale and its major associations: lateral superior temporal gyrus (STG), superior temporal sulcus (STS), middle temporal gyrus (MTG), parieto-temporal operculum (PTO), inferior parietal lobe (IPL). 24 Functional mapping of auditory processing Auditory objects are categorized into human voices, musical instruments, animal sounds, etc. Auditory objects are learned over our lifetime, and associations are stored in the memory. Auditory areas in superior temporal cortex are activated both by recognized and unrecognized sounds. Recognized sounds also activate superior temporal sulcus and middle temporal gyrus (MTG). Fig. (c) shows difference between Activations for recognized sounds and unrecognized sounds 25 Functional mapping of auditory processing Binder and colleagues propose that middle temporal gyrus (MTG) is the region that associates sounds and images. This is in agreement with case studies of patients who suffered from auditory agnosia (inability to recognize sounds). Research results showed that auditory object perception is a complex process and involves multiple brain regions in both hemispheres. Brain activities in auditory processing – cross sections at different depth 26 Function source : Palmer & Hall, 2002 • Numerous bilateral regions are frequency-dependent • Overlapping regions are sensitive to intensity and to the temporal changes in sound • One 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 important 27 L L H H H L L Right hemisphere Slow-rate temporal pattern in sound Auditory Cortex • Tonotopic Organization – Different frequencies of sound are mapped to different regions of the auditory cortex – Extends to the level of the cochlea Zhou, X. and M. M. Merzenich (2007). "Intensive training in adults refines A1 representations degraded in an early postnatal critical period." Proceedings of the National Academy of Sciences 104(40): 15935-15940. • 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 TTG • Medium intensity (82 dB) sounds activated intermediate area (green). (NeuroImage 2002;17: 710) 29 Auditory agnosia • A deficit in recognition Perception Auditory input Acoustical analysis Representations “Apperceptive agnosia” “Associative agnosia” Auditory agnosia is of this type 30 Recognition • sordità verbale pura ( lesione bitemporale o sinistra, area 22) • agnosia acustica ( area acustica secondaria) What is Language? • Grammar – Phonetics, morphology, syntax, semantics • • • • • Symbol usage Ability to represent real-world situations Ability to articulate something new Intention to communicate Duality, productivity, arbitrariness, interchangeability, specialization, displacement, and cultural transmission (Linden 1974) “An infinitely open system of communication” Rumbaugh, 1977 Schematic representation of the components that are involved in spoken and written language comprehension. Input can enter via either auditory (spoken word) or visual (written word) modalities. The flow of info is bottom up, from Perceptual identification to “higher-level” word and lemma activation. Interactive models of language Understanding would predict top-down influence to play a role as well. W. W. Norton Outline of the theory of speech production developed by William Levelt (1999) Adapted from Levelt, W.J.M., The Architecture of Normal Spoken Language Use, in Blanken, G., Dittman, J., Grimm, H., Marshall, J.C., and Wallesh, C-W. (Eds.), Linguistic Disorders and Pathologies: An International Handbook. Berlin: Walter de Gruyter, 1993 Speech Production Brain areas involved in Language Wernicke-Geschwind Model 1. Repeating a spoken word • Arcuate fasciculus is the bridge from the Wernicke’s area to the Broca’s area Wernicke-Geschwind Model 2. Repeating a written word • Angular gyrus is the gateway from visual cortex to Wernicke’s area • This is an oversimplification of the issue: – not all patients show such predicted behavior (Howard, 1997) Functional neuroimaging of the language network One to many, many to one CJ Price, J Anat 2002 Language function: Using neuroimaging to test hypotheses CJ Price, J Anat 2002 Brain Imaging fMRI:Record during language tasks – Activated brain areas consistent with temporal and parietal language areas – More activity than expected in nondominant hemisphere Generate words from a category Silently repeat a heard sentence Listen to a story Slide 42 Neuroscience: Exploring the Brain, 3rd Ed, Bear, Connors, and Paradiso Copyright © 2007 Lippincott Williams & Wilkins A functional MRI protocol for localizing language comprehension in the human brain Gary W. Thickbrooma ,*, Michelle L. Byrnesa, David J. Blackerb, Ian T. Morrisc, a,b,d Frank L. Mastaglia Brain Research Protocols 10 (2003) 175–180 TONE DECISION TASK Semantic decision PET by Damasio’s • Different areas of left hemisphere (other than Broca’s and Wernicke’s regions) are used to name (1) tools, (2) animals, and (3) persons • Stroke studies support this claim • Three different regions in temporal lobe are used • ERP studies support that word meaning are on temporal lobe (may originate from Wernicke’s area): – “the man started the car engine and stepped on the pancake” – Takes longer to process if grammar is involved LETTURA Neural systems for reading • Converging evidence indicates three important systems in reading, all primarily in the left hemisphere– Some right hemisphere activation now implicated; • These include an anterior system and two posterior systems: 1) anterior system in the left inferior frontal region; 2) dorsal parietotemporal system involving angular gyrus, supramarginal gyrus and posterior portions of the superior temporal gyrus; 3) ventral occipitotemporal system involving portions of the middle temporal gyrus and middle occipital gyrus 50 Dorsal (Green) and Ventral Pathways (Purple) Written language neural pathway • Visual input transmitted from lateral geniculate to primary cortex in striate areas and secondary extrastriate cortex. • From here 2 streams– Ventral (what): unimodal visual area of fusiform gyrus (may contain ortho reps of words) – Dorsal (where): superior parietal lobule for spatial aspects of reading. 52 Heteromodal areas • Wernicke’s including angular gyrus, supramarginal gyrus – Likely responsible for integration of written & spoken word forms. – Wernicke’s is massively connected to inferior temporal category specific areas for faces animals, tools – Also with frontal areas for overt speech production (Broca’s), and reciprocal connections for memory and manipulating verbal information. 53 Confirmed two systems for reading • word analysis – operating on individual units of words such as phonemes, requiring attentional resources and processing relatively slowly • Parietotemporal area and • visual word attention – an obligatory system that does not require attention and processes very rapidly, on the order of 150 msec after a word is read; Price et al 1996. • Occipitotemporal area • visual word form area appears to respond preferentially to rapidly presented stimuli (Price et al 1996) and is engaged even when the word has not been consciously perceived (Dehaene et al 2001). 54 Due sistemi corticali: dorsale e ventrale Cellular responses in ventral stream • V2-V4 especially orientation & color sensitive – V4 contains color sensitive cells – some have complex preferred stimuli • Inferior temporal cortex (IT) is form sensitive – big receptive fields (up to entire visual field) – face selective cells in some regions – others respond best to complex, 3D stimuli (Tanaka) www.cnbc.cmu.edu/~mgilzen/85219/slides/Week8a.ppt Effects of lesions to ventral stream • Animal studies – Damage to P pathway inputs abolishes color perception – Damage to IT impairs discrimination between objects and identification • Human patients – Achromatopsia • Loss of color perception • Associated with damage to lingual and fusiform gyri – Agnosia • Apperceptive • Associative • Prosopagnosia www.cnbc.cmu.edu/~mgilzen/85219/slides/Week8a.ppt PERCEZIONE VISIVA Visual processing streams: Confirmation of hypotheses using neuroimaging Ungerleider LG, PNAS 1998 Visual processing: Attention influences which stream is used Ungerleider LG, PNAS 1998 Encoding & recall of categoryspecific information Faces: Fusiform gyrus Places: Parahippocampal gyrus Encoding of category-specific information activates relevant areas of cortex Polyn SM et al., Science, 2005 Visual object recognition: Faces & places Kanwisher N, Science, 2006 Fusiform Face Area (FFA) • functional brain imaging investigations of the normal human brain show that a region in the fusiform gyrus is not only activated when subjects view faces, but is activated twice as strongly for faces as for a wide range of nonface stimuli (Kanwisher et al., 1997) L’altra faccia della luna IL LOBO LIMBICO TEMPORALE Functions of The Limbic System LOBO LIMBICO giro del cingolo giro paraippocampale giro paraterminale ( lamina terminalis) giro subcallosale ippocampo, paraippocampo, uncus Emozioni, apprendimento , memoria Medial Temporal Lobe (MTL) • Hippocampus • Connected areas – Entorhinal cortex – Perirhinal cortex – Parahippocampal cortex Bird & Burgess, Nature Reviews Neuroscience 9, 182-1 Function of MTL • Principally concerned with memory • Operates with neocortex to establish and maintain long-term memory • Ultimately, through a process of consolidation, becomes independent of long-term memory MTL Simons & Spiers (2003) Nature Reviews Neuroscience 4; 637 STRUTTURA DELL’IPPOCAMPO A: ricostruzione dell’ippocampo e del fornice. B : struttura del giro dentato e del corno d’Ammone C, sezione coronale del giro dentato, ippocampo e subiculum D, diagramma delle cellule e fibre dell’ippocampo Le fibre efferenti dall’ippocampo si originano da cellule piramidali . La maggior parte delle efferenze originano dal subiculum. Il flusso di informazioni proviene maggiormente dalla corteccia entorinale > giro dentato> cellule piramidali dell’ippocampo e subiculum> fornice Nel giro dentato dono presenti cellule staminali. (B, modified from Duvernoy HM: The human hippocampus: functional anatomy, vascularization and serial sections with MRI, ed 3, Berlin, 2005, Springer-Verlag. C, modified from Nolte J, Angevine JB Jr: The human brain in photographs and diagrams, ed 3, St. Louis, 2007, Mosby.) • CA1 > CA4 = zone del corno d’Ammone ( ippocampo = corno d’Ammone + giro dentato) • Paraippocampo = subiculum + corteccia entorinale Settore CA1 del C.Ammone = settore di SOMMER : compostoa da grandi cellule piramidali, Con metabolismo molto attivo, alta densita di recettori NMDA, estrema sensibillità all’ipossia, Marcato potenziamento post-sinaptico ( 2 ore per singolo stimolo in arrivo) Memoria sinaptica Neuroplasticità Crisi epilettiche Connessioni anatomiche dell’ippocampo Input e out dell’ippocampo: è una struttura nervosa della regione temporale mediale Gli input all’ippocampo: arrivano tramite stazioni di ritrasmissione nelle corteccie peririnali, paraippocampali ed entorinali Le uscite dall’ippocampo: seguono il percorso opposto. Gli inputoutput sotto-corticali dell’ippocampo viaggiano nel fornice AFFERENZE DELL’IPPOCAMPO: • corteccia entorinale ( bulbo olfattorio) • amigdala • nuclei settali • ippocampo controlaterale • locus coerulus . (Modified from an illustration in Warwick R, Williams PL: Gray's anatomy, Br ed 35, Philadelphia, 1973, WB Saunders.) EFFERENZE DELL’IPPOCAMPO • Fornice ( attraverso il fornice le informazioni provenienti dall’ippocampo raggiungono i corpi mammilari,il nucleo anteriore e mediodorsale del talamo, il cingolo) Papez Circuit (memory) Fornix Mammillary bodies Other hypothalamic nuclei Septal nuclei Substantia innominata (Basal nucleus of Meynert) Hippocampal Formation (hippocampus and dentate gyrus) Parahippocampal Gyrus Neocortex Mammillothalamic tract Anterior Thalamic nuclear group Cortex of Cingulate Gyrus The Papez circuit. The shortcut from the hippocampus directly to the anterior thalamic nucleus, not part of the circuit as originally proposed, is indicated by a dashed line. A, anterior thalamic nucleus; CA, hippocampus proper; D, dentate gyrus; MB, mammillary body. (Modified from an illustration in Warwick R, Williams PL: Gray's anatomy, Br ed 35, Philadelphia, 1973, WB Saunders.) IPPOCAMPO E MEMORIA Nel 1957 un paziente di nome H.M. fu trattato chirurgicamente per epilessia resistente alla terapia farmacologica con asportazione bilaterale dell’ippocampo e dell’amigdala. HM soffri da quel momento di un grave disturbo della memoria episodica Dopo molti anni dall’intervento il paziente fu sottoposto a RM encefalo. Il suo caso dimostrò in modo conclusivo il ruolo fondamentale della parte mediale del lobo temporale nella memoria HM AMIGDALA L’amigdala è un insieme di circa 12 nuclei situati in prossimità dell’uncus, vicino alla parte anteriore dell’ippocampo, e accanto al corno temporale del ventricolo laterale . Origina dalla corteccia periamigdaloidea , che forma parte della superficie del’uncus. I nuclei dell’amigdala vengono classificati in 3 principali gruppi : mediale, centrale e basolaterale, ognuno con diverse funzioni. Nucleo mediale: è connesso con il sistema olfattorio ed è piuttosto piccolo negli esseri umani. Nucleo centrale : collegato all’ipotalamo e alla sostanza grigia periacqueduttale ha funzioni emozionali Nucleo basolaterale: è il nucleo più grande negli esseri umani, contiene cellule piramidali ed è strettamente collegato agli altri nuclei Suddivisione dell’amigdala nei principali nuclei (basolaterale, centrale e mediale) con le principali connessioni. ( PH, parahippocampal gyrus) Amygdala Connections Cerebral cortex Olfactory system Thalamus Brainstem reticular formation Stria terminalis Hypothalamus AMYGDALA Ventral Amygdalofugal fibers Amygdala Inputs Olfactory System Temporal Lobe (associated with visual, auditory, tactile senses) AMYGDALA Corticomedial Nuclear Group Basolateral Nuclear Group Central Nucleus Brainstem (viscerosensory relay Nuclei: solitary nucleus and parbrachial nucleus) Ventral Amygdalofugal Fibers Maggiori input ai nuclei basolaterale(blu), centrale(rosso) e mediale (verde) dell’amigdala. Sono evidenziati gli input dalle aree visive , ma simili proiezioni esistono dalle principali aree unimodali e dalle altre aree limbiche. B, brainstem (periaqueductal gray, parabrachial nuclei, other nuclei); Hy, hypothalamus; S, septal nuclei; T, thalamus (multiple nuclei). (Modified from Warwick R, Williams PL: Gray's anatomy, Br ed 35, Philadelphia, 1973, WB Saunders.) Principali efferenze dai nuclei dell’amigdala ( blu= basolaterale;rosso=centrale; verde= mediale) (1) stria terminalis: verso i nuclei del setto e l’ipotalamo (2) Via amigdalofugale e Via amigdalofugale ventrale ( ipotalamo e talamo MD) (3) proiezioni diffuse alla corteccia frontale ventromediale , corteccia insulare, striato ventrale, aree olfattorie , tronco encefalico . (4) connessioni dirette con l’ippocampo e il lobo temporale. (Modified from Warwick R, Williams PL: Gray's anatomy, Br ed 35, Philadelphia, 1973, WB Saunders.) Riepilogando: L’amigdala riceve una gran quantità di input : 1. Molti di questi sono semplici e famigliari, riguardando stimoli visivi, suoni, stimoli tattili, odori e. Gli stimoli olfattivi raggiungono direttamente il nucleo mediale , dalla corteccia e bulbo olfattorio ( tratto olfattorio). Gli altri tipi di stimoli raggiungono il nucleo basolaterale dal talamo e dalle cortecce unimodali visiva, uditiva, gustativa e somatosensitiva. 2. Un altro tipo di stimolo arriva all’amigdala dall’insula, dal cingolo e dalla corteccia orbitofrontale ( via amigdalofugale) 3.Infine l’amigdala riceve input viscerali provenienti dall’ipotalamo ( stria terminalis) e dal tronco encefalico ( grigio periacqueduttale , n.parabrachiale) Le efferenze dall’amigdala raggiungono le stesse aree da cui l’amigdala riceve degli input Limbic System and Basal Nuclei Anterior Cingulate Gyrus Orbitofrontal Areas (10, 11) Medial and lateral temporal lobe Hippocampus Amygdala Entorhinal cortex (24) Ventral Pallidum Medial Globus Pallidus Pars Reticularis (Substantia nigra) Ventral Striatum (nucleus accumbens) Caudate Nucleus (head) Ventral Anterior Nucleus Dorsomedial Nucleus Functions of the Amygdala • Relate environmental stimuli to coordinated behavioral autonomic and endocrine and motor responses seen in species-preservation. • Responses include: Feeding and drinking Agnostic (fighting) behavior Mating and maternal care Responses to physical or emotional stresses. La connessione limbica tra amigdala e gangli della base influenza le decisioni riguardo al movimento e, più in generale, è necessaria per l’associazione stimolo-ricompensa. Qualsiasi cosa possa apparire piacevole determina un aumento di rilascio di dopamina nello striato che si traduce successivamente in un’azione; avviene il contrario per ciò che consideriamo spiacevole Neuro-anatomy of hedonic regulation of food intake; reward (or pleasure) seeking areas of the brain control orexigenic neurons of LHA L’amigdala, attraverso le sue connessioni , in particolare con la corteccia frontoventromediale e l’ipotalamo, svolge un ruolo fondamentale nella vita emotiva. Quando l’amigdala di un animale viene stimolata l’animale si ferma e rimane attento: a ciò può far seguito una risposta di difesa, rabbia, aggressione, o di fuga La stimolazione dell’amigdala nell’uomo evoca più spesso paura accompagnata dalle reazioni vegetative corrispondenti . La distruzione bilaterale dell’amigdala determina un assenza di paura e di aggressività con mancanza di reazioni vegetative . L’assenza congenita dell’amigdala impedisce l’apprendimento di reazioni comportamentali complesse in risposta a determinate situazioni LA PATOLOGIA Symptoms of Temporal-Lobe Lesions Pathologies (lesions) • Voracious appetite • Increased (perverse) sexual activity • Docility: Loss of normal fear/anger response • Memory loss: Damage to hippocampus portion: Cells undergoing calcium-induced changes associated with memory Temporal lobectomy • Brown and Schäfer (1888) reported behaviour of monkey ‘Tame one’ after bilateral temporal lobectomy. • Preop: wild, fierce • Postop: – Does not retaliate or escape if slapped, tame – Poor memory and intelligence – Evidence of hearing and seeing, but ‘no longer clearly understands meaning of sights, sounds.’ – Does not select raisins from other food in dish: does not seem to visually recognize 97 items. Kluver-Bucy Syndrome: • Results from bilateral destruction of amygdala. • Characteristics: Increase in sexual activity. Compulsive tendency to place objects in mouth. Decreased emotionality. Changes in eating behavior. Visual agnosia. Delusional Misidentification Syndromes • 3 DMS: 1. Pick [1903] “reduplicative paramnesia” • Misidentifies familiar places as replica 2. Capgras Syndrome [1923] • Familiar people described as doppelgangers – visual but not emotional recognition 3. Frégoli Syndrome [1927] • • DMS are rare – – 99 Person misidentified as someone else with totally different appearance. Rare enough to be of little clinical importance Yet, may still reveal how emotions are processed Hirstein and Ramachandran [1997] • H&R postulate that DMS is caused by disconnection between visual recognition system and emotional system. • E.G. Capgras syndrome due to disconnection between fusiform gyrus [face area] and amygdala [limbic system] 100 Hudson and Grace • 71 women suffered lesion to anterior fusiform gyrus (between face area and amygdala) – Frégoli Syndrome • Identified husband as elder sister (who had died 3 years previously) • Only visual misidentification (fine on phone) • Home was ‘replica’ would pack bags to return to ‘real’ home. • Support for H&R 101 Pain asymbolia • Patient’s report they can feel pain, but it no longer hurts. • Ramachandran (1998): speculates disconnection of insula from cingulate (part of limbic system) – Insula identifies pain – Cingulate does not receive signal, so discounts threat 102 I DISTURBI ACUSTICI E DI LINGUAGGIO Auditory agnosia source : Griffiths et al. 1999 • Normal brainstem processing • Midbrain impairment questionable • Cortical deficit in perception - Preserved hearing (pure tones) - Disordered perception of certain sounds : Speech - word deafness Music - amusia Environmental sounds - environmental sound agnosia 104 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 105 Word deafness source : Ellis & Young, 1988 • Associated symptoms - Some hearing loss (> 20 dB HL) - Production (Broca’s) aphasia - Perception of melody - Perception of environmental sounds • Lesion 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) 106 A case of amusia source : Peretz, 1993 Patient CN • Symptoms - 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 sounds • Lesion site - Bilateral temporal lobe damage - When unilateral, it’s more often the right hemisphere 107 Amusia source : Peretz, 1993 • Dissociation 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 108 Environmental sound agnosia source : Griffiths et al. 1999 • Deficit rarely occurs in isolation Environmental sounds contain fewer changes in acoustic structure over time than an equivalent length segment of speech or music 109 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) 110 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 tested • All 3 types of sound contain a mixture of acoustic features • Deficit in an intermediate level of analysis, which is rarely tested - Analysing the spectro-temporal pattern in sound 111 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 side Failure to detect contralesional sound, when presented concurrently Poor allocation of attention to sounds separated in time 112 Auditory neglect Lesion site – usually right hemisphere - inferior parietal lobe - superior temporal gyrus - temporo-parietal junction 113 source : Pavani et al., 2003 Quadro riassuntivo Summary: Correlations of symptoms with areas of lesion Aphasic Syndrome Area of Damage Broca’s Broca’s area Wernicke’s Wernicke’s area Conduction SMG, Insula, Arcuate fasciculus Transcortical motor Transcortical sensory Areas anterior and/or superior to Broca’s area Areas posterior and/or superior to Wernickes a. Cf. H. Damasio 1998: 43-44 Two different patients with anomia Inability to retrieve words for unique entities Deficit in retrieval of animal names (Left temporal lobectomy) (Damage from stroke) Two more patients with anomia Deficit in retrieval of words for man-made manipulable objects (Damage from stroke) Severe deficit in retrieval of words for concrete entities (Herpes simplex encephalitis) I DISTURBI AGNOSICI Cerebral Achromatopsia Ishyhara plates Color Agnosia • What causes it? – Damage to the extrastriate visual cortex, specifically the V4 area. – The visual area 4 (V4) is cut off from sending information to the lingual gyrus, fusiform gyrus and inferior temporal gyrus. – Without the ability to send that information to the “What” pathway of the temporal lobe, the color cannot be recognized. Cerebral Achromatopsia Usually caused by bilateral damage to V4 (lingual and fusiform gyri (occipitotemporal junction)) characterized by an inability to identify or discriminate colour Usually full field deficit but hemiachromatopsia possible if damage is unilateral Still able to perceive form and motion - dissociation with akinetopsia and visual form agnosias Achromatopsia is not: a) b) c) d) due to peripheral damage (e.g. retina) due to primary visual area damage colour agnosia: disorder of colour categorization colour anomia: disorder of colour naming Which part of V4? Damasio et al., (1989b) in Heilman and Rothi (1993) 42 patients. achromatopsia associated with lesions below calcarine sulcus that damaged middle third of lingual gyrus, but not fusiform gyrus Calcarine sulcus Lingual Gyrus Fusiform Gyrus Prosopagnosia • Special nature of face processing originates from reports of patients who are reportedly unable to recognize familiar faces while maintaining the ability to recognize objects. – This impairment is often accompanied by focal damage to the ventral occipitotemporal and temporal cortices. – Reported to process faces similar to objects (sometimes absence of inversion effects). Prosopagnosia Why just faces? What about objects? Fusiform gyrus responds to faces Parahippocampal gyrus responds to inanimate objects Double Dissociation- The areas for recognizing faces and inanimate objects are separate therefore agnosia for objects and prosopagnosia do not occur together Face Processing: Cognitive and Neural Components Bruce & Young (1986) Haxby et al., (2000 Importance of Facial Configuration • the importance of the overall configuration of the face can help us understand why face recognition can be remarkably robust despite a variety of natural (change in expression, orientation etc) as well as unnatural (cartoons) transformations in faces. Prosopagnosia • Marotta, Genovese, & Behrmann (2001) – fMRI of prosopagnosic patient did not show normal activation of fusiform. – Did show left hemisphere posterior fusiform activation, suggesting faces are being processed featurally. Fig. 4-18, p. 83 Prosopagnosia Brain region involved George et al. (1999). fMRI study of positive and reverse contrast faces. Bilateral fusiform gyri response to faces Right fusiform gyrus only when face became familiar (Note contrast to Alexia) Image of Left Fusiform Gyrus (Visual Word Form Area -VWFA) and Right Fusiform Gyrus (Weiner et al, 2004.) The left fusiform gyrus ( referred to as the visual word form area) is responsible for word recognition. The right fusiform gyrus is (referred to as the fusiform face area) responsible for facial recognition. What Causes Prosopagnosia? • Damage to occipitaltemporal regions of the brain – Specifically the fusiform gyrus of the interior temporal cortex – The cause of damage can be from head injury, degenerative diseases (ex. Alzheimer’s and Parkinson’s disease), right temporal lobe atrophy, encephalitis, or strokes (ex. Posterior cerebral artery stroke or transient ischemic attack). • There is also a genetic form of prosopagnosia that can be passed down from a parent to child. Prosopagnosia can also be the result of a developmental disorder. • Evolutionarily facial recognition is important – There must be an adaptive benefit for facial recognition. Newborns show great preference for human faces despite their poor visual acuity. Newborns use eye contact and facial expressions to engage caretakers to take care of their needs. Socially facial recognition became essential to survival because it provides self-identity, identity for group members, and identity for non-group member (could be an enemy?). LA MEMORIA Memory and Forgetting Classificazione dei tipi di memoria. Memoria Esplicita (dichiarativa) Episodica Semantica Implicita Procedurale Emotiva Vegetativa (condizionamento) • cos’è una bicicletta • ieri sono andato in bicicletta • andare in bicicletta dopo 10 anni • ho paura della bicicletta ( non so perché) memoria semantica memoria episodica memoria procedurale memoria emotiva Brain regions in Learning and Memory Patient H.M. • Bilateral temporal lobe resection for treatment of epilepsy. • Included removal of hippocampus and amygdala from both sides. • Various etiologies lead to symptoms like H.M.'S, including stroke and herpes Case of Patient H. M. • Age 9, knocked over by a bicycle rider, sustained brain damage • Age 16, suffered bilateral temporal lobe seizures which became uncontrollable – Unable to work and lead a normal life • Age 27, underwent bilateral removal of hippocampal formation, amygdala, parts of multimodal association areas of temporal cortex (1953) Consequences of Psychosurgery for H.M. • Positive - seizures better controlled - IQ unaffected; bright - good long term memory for events before the surgery - good command of language including vocabulary - remembered his name and job he held • Negative • suffered anterograde amnesia unable to transfer new short-term memory into long term memory – unable to retain for more than a minute new people, places or objects – unable to recognize people he met during surgery including his neurosurgeons – took a year to learn his way around a new house - suffered retrograde amnesia for information acquired a few years before surgery Patient HM • Revealed declarative/ nondeclarative distinction – Declarative memories (explicit memories) involve conscious recollection of events and information. • H.M. Lost this ability. – Nondeclarative memories (implicit memories) involve ability to acquire and perform new behaviors or associations. • H.M. Retained this ability. – could perform mirror-tracing after training but could not remember doing the task before Semantic Dementia (Snowden 1989) • Semantic memory (Warrington 1975): • Term applied to the component of long-term memory which contains the permanent representation of our knowledge about things in the world: facts, concepts and words • Culturally shared, acquired early in life. Semantic Dementia (Snowden 1989) • Affects fundamental aspects of language, memory and object recognition. Semantic Dementia • Progressive anomia, not an aphasia, but a loss of semantic memory. • Impaired: naming, word comprehension, object recognition and understanding of concepts. • characterized by preserved fluency and impaired language comprehension: “phonologically and syntactically correct” Assessement • Category fluency • Generation of definitions – Lion: ” it has little legs and big ears, they sleep a lot, see them in shops” • Word-picture matching • Famous faces test • Normal episodic memory, normal visuospatial skills • Nature of error Semantic-type naming errors:initially within-category, “elephant” for hippopotamus, then superordinate “dog” for everything, then “animal”… • Profound and complete anomia • Circumlocutions and semantic paraphasias – semantic paraphasias, in which the wrong word is produced, one that is usually related to the target (eg, "pliers" for "hammer"). • In semantic dementia the most contextfree levels of knowledge (constituting traditional notions of semantic memory) are most compromised. • In contrast, patients may retain knowledge tied to specific experiences or routines SD and memory • Can relate details ( in a rather anomic fashion) of recent events, but there is impaired recall of distant life events. SD • In most cases, neuroradiological studies reveal selective damage to the inferolateral temporal gyri(inferior and middle) of one or both temporal lobes, with sparing of the hippocampi, parahippocampal gyri, and subiculum. • Note: AD: inferior and middle temporal gyri Semantic dementia patient with severe focal atrophy of the left temporal lobe see arrow, right-hand side of MRI scan) involving the pole, inferior, and middle temporal gyri with relative sparing of the hippocampal complex (H) and of the superior temporal gyrus. fine Grazie per l’attenzione