NEUROANATOMY OF LANGUAGE 1 SEPT 13, 2013 – DAY 8 Brain & Language LING NSCI Harry Howard Tulane University

Course organization The syllabus, these slides and my recordings are available at If you want to learn more about EEG and neurolinguistics, you are welcome to participate in my lab. This is also a good way to get started on an honor's thesis. The grades are posted to Blackboard. 9/13/13Brain & Language - Harry Howard - Tulane University 2

Review Syntax 1. Kiss Mary, I would never do. 2. [ VP kiss Mary] I would never do 3. *Kiss, I would never do Mary. 4. *[ VP kiss] I would never do [ VP Mary] By the way, how do you know which ones are bad? Because you are an expert in the grammar of your native language. 9/13/13Brain & Language - Harry Howard - Tulane University 3

NEUROANATOMY OF LANGUAGE Ingram §3 9/13/13Brain & Language - Harry Howard - Tulane University 4

9/13/13Brain & Language - Harry Howard - Tulane University 5 Short history of research DateEvent 1836Abercrombie? 1836 Marc Dax claimed that the LH of right-handers has “ memory for words ” 1861 Paul Broca claimed that the LH of right-handers has “ faculty of articulate speech ” 1874 Karl Wernicke discovered that damage to a certain area could cause receptive aphasia. John Hughlings Jackson claimed that the LH is responsible for language, while the RH is responsible for visual cognition (recognition, discrimination, recall). WWI- II Many observations of the cognitive results of head injuries end WWII Juhn A. Wada developed test for cerebral dominance for speech by injecting an anesthetic into the right or left internal carotid artery 1950s Penfield & Wilder use cortical stimulation to map the cortex > treat epilepsy, discover the motor-sensory homunculus 1960sCorpus callosotomy (commissurotomy) > split-brain patients 1970sHemifield tachistoscopy, dichotic listening > laterality research 1980sNoninvasive imaging techniques

9/13/13Brain & Language - Harry Howard - Tulane University 6 Overview of methodologies Lesions Wada test Craniotomy & cortical stimulation Corpus callosotomy & split-brain patients Hemifield tachistoscopy Dichotic listening Imaging: C(A)T, PET, (f)MRI Electromagnetic: EEG, MEG - not today Transcranial magnetic stimulation (TMS)

9/13/13Brain & Language - Harry Howard - Tulane University 7 Lesions A lesion is a non- specific term referring to abnormal tissue in the body. It can be caused by any disease process including trauma (physical, chemical, electrical), infection, neoplasm, metabolic and autoimmune.

9/13/13Brain & Language - Harry Howard - Tulane University 8 Wada test The Wada test (named for a neurologist, Juhn A. Wada) consists of behavioral testing after the injection of an anesthetic (such as sodium amobarbital or sodium methohexital) into the right or left internal carotid artery. Depending on how the injection is made (and the quantity), there is a certain amount of time during which the activities of one of the cerebral hemispheres are suspended, so the abilities subserved by the other hemisphere can be tested in isolation.

9/13/13Brain & Language - Harry Howard - Tulane University 9 Craniotomy & cortical stimulation A craniotomy is a surgical operation in which part of the skull, called a skull flap, is removed in order to access the brain. Craniotomies are necessary for many types of surgery; they are also widely used in neuroscience in techniques such as extracellular recording, brain imaging, and manipulations such as electrical stimulation and chemical titration. Human craniotomy is usually performed under general anesthesia but can be also done with the patient awake using a local anaesthetic and generally does not involve significant discomfort for the patient.

9/13/13Brain & Language - Harry Howard - Tulane University 10 Corpus callosotomy (split-brain patients) “Split-brain” is a lay term to describe the result of severing the corpus callosum to some degree. The surgical operation to produce this condition is called corpus callosotomy. It is rarely performed, usually only in the case of epilepsy, in order to mitigate the risk of accidental physical injury by reducing the severity and violence of epileptic seizures.

9/13/13Brain & Language - Harry Howard - Tulane University 11 Corpus callosotomy (split-brain patients), cont. A patient with a split brain, when shown an image in his or her left visual field (the left half of what each eye sees), will be unable to name what he or she has seen. This is because the speech control center is in the left side of the brain in most people and the image from the left visual field is sent only to the right side of the brain. Since the two sides of the brain cannot communicate, the patient can't name what he or she is seeing. The person can, however, pick up a corresponding object with their left hand, since that hand is controlled by the right side of their brain.

Divided visual-field (hemifield) tachistoscopy A tachistoscope is a device that displays an image for a specific amount of time. It can be used to increase recognition speed, to show something too fast to be consciously recognized, or to test which elements of an image are memorable. 9/13/13Brain & Language - Harry Howard - Tulane University 12

Dichotic listening Dichotic listening is a procedure commonly used for investigating selective attention in the auditory domain. Two messages are presented to both the left and right ears (one message to each ear), normally using a set of headphones. Normally, participants are asked to pay attention to either one, or both (divided attention condition) of the messages and may later be asked about the content of both. 9/13/13Brain & Language - Harry Howard - Tulane University 13

The cocktail party effect Dichotic listening was developed to mimic processing demands in the natural world, where sensory overload is common. Consider the cocktail party or, more appropriate for today, the wine-tasting party. We may attempt to speak with one individual, but the speaker's voice is intermixed with a multitude of incoming auditory signals: conversations going on about us, music from the compact disc player, the clatter of plates being filled at the buffet table, the children watching a video in the next room. Despite this cacophony of sound, we are quite proficient at focusing on the relevant signal—the words being spoken by our conversational partner. 9/13/13Brain & Language - Harry Howard - Tulane University 14

9/13/13Brain & Language - Harry Howard - Tulane University 15 The right-ear advantage In the seminal study of Kimura (1961), the stimuli were digits, presented so that one digit was heard in the left ear at the same time as a second digit was heard in the right ear. Kimura found that people were much more likely to report having heard the stimuli presented to the right ear, an effect dubbed the right- ear advantage. Kimura’s usage of dichotic listening to confirm the lateralization of language to the left hemisphere in normal subjects was soon substantiated by studies with lesioned subjects. Kimura (1961b) showed that patients with left temporal lobe lesions performed worse at the task than did patients with right temporal lobe lesions. In addition, split-brain patients showed a considerable right ear advantage in a study using words as stimuli. They succeeded in recognizing words presented to the right ear, but performed no better than chance for words presented to the left ear.

Computerized (Axial) Tomography (CT/CAT) Computed tomography (CT), originally known as computed axial tomography (CAT or CT scan), employs tomography (digital geometry processing) to generate a 3D image of the internals of an object from a large series of two- dimensional X-ray images taken around a single axis of rotation. 9/13/13Brain & Language - Harry Howard - Tulane University 16

9/13/13Brain & Language - Harry Howard - Tulane University 17 Positron Emission Tomography (PET) To produce a PET scan, a patient is administered a solution of a metabolically-active substance, such as glucose, tagged with a positron-emitting isotope. The substance eventually makes its way to the brain and concentrates in areas of high metabolism and blood flow, which are presumably triggered by increased neural activity. The positrons emitted by the isotopes are collected by detectors arrayed around the patients’ body and converted into signals which are amplified and sent to a computer for construction of an image.

9/13/13Brain & Language - Harry Howard - Tulane University 18 PET vs CT PET differs from CT in that it uses the body’s basic biochemistry to produce images. The positron-emitting isotope is chosen from elements that the body already uses, such as carbon, nitrogen, oxygen, and fluorine. By relying on normal metabolism, PET is able to show a biochemical change even in diseases such as Alzheimer’s in which there is no gross structural abnormality.

9/13/13Brain & Language - Harry Howard - Tulane University 19 Magnetic Resonance Imaging (MRI) In 1977, a team lead by Raymond Damadian produced the first image of the interior of the human body with a prototype device using nuclear magnetic resonance. Damadian’s device uses liquid helium to supercool magnets in the walls of a cylindrical chamber. A subject is introduced into the chamber and so exposed to a powerful magnetic field. This magnetic field has a particular effect on the nuclei of hydrogen atoms in the water which all cells contain that forms the basis of the imaging technique.

9/13/13Brain & Language - Harry Howard - Tulane University 20 Magnetic Resonance Imaging (MRI) All atoms spin on their axes. Nuclei have a positive electronic charge, and any spinning charged particle will act as a magnet with north and south poles located on the axis of spin. The spin-axes of the nuclei in the subject line up with the chamber’s field, with the north poles of the nuclei pointing in the ‘southward’ direction of the field. Then a radio pulse is broadcast toward the subject. The pulse causes the axes of the nuclei to tilt with respect to the chamber’s magnetic field, and as it wears off, the axes gradually return to their resting position (within the magnetic field). As they do so, each nucleus becomes a miniature radio transmitter, giving out a characteristic pulse that changes over time, depending on the microenvironment surrounding it. For example, hydrogen nuclei in fats have a different microenvironment than do those in water, and thus transmit different pulses. Due to such contrasts, different tissues transmit different radio signals. These radio transmissions can be coordinated by a computer into an image. This method is known as magnetic resonance imaging (MRI), and it can be used to scan the human body safely and accurately

9/13/13Brain & Language - Harry Howard - Tulane University 21 functional Magnetic Resonance Imaging (fMRI) An elaboration of MRI called functional MRI (fMRI) has become the dominant technique for the study of the functional organization of the human brain during cognitive, perceptual, sensory, and motor tasks. As Gregg (2002) explains it, the principle of fMRI imaging is to take a series of images in quick succession and then to analyze them statistically for differences. For example, in the blood-oxygen-level dependent (BOLD) method introduced by Ogawa et al. (1990), the fact that hemoglobin and deoxyhemoglobin are magnetically different is exploited. Hemoglobin shows up better on MRI images than deoxyhemoglobin, which is to say that oxygenated blood shows up better then blood whose oxygen has been depleted by neural metabolism. This has been exploited in the following type of procedure: a series of baseline images are taken of the brain region of interest when the subject is at rest. The subject then performs a task, and a second series is taken. The first set of images is subtracted from the second, and the areas that are most visible in the resulting image are presumed to have been activated by the task.

NEXT TIME Ingram §3: Neuroanatomy of language, any leftovers ☞ Go over questions at end of chapter. 9/13/13Brain & Language - Harry Howard - Tulane University 22