The response properties of frequency-tuned neurons and or
|
|
- Martha Henderson
- 8 years ago
- Views:
Transcription
1 Centripetal and centrifugal reorganizations of frequency map of auditory cortex in gerbils Masashi Sakai and Nobuo Suga* Department of Biology, Washington University, One Brookings Drive, St. Louis, MO Contributed by Nobuo Suga, March 21, 2002 As repetitive acoustic stimulation and auditory conditioning do, electric stimulation of the primary auditory cortex (AI) evokes reorganization of the frequency map of AI, as well as of the subcortical auditory nuclei. The reorganization is caused by shifts in best frequencies (BFs) of neurons either toward (centripetal) or away from (centrifugal) the BF of stimulated cortical neurons. In AI of the Mongolian gerbil, we found that focal electrical stimulation evoked a centripetal BF shift in an elliptical area centered at the stimulated neurons and a centrifugal BF shift in a zone surrounding it. The 1.9-mm long major and 1.1-mm long minor axes of the elliptical area were parallel and orthogonal to the frequency axis, respectively. The width of the surrounding zone was mm. Such center-surround reorganization has not yet been found in any sensory cortex except AI of the gerbil. The ellipse is similar to the arborization pattern of pyramidal neurons, the major source of excitatory horizontal connections in AI, whereas the surrounding zone is compatible to the arborization range of small basket cells (inhibitory neurons) in AI. frequency tuning hearing plasticity tonotopic map The response properties of frequency-tuned neurons and or the cochleotopic (frequency) map of the primary auditory cortex (AI) can be changed by repetitive acoustic stimulation (1, 2), auditory conditioning (3 9), learning of a discrimination task (10), focal electric stimulation of AI (1, 2, 11), or electric stimulation of the basal forebrain paired with acoustic stimulation (6, 12). The response properties of frequency-tuned neurons and the cochleotopic map of the inferior colliculus (IC) also can be changed by repetitive acoustic stimulation (2, 13, 14), auditory conditioning (8, 9, 14), or focal electric stimulation of AI (2, 13, 15, 17). In the big brown bat, Eptesicus fuscus, cortical and collicular changes in the cochleotopic map are caused by the shift in best frequency (BF) of single neurons toward the frequency of a repetitively delivered acoustic stimulus (1, 13, 14), the frequency of a conditioned sound (8, 9, 14), or the BF of electrically stimulated AI neurons (1, 11, 13, 15). BF shifts resulting in reorganization of the frequency map are basically the same regardless of the means that evoked them and are attributable to activity within the corticofugal system (8, 9, 14). This means that focal electric stimulation of AI activates the essential portion of the neural mechanism for reorganization (plasticity) of the central auditory system and that it can be an appropriate method for the exploration of the plasticity of neurons in the IC, medial geniculate body, and AI (18). BF shifts toward and away from the BF of electrically stimulated cortical neurons or the frequency of a repetitively delivered tone burst are called centripetal and centrifugal BF shifts, respectively (18). A centripetal BF shift evoked by focal electric stimulation of AI has been found not only in the big brown bat, but also in the posterior division (nonspecialized portion) of AI of the mustached bat and AI of the Mongolian gerbil (11). Furthermore, centripetal shift of the receptive fields of neurons in the somatosensory cortex has been found in monkeys (19) and cats (20). Focal electric stimulation of AI also evokes centripetal shift in sound-duration tuning (15) of collicular neurons. Therefore, the corticofugal system modulates the functional organization of the IC not only in the frequency domain but also in the time domain. Centripetal reorganization is assumed to be widely shared with mammalian sensory systems. Reorganization of the frequency map is, however, different between specialized and nonspecialized areas of the auditory cortex of the mustached bat (11, 18). The Doppler-shifted constant frequency (DSCF) and frequency modulation (FM) FM areas of the auditory cortex of the mustached bat are specialized for processing either velocity and relative size information carried by 61 khz sound (21) or target-distance information carried by a pair of FM sounds (23, 24), respectively. Focal electric stimulation of the DSCF or FM FM area revokes centrifugal shift in the BFs of collicular, thalamic, (16, 25) and cortical (11) DSCF neurons or in the best delays of collicular and thalamic FM FM neurons (26), respectively. The direction of these shifts in BF and best delay is opposite to that in BF observed in the posterior division of AI of the mustached bat (11). Such a difference appears to be related to corticofugal facilitation and lateral inhibition of auditory responses of subcortical auditory neurons, so that the following working hypothesis has been proposed (18). If the positive feedback evoking facilitation is strong and widespread to neighboring neurons and negative feedback evoking lateral inhibition is weak, a focal activation of the auditory cortex evokes a prominent centripetal BF shift in a large area that is surrounded by a narrow zone of centrifugal BF shift. On the contrary, if the positive feedback is highly focused and negative feedback is strong and widespread to neighboring neurons, a focal activation of the auditory cortex evokes a prominent centrifugal BF shift in a large area surrounding a narrow zone where a small centripetal BF shift is evoked. In AI, there are excitatory neurons for horizontal connections and inhibitory neurons for local connections (27, 28), so that facilitation and inhibition for BF shifts may also occur. If the corticofugal system evokes facilitation and inhibition of subcortical neurons for processing auditory information and reorganizing the central auditory system, reorganization of AI of the Mongolian gerbil may consist of two types of areas: an area for centripetal BF shift at the center and an area for centrifugal BF shift in the surround. The aim of the current article is to report the data showing such a center-surround reorganization of the gerbil s AI. Materials and Methods Animal preparation and experimental procedures were exactly the same as those in our previous article (11). Eleven adult Mongolian gerbils (Meriones unguiculatus; g body weight) were used. Under anesthesia with ketamine (40 mg kg Abbreviations: AI, primary auditory cortex; BF, best frequency; IC, inferior colliculus; DSCF, Doppler-shifted constant frequency; FM, frequency modulation. *To whom reprint requests should be addressed. suga@biology.wustl.edu. The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked advertisement in accordance with 18 U.S.C solely to indicate this fact PNAS May 14, 2002 vol. 99 no cgi doi pnas
2 body weight) and meditomidine (0.26 mg kg body weight), the dorsal surface of the animal s skull was exposed, and four plastic sockets (1.5 mm in diameter and 2.0 mm in length) were attached onto the lateral sides of the skull with glue and dental cement. After the surgery (5 7 d), the animal was lightly anesthetized with the above drugs and placed in a polyethylene-foam bodymold suspended by an elastic band at the center of a soundproof room. The animal s head was immobilized by inserting metal rods into the sockets and adjusted to face directly at a loudspeaker located 74 cm away. Holes ( m in diameter) were made on the skull covering AI. Tungsten-wire electrodes ( 7 m in tip diameter) for electric stimulation or recording action potentials were orthogonally inserted into AI through the holes. The protocol of our research was approved by the Animal Studies Committee of Washington University. A tone burst (20 ms-long, 0.5-ms rise decay time) was delivered at a rate of 5 s from the loudspeaker. The BF, minimum threshold, and frequency-tuning curve of a single neuron were first measured audiovisually. A computer-controlled frequency scan (22 time blocks, 200 ms long for each) was then delivered, in which the frequency of the tone burst was shifted in steps of a given frequency ( khz) across the BF of the neuron. The amplitude of tone bursts in the scan was set at 10 db above the minimum threshold of a given neuron. Identical frequency scans were repeated at a rate of 1 5 s. A pair of tungsten-wire microelectrodes ( 7 m in tip diameter; 150- m apart, one proximal to the other) was inserted into cortical layer V ( m in depth). Action potentials originating from 2 3 single neurons were recorded and the BFs of the neurons were measured. Then they were electrically stimulated through the pair of electrodes. The electric stimulation was a 6.2-ms long train of four monophasic electric pulses (100 na, 0.2-ms duration, 2.0-ms intervals). The train was delivered at a rate of 10 s for 30 min. Action potentials of a single cortical neuron were selected with an amplitude-window discriminator (BAK Electronic, Rockville, MD; model DLS-1), and an action potential of the neuron was stored on the screen of a digital storage oscilloscope at the beginning of data acquisition. The responses of a cortical neuron to 50 identical frequency scans were displayed as an array of peri-stimulus-time histograms as a function of frequency. Acquisition of peri-stimulus-time histograms was continued as long as action potentials visually matched the template, the stored action potential of the neuron. Data were stored on a computer hard drive and analyzed off-line for BF shifts. If a shifted BF and frequency-response curve did not shift back by more than 50% of the change, the data were excluded from the analysis. In stable recordings, all BFs and curves shifted by the electric stimulation recovered by 50%. This recovery itself helped to prove that the shift was significant. Results In AI of the gerbil, frequencies between 0.2 and 43 khz are systematically mapped along the caudo-rostral axis, and frequencies from 0.5 to 5 khz are over-represented (ref. 29; Fig. 1). The BFs of electrically stimulated cortical neurons ranged between 0.88 and 1.2 khz (mean SE: khz, n 97) and those of recorded cortical neurons ranged between 0.1 and 37.0 khz ( khz, n 338). Electric stimulation of cortical neurons evoked a BF shift of other cortical neurons located near those stimulated. When stimulated and recorded neurons were tuned to 1.00 and 1.60 khz, respectively (Fig. 2Aa1), electric stimulation suppressed the response of the recorded neuron at 1.60 khz (Fig. 2Ab2) and facilitated its response at 1.20 khz (Fig. 2Ac2). As a result, the BF and the frequency-response curve of the neuron shifted toward the BF of the stimulated neurons (Fig. 2Aa2). In other words, centripetal BF shift was evoked. When stimulated and recorded neurons were tuned to 1.10 and 1.45 Fig. 1. Frequency map of the AI of the Mongolian gerbil. Dashed lines: iso-bf lines [based on Thomas et al. (29)]. Electric stimulation was delivered to cortical neurons within the shaded area. (Inset) Dorsolateral view of the gerbil brain. The auditory cortex (AC) consists of seven areas. The black area is the AI. khz, respectively (Fig. 2Ba1), the electric stimulation suppressed response of the recorded neuron at 1.45 khz (Fig. 2Bb2) and facilitated its response at 1.65 khz (Fig. 2Bc2). As a result, the BF and frequency-response curve of the neuron shifted away Fig. 2. Shifts in frequency-response curve evoked by cortical electric stimulation. (Aa and Ba) Array of peristimulus-time cumulative histograms representing frequency-response curves. (Ab, Ac, Bb, and Bc) Peristimulus-time histograms representing responses at BF in the control (BF c ) or shifted (BF s ) condition. (A) The BF at 1.60 khz shifted toward the 1.00 khz BF of the stimulated cortical neurons (centripetal BF shift). (B) The BF at 1.45 khz shifted away from the 1.10-kHz BF of the stimulated cortical neurons (centrifugal BF shift). (Aa2 and Ba2, arrow) The BF of the stimulated cortical neurons. (Aa and Ba) The BFs of the recorded cortical neuron in the control F, shifted, and recovered E conditions, respectively. 1 4: The data obtained before, and 30, 80, and 180 min after the onset of electric stimulation. The horizontal bars at the bottom of b and c represent 20 ms-long tone bursts. NEUROBIOLOGY Sakai and Suga PNAS May 14, 2002 vol. 99 no
3 Fig. 3. Distributions of cortical neurons that exhibited centripetal or centrifugal BF shift. (A) Centripetal BF shift, F. (B) Centrifugal BF shift, Œ and no BF shift,. Locations of recorded neurons along the cortical surface are plotted relative to that of stimulated cortical neurons at the origin of the coordinates. x and y axes, directions parallel or orthogonal to the frequency axis of the AI. Data are pooled from 16 hemispheres of 11 animals. BF e,bfof electrically stimulated cortical neurons. Ellipse, area for centripetal or centrifugal BF shifts. from the BF of the stimulated neuron (Fig. 2Ba2). In other words, centrifugal BF shift was evoked. The shifted BF and changed responses returned to the control condition 180 min after the electric stimulation (a4, b4, and c4 in Fig. 2 A and B). Whether a recorded neuron showed a centripetal or centrifugal BF shift depended on the distance between the recorded and stimulated neurons along the surface of AI. Neurons showing a centripetal BF shift were distributed in an elliptical area centered at the stimulated neurons that were tuned to 1.1 khz on the average (Fig. 3A). Their confidence ellipse (based on Mahalanobis s equation P 0.95) crossed the rostro-caudal (x) axis at approximately 0.56 mm and the 1.1-kHz iso-bf line (y) axis at approximately 0.95 mm. On the other hand, neurons showing centrifugal BF shift distributed in a zone surrounding the elliptical area (Fig. 3B). The confidence ellipse crossed the x axis at approximately 0.72 mm and the y axis at approximately 1.23 mm. This surround was mm wide. In this surrounding zone, many neurons ( 57%) did not show a BF shift (see Discussion). The amount of BF shift varied depending on the distance and the BF difference between recorded and stimulated neurons. BF shifts measured in a 0.6-mm wide horizontal zone containing the electrically stimulated neurons at its center (zone B in Fig. 4A) were pooled because the edge of the ellipse in this zone was somewhat parallel to the 1.1-kHz iso-bf line. The pooled data were plotted as a function of the distance between stimulated and recorded neurons along the frequency axis. The great majority of neurons within 0.45 mm from the 1.1-kHz iso-bf line showed centripetal BF shift, and the amounts of their BF shifts were nonmonotonically related to the distance (Fig. 4B, circles). The largest negative centripetal BF shift occurred at 0.21 mm rostral to the 1.1-kHz iso-bf line (corresponding to a BF 1.8 khz higher than 1.1 khz), whereas the largest positive centripetal BF shift occurred at 0.27 mm caudal (corresponding to a BF 0.36 khz lower than 1.1 khz). The largest positive centrifugal BF shift occurred at 0.48 mm rostral to the 1.1-kHz iso-bf line, whereas the largest negative centrifugal BF shift occurred at 0.62 mm caudal (Fig. 4B, triangles). Both centripetal and centrifugal BF shifts were significantly larger on the rostral side than on the caudal side (P 0.05). However, there was no Fig. 4. Amount of BF shift in a zone parallel or orthogonal to the frequency axis of the AI. (A) Confidence ellipses for neurons that showed centripetal (open area) or centrifugal BF shift (shaded area) (see Fig. 3). The directions and amounts of BF shifts of neurons in the rostro-caudal (B) and dorso-ventral (C and D) zones in A were respectively plotted in B D as a function of distance along the cortical surface. BF e, BF of electrically stimulated cortical neuron and BF r, BF of recorded cortical neuron. See Inset at the top right for symbols. difference in distance for evoking a BF shift between the rostral and caudal sides (Fig. 4B). BF shifts measured within a 0.30-mm wide vertical zone caudal or rostral to the 1.1-kHz iso-bf line (C and D in Fig. 4A) were pooled because the edge of the ellipse in this zone was somewhat parallel to the frequency axis. The pooled data were plotted as a function of distance between stimulated and recorded neurons along the 1.1-kHz iso-bf line. The amount of BF shift was larger on the rostral side of the 1.1-kHz iso-bf line (Fig. 4D) than on its caudal side (Fig. 4C). Centripetal and centrifugal BF shifts both occurred symmetrically in amount on the dorsal and ventral sides of the stimulated cortical neurons. Within 0.60 mm, the great majority of the neurons showed centripetal BF shift, and there was a tendency that the farther the distance away from the stimulated neurons along the iso-bf line, the smaller the centripetal BF shift. Large BF shifts ( khz) occurred within 0.10 mm. This finding indicated that BF shift was the largest along the frequency axis crossing the electrically stimulated cortical neurons. Centrifugal BF shift occurred at distances between 0.5 and 1.1 mm from the stimulated neurons (Fig. 4 C and D). In the normal (i.e., control) condition, a BF systematically increased from the caudal end to the rostral end of AI (Fig. 5A, E). There was no sign that 1.1 khz was over-represented. However, BF shift evoked by electric stimulation of 1.1-kHztuned cortical neurons resulted in the prominent overrepresentation of 1.1-kHz sound, so that the iso-1.1-khz BF line changed into a 0.33-mm wide band (Fig. 5A, F). A small under-representation also occurred for frequencies lower than 0.52 khz and higher than 4.7 khz (Fig. 5A, Œ). Because a BF systematically changed along the caudo-rostral axis of AI, the amount of BF shifts shown in Fig. 4B could be expressed in mm (Fig. 5B). When expressed in this manner, the maximum amount of centripetal BF shift in mm was 0.21 or 0.22 mm on the rostral side and on the caudal side, respectively. Therefore, the prominent asymmetrical BF shift in khz (Fig. 4B) was related to the nonlinear frequency axis in AI: a small BF cgi doi pnas Sakai and Suga
4 Fig. 5. Change in the frequency axis (A) and amount of BF shift expressed in millimeters (B). (A) Over- (between the arrows) and under- (between the vertical lines) representations evoked by cortical electric stimulation. Distribution of BFs along the caudo-rostral axis of the AI was studied before (E) and after (F and Œ) electric stimulation of 1.1-kHz-tuned cortical neurons. (B) Distributions of centripetal (F) and centrifugal (Œ) BF shifts expressed in millimeters along the caudo-rostral axis of the AI. change unit distance below 1.1 khz and a large BF change unit distance above 1.1 khz. Discussion Centripetal and Centrifugal BF Shifts. The amount of BF shifts we observed was related to differences in both distance and BF between recorded and stimulated cortical neurons. Iso-BF lines are not necessarily parallel to each other and show some variations among animals (22, 29). Accordingly, in our data pooled from 11 animals, the area for the centripetal BF shift overlapped with the surround for the centrifugal BF shift, and neurons showing centrifugal BF shift were intermixed with those showing no BF shift. Irrespective of these overlap and intermixture, our present data clearly show center-surround reorganization and favor the hypothesis stating that facilitation and lateral inhibition, respectively, evoke centripetal and centrifugal BF shifts (18). In AI of the big brown bat (1, 2) and the nonspecialized posterior division of AI of the mustached bat (11), cortical electric stimulation evokes a centripetal BF shift around the stimulated cortical neurons and a centrifugal BF shift at the rostral or caudal edge of the centripetal BF shift area. However, in previous studies, the number of neurons showing centrifugal BF shift was so small that the center-surround reorganization was not clear. In rats (12), electric stimulation of the basal forebrain paired with an acoustic stimulus evokes a centripetal BF shift. Centrifugal BF shift has not been reported in any species of animals except the above two species of bats. In our previous studies of the gerbil AI (11), we found 65 centripetal BF shifts and two centrifugal BF shifts, so that center-surround reorganization was unclear. However, it became clear in our present detailed studies. Therefore, it is expected that centersurround reorganization will also be found in the auditory cortex of other species of animals by further studies. In rats, a train of tone bursts paired with electric stimulation of dopaminergic neurons in the ventral tegmental area evokes over-representation of the frequency of these tones and under-representation of frequencies lower or higher than that in AI (30). However, it has not yet been shown whether this under-representation is partly caused by centrifugal BF shifts. In a nonspecialized auditory cortex such as the gerbil s AI, reorganization is predominantly based on centripetal BF shift. A centripetal BF shift would result in over-representation of a particular value of a parameter characterizing an acoustic signal (1, 2, 11, 13, 14). A specialized auditory cortex such as the DSCF area of the mustached bat over-represents particular values of frequency in a narrow range in the natural condition (21, 22). In the DSCF area, reorganization is predominantly based on centrifugal BF shift, which would increase contrast in neural representation of a particular value of frequency characterizing an acoustic signal (16, 25). Receptive-Field Shift in Nonauditory Sensory Systems. In monkeys (19), excess stimulation of a digit evokes over-representation of that digit in the primary somatosensory cortex. This overrepresentation is caused by centripetal shift of the receptive fields of cortical neurons representing a cutaneous area around that digit. Thus far, centrifugal shift of a receptive field has not been reported for reorganization of the somatosensory system. In cats, electric stimulation of the primary visual cortex evokes lateral inhibition associated with focused facilitation in the lateral geniculate nucleus (31) as in the auditory system of the mustached bat (16, 25, 26). However, shift in receptive field has not been reported. Neurons in the primary visual cortex have excitatory inputs surrounded by inhibitory inputs (32, 33). However, the diameter of the area for excitatory inputs ( 0.25 mm) is almost one-half of the minor axis of the elliptical area for a centripetal BF shift in the gerbil s AI( 0.60 mm). The estimated density of horizontal axon collaterals of pyramidal neurons in the visual cortex is approximately one-half of that in the auditory cortex (34). Because the most basic cortical neural circuit (27, 35) and the corticothalamic feedback loop are presumably shared by the auditory (36), visual (37), and somatosensory (38) cortices, it is likely that center-surround reorganization also may be shared by these sensory systems. Anatomical Correlates to Centripetal and Centrifugal BF Shifts. Axon collaterals of pyramidal neurons in the primary auditory, somatosensory, visual, and motor cortices extend 1,500 m (27) for the integration of neural signals and the synchronization of the activities of distant neurons (39, 40). Horizontal excitatory connections are mostly under tonic inhibition produced by GABAergic interneurons (41). The balance between excitatory and inhibitory influences on single neurons is the primary determinant of the representation of the somatosensory (42), visual (40), and motor cortices (43). In the somatosensory cortex, expansion of representation of a body surface caused by training (10) or cortical electric stimulation (19), especially its rapid change, is the result of unmasking, i.e., conversion of subthreshold excitatory-horizontal connections to suprathreshold connections. In a slice preparation of AI, synaptic efficiency between horizontal fibers is highly modifiable (44). In the bat s AI, BF shifts evoked by cortical electric stimulation occur within several minutes and disappears within 180 min after the cessation of the NEUROBIOLOGY Sakai and Suga PNAS May 14, 2002 vol. 99 no
5 stimulation (Fig. 2; 1, 2). This short-term plasticity is probably not caused by to a morphological change such as axonal sprouting or dendritic spine proliferation, but by a change in efficiency of existing synapses. Horizontal arborizations of pyramidal neurons for excitation are classified into short- and long-range projections. The former is up to 0.6 mm long and omnidirectional, whereas the latter is several millimeters long and preferentially parallel to iso-bf contour lines (45 48). The minor and major axes of the ellipse for centripetal BF shift may, respectively, be related to the shortand long-range projections. The recurrent fibers of pyramidal neurons form an intracortical positive feedback loop. In the big brown bat, electric stimulation of AI evokes a BF shift in the IC, which is very similar to cortical BF shift (2), and the cortical BF shift greatly depends on the collicular BF shift (8, 9). Because centripetal BF shift in the IC is evoked by the corticofugal system (9, 13, 14, 18), the recurrent circuit within AI and the corticofugal feedback loop both presumably play an important role in cortical plasticity. The recurrent fibers of pyramidal neurons spread over 0.6 to several millimeters within the cortex and form minute connections with inhibitory interneurons. Most inhibitory interneurons (small basket cells) in AI radially project mm, forming uniformly dense arbors (28, 49). The surrounding zone for centrifugal BF shift was mm away from the stimulated cortical neurons and mm wide. Therefore, the inhibitory interneurons activated by the recurrent fibers may produce lateral inhibition that evokes a centrifugal BF shift. We thank N. Laleman and K. K. Ohlemiller for comments on the manuscript. Our research has been supported by a research grant from the National Institute on Deafness and Other Communicative Disorders (DC 00175). 1. Chowdhury, S. A. & Suga, N. (2000) J. Neurophysiol. 83, Ma, X. & Suga, N. (2001) J. Neurophysiol. 85, Disterhoft, J. F. & Olds, J. (1972) J. Neurophysiol. 35, Diamond, D. M. & Weinberger, N. M. (1986) Brain Res. 372, Edeline, J. M. & Weinberger, N. M. (1993) Behav. Neurosci. 107, Bakin, J. S. & Weinberger N. M. (1996) Proc. Natl. Acad. Sci. USA 93, Ohl, F. W. & Scheich, H. (1996) Eur. J. Neurosci. 8, Gao, E. & Suga, N. (2000) Proc. Natl. Acad. Sci. USA 97, Ji, W., Gao, E. & Suga, N. (2001) J. Neurophysiol. 86, Recanzone, G. H., Schreiner, C. E. & Merzenich, M. M. (1993) J. Neurosci. 13, Sakai, M. & Suga, N. (2001) Proc. Natl. Acad. Sci. USA 98, Kilgard, M. P. & Merzenich, M. M. (1998) Science 279, Gao, E. & Suga, N. (1998) Proc. Natl. Acad. Sci. USA 95, Ma, X. & Suga, N. (2001) Proc. Natl. Acad. Sci. USA 98, Yan, W. & Suga, N. (1998) Nat. Neurosci. 1, Zhang, Y. & Suga, N. (2000) J. Neurophysiol. 84, Jen, P. H.-S., Chen, Q. C. & Sun, X. D. (1998) J. Comp. Physiol. A 183, Suga, N., Gao, E., Zhang, Y., Ma, X. & Olsen, J. F. (2000) Proc. Natl. Acad. Sci. USA 97, Recanzone, G. H., Merzenich, M. M. & Dinse, H. R. (1992) Cereb. Cortex 2, Kano, M., Iino, K. & Kano, M. (1991) NeuroReport 2, Suga, N. & Jen, P. H.-S. (1976) Science 194, Suga, N. & Manabe, T. (1982) J. Neurophysiol. 47, Suga, N. & O Neill, W. E. (1979) Science 203, O Neill, W. E. & Suga, N. (1979) Science 203, Yan, J. & Suga, N. (1996) Science 273, Zhang, Y., Suga, N. & Yan, J. (1997) Nature (London) 387, Szentagothai, J. (1975) Brain Res. 95, Prieto, J. J., Peterson, B. A. & Winer, J. A. (1994) J. Comp. Neurol. 344, Thomas, H., Tillein, J., Heil, P. & Scheich, H. (1993) Eur. J. Neurosci. 5, Bao, S., Chan, V. T. & Merzenich, M. M. (2001) Nature (London) 412, Tsumoto, T., Creutzfeldt, O. D. & Legendy, C. R. (1978) Exp. Brain Res. 32, Dalva, M. B., Weliky, M. & Katz, L. C. (1997) Neuron 19, Hirsch, J. A. & Gilbert, C. D. (1991) J. Neurosci. 11, Kudoh, M. & Shibuki, K. (1997) J. Neurosci. 17, Creutzfeldt, O. D. (1977) Naturwissenschaften 64, Huffman, R. F. & Henson, O. W., Jr. (1990) Brain. Res. Rev. 15, Murphy, P. C., Duckett, S. G. & Sillito, A. M. (1999) Science 286, Krupa, D. J., Ghazanfar, A. A. & Nicolelis, M. A. (1999) Proc. Natl. Acad. Sci. USA 96, Buonomano, D. V. & Merzenich, M. M. (1998) Annu. Rev. Neurosci. 21, Gilbert, C. D. (1998) Physiol. Rev. 78, Jones, E. G. (1993) Cereb. Cortex 3, Merzenich, M. M. & Jenkins, W. M. (1993) J. Hand Ther. 6, Donoghue, J. P. (1995) Curr. Opin. Neurobiol. 5, Buonomano, D. V. (1999) J. Neurosci. 19, Reale, R. A., Brugge, J. F. & Feng, J. Z. (1983) Proc. Natl. Acad. Sci. USA 80, Winer, J. A. (1984) J. Comp. Neurol. 229, Matsubara, J. A. & Phillips, D. P. (1988) J. Comp. Neurol. 268, Ojima, H., Honda, C. N. & Jones, E. G. (1991) Cereb. Cortex 1, Hendry, S. H. & Jones, E. G. (1991) Brain Res. 543, cgi doi pnas Sakai and Suga
In the big brown bat (Eptesicus fuscus), auditory responses,
Plasticity of the cochleotopic (frequency) map in specialized and nonspecialized auditory cortices Masashi Sakai and Nobuo Suga* Department of Biology, Washington University, One Brookings Drive, St. Louis,
More informationLateral Inhibition for Center-Surround Reorganization of the Frequency Map of Bat Auditory Cortex
J Neurophysiol 92: 3192 3199, 2004; doi:10.1152/jn.00301.2004. Lateral Inhibition for Center-Surround Reorganization of the Frequency Map of Bat Auditory Cortex Xiaofeng Ma and Nobuo Suga Department of
More informationPlasticity of Bat s Central Auditory System Evoked by Focal Electric Stimulation of Auditory and/or Somatosensory Cortices
Plasticity of Bat s Central Auditory System Evoked by Focal Electric Stimulation of Auditory and/or Somatosensory Cortices XIAOFENG MA AND NOBUO SUGA Department of Biology, Washington University, St. Louis,
More informationThis contribution is part of the special series of Inaugural Articles by members of the National Academy of Sciences elected on April 28, 1998.
Proc. Natl. Acad. Sci. USA Vol. 95, pp. 12663 12670, October 1998 Neurobiology This contribution is part of the special series of Inaugural Articles by members of the National Academy of Sciences elected
More informationAuditory neuroanatomy: the Spanish heritage. Santiago Ramón y Cajal, 1852 1934
Auditory neuroanatomy: the Spanish heritage Santiago Ramón y Cajal, 1852 1934 Rafael Lorente de Nó, 1902 1990 3 The nervous system is made up of cells. Estimates of the number of cells vary from
More informationMasters research projects. 1. Adapting Granger causality for use on EEG data.
Masters research projects 1. Adapting Granger causality for use on EEG data. Background. Granger causality is a concept introduced in the field of economy to determine which variables influence, or cause,
More informationCHAPTER 6 PRINCIPLES OF NEURAL CIRCUITS.
CHAPTER 6 PRINCIPLES OF NEURAL CIRCUITS. 6.1. CONNECTIONS AMONG NEURONS Neurons are interconnected with one another to form circuits, much as electronic components are wired together to form a functional
More informationTHE HUMAN BRAIN. observations and foundations
THE HUMAN BRAIN observations and foundations brains versus computers a typical brain contains something like 100 billion miniscule cells called neurons estimates go from about 50 billion to as many as
More informationSynaptic depression creates a switch that controls the frequency of an oscillatory circuit
Proc. Natl. Acad. Sci. USA Vol. 96, pp. 8206 8211, July 1999 Neurobiology Synaptic depression creates a switch that controls the frequency of an oscillatory circuit FARZAN NADIM*, YAIR MANOR, NANCY KOPELL,
More information2 Neurons. 4 The Brain: Cortex
1 Neuroscience 2 Neurons output integration axon cell body, membrane potential Frontal planning control auditory episodes soma motor Temporal Parietal action language objects space vision Occipital inputs
More informationVisual area MT responds to local motion. Visual area MST responds to optic flow. Visual area STS responds to biological motion. Macaque visual areas
Visual area responds to local motion MST a Visual area MST responds to optic flow MST a Visual area STS responds to biological motion STS Macaque visual areas Flattening the brain What is a visual area?
More informationMotor dysfunction 2: Spinal cord injury and subcortical motor disorders ANATOMY REVIEW: Basal Ganglia
Motor dysfunction 2: Spinal cord injury and subcortical motor disorders ANATOMY REVIEW: Basal Ganglia A group of subcortical nuclei caudate, putamen, globus pallidus Caudate & Putamen = Neostriatum caudate
More informationHow songbirds sing birdsongs?
How songbirds sing birdsongs? Liora Las Michale Fee (MIT) Veit et al. (J Neurophysiol, 2011) Outline: 1) Introduction to songbirds as a model. 2) Neuronal circuits underlying mature song production (motor
More informationProcessing the Image or Can you Believe what you see? Light and Color for Nonscientists PHYS 1230
Processing the Image or Can you Believe what you see? Light and Color for Nonscientists PHYS 1230 Optical Illusions http://www.michaelbach.de/ot/mot_mib/index.html Vision We construct images unconsciously
More informationNervous System: Spinal Cord and Spinal Nerves (Chapter 13) Lecture Materials for Amy Warenda Czura, Ph.D. Suffolk County Community College
Nervous System: Spinal Cord and Spinal Nerves (Chapter 13) Lecture Materials for Amy Warenda Czura, Ph.D. Suffolk County Community College Primary Sources for figures and content: Eastern Campus Marieb,
More informationIn a number of recent studies investigators have shown that
Cortical correlates of learning in monkeys adapting to a new dynamical environment F. Gandolfo, C.-S. R. Li*, B. J. Benda, C. Padoa Schioppa, and E. Bizzi Department of Brain and Cognitive Sciences, Massachusetts
More informationWiring optimization in the brain
Wiring optimization in the brain Dmitri B. Chklovskii Sloan Center for Theoretical Neurobiology The Salk Institute La Jolla, CA 92037 mitya@salk.edu Charles F. Stevens Howard Hughes Medical Institute and
More informationThe Visual Cortex 0 http://www.tutis.ca/neuromd/index.htm 20 February 2013
T he Visual Cortex 0 Chapter contents Contents Chapter 2... 0 T he Visual Cortex... 0 Chapter Contents... 1 Introduction... 2 Optic Chiasm... 2 Where do the eye's ganglion cells project to?... 3 To where
More informationNeu. al Network Analysis of Distributed Representations of Dynamical Sensory-Motor rrransformations in the Leech
28 Lockery t Fang and Sejnowski Neu. al Network Analysis of Distributed Representations of Dynamical Sensory-Motor rrransformations in the Leech Shawn R. LockerYt Van Fangt and Terrence J. Sejnowski Computational
More informationTinnitus and the Brain
Tinnitus and the Brain Dirk De Ridder & Berthold Langguth Moving animals have developed a brain in order to reduce the inherent uncertainty present in an ever changing environment. The auditory system
More informationLearning with Your Brain. Teaching With the Brain in Mind
Learning with Your Brain Should what (and how) we teach be associated with what we know about the brain and the nervous system? Jonathan Karp, Ph.D. Dept of Biology 5/20/2004 Teaching With the Brain in
More informationAgent Simulation of Hull s Drive Theory
Agent Simulation of Hull s Drive Theory Nick Schmansky Department of Cognitive and Neural Systems Boston University March 7, 4 Abstract A computer simulation was conducted of an agent attempting to survive
More informationPlate waves in phononic crystals slabs
Acoustics 8 Paris Plate waves in phononic crystals slabs J.-J. Chen and B. Bonello CNRS and Paris VI University, INSP - 14 rue de Lourmel, 7515 Paris, France chen99nju@gmail.com 41 Acoustics 8 Paris We
More informationChapter 14: The Cutaneous Senses
Chapter 14: The Cutaneous Senses Skin - heaviest organ in the body Cutaneous System Epidermis is the outer layer of the skin, which is made up of dead skin cells Dermis is below the epidermis and contains
More informationAppendix 4 Simulation software for neuronal network models
Appendix 4 Simulation software for neuronal network models D.1 Introduction This Appendix describes the Matlab software that has been made available with Cerebral Cortex: Principles of Operation (Rolls
More informationThe Physiology of the Senses Lecture 1 - The Eye www.tutis.ca/senses/
The Physiology of the Senses Lecture 1 - The Eye www.tutis.ca/senses/ Contents Objectives... 2 Introduction... 2 Accommodation... 3 The Iris... 4 The Cells in the Retina... 5 Receptive Fields... 8 The
More informationModels of Cortical Maps II
CN510: Principles and Methods of Cognitive and Neural Modeling Models of Cortical Maps II Lecture 19 Instructor: Anatoli Gorchetchnikov dy dt The Network of Grossberg (1976) Ay B y f (
More informationSelf Organizing Maps: Fundamentals
Self Organizing Maps: Fundamentals Introduction to Neural Networks : Lecture 16 John A. Bullinaria, 2004 1. What is a Self Organizing Map? 2. Topographic Maps 3. Setting up a Self Organizing Map 4. Kohonen
More informationComparative collicular tonotopy in two bat species adapted to movement detection, Hipposideros speoris and Megaderma lyra
J Comp Physiol A (1988) 163:271-285 Journal of Comparative Physiology A Springer-Verlag 1988 Comparative collicular tonotopy in two bat species adapted to movement detection, Hipposideros speoris and Megaderma
More informationNeurotrophic factors and Their receptors
Neurotrophic factors and Their receptors Huang Shu-Hong Institute of neurobiology 1 For decades, scientists believed that brain cells of the central nervous system could not regrow following damage due
More informationAIR RESONANCE IN A PLASTIC BOTTLE Darrell Megli, Emeritus Professor of Physics, University of Evansville, Evansville, IN dm37@evansville.
AIR RESONANCE IN A PLASTIC BOTTLE Darrell Megli, Emeritus Professor of Physics, University of Evansville, Evansville, IN dm37@evansville.edu It is well known that if one blows across the neck of an empty
More informationPrevention & Recovery Conference November 28, 29 & 30 Norman, Ok
Prevention & Recovery Conference November 28, 29 & 30 Norman, Ok What is Addiction? The American Society of Addiction Medicine (ASAM) released on August 15, 2011 their latest definition of addiction:
More informationChapter 7: The Nervous System
Chapter 7: The Nervous System Objectives Discuss the general organization of the nervous system Describe the structure & function of a nerve Draw and label the pathways involved in a withdraw reflex Define
More informationBrain Maps The Sensory Homunculus
Brain Maps The Sensory Homunculus Our brains are maps. This mapping results from the way connections in the brain are ordered and arranged. The ordering of neural pathways between different parts of the
More informationA Game of Numbers (Understanding Directivity Specifications)
A Game of Numbers (Understanding Directivity Specifications) José (Joe) Brusi, Brusi Acoustical Consulting Loudspeaker directivity is expressed in many different ways on specification sheets and marketing
More informationVideo-Based Eye Tracking
Video-Based Eye Tracking Our Experience with Advanced Stimuli Design for Eye Tracking Software A. RUFA, a G.L. MARIOTTINI, b D. PRATTICHIZZO, b D. ALESSANDRINI, b A. VICINO, b AND A. FEDERICO a a Department
More informationVocabulary & General Concepts of Brain Organization
Vocabulary & General Concepts of Brain Organization Jeanette J. Norden, Ph.D. Professor Emerita Vanderbilt University School of Medicine Course Outline Lecture 1: Vocabulary & General Concepts of Brain
More informationNeurobiology of Depression in Relation to ECT. PJ Cowen Department of Psychiatry, University of Oxford
Neurobiology of Depression in Relation to ECT PJ Cowen Department of Psychiatry, University of Oxford Causes of Depression Genetic Childhood experience Life Events (particularly losses) Life Difficulties
More informationStructure and Function of Neurons
CHPTER 1 Structure and Function of Neurons Varieties of neurons General structure Structure of unique neurons Internal operations and the functioning of a neuron Subcellular organelles Protein synthesis
More informationMeasurement of Output Power Density from Mobile Phone as a Function of Input Sound Frequency
, pp. 270-279. A Publication of the Measurement of Output Power Density from Mobile Phone as a Function of Input Sound Frequency Emanuele Calabrò and Salvatore Magazù Department of Physics, University
More informationCSE511 Brain & Memory Modeling. Lect04: Brain & Spine Neuroanatomy
CSE511 Brain & Memory Modeling CSE511 Brain & Memory Modeling Lect02: BOSS Discrete Event Simulator Lect04: Brain & Spine Neuroanatomy Appendix of Purves et al., 4e Larry Wittie Computer Science, StonyBrook
More informationA Biophysical Network Model Displaying the Role of Basal Ganglia Pathways in Action Selection
A Biophysical Network Model Displaying the Role of Basal Ganglia Pathways in Action Selection Cem Yucelgen, Berat Denizdurduran, Selin Metin, Rahmi Elibol, N. Serap Sengor Istanbul Technical University,
More informationRock Bolt Condition Monitoring Using Ultrasonic Guided Waves
Rock Bolt Condition Monitoring Using Ultrasonic Guided Waves Bennie Buys Department of Mechanical and Aeronautical Engineering University of Pretoria Introduction Rock Bolts and their associated problems
More informationA Segmentation Algorithm for Zebra Finch Song at the Note Level. Ping Du and Todd W. Troyer
A Segmentation Algorithm for Zebra Finch Song at the Note Level Ping Du and Todd W. Troyer Neuroscience and Cognitive Science Program, Dept. of Psychology University of Maryland, College Park, MD 20742
More informationTheta, Gamma, and Working Memory
Theta, Gamma, and Working Memory Lecture 3.8 David S. Touretzky November, 2015 Outline Theta rhythm and gamma rhythms Phase precession in hippocampus Theta and gamma in entorhinal cortex Lisman working
More informationBi 360: Midterm Review
Bi 360: Midterm Review Basic Neurobiology 1) Many axons are surrounded by a fatty insulating sheath called myelin, which is interrupted at regular intervals at the Nodes of Ranvier, where the action potential
More informationEXPERIMENTAL ERROR AND DATA ANALYSIS
EXPERIMENTAL ERROR AND DATA ANALYSIS 1. INTRODUCTION: Laboratory experiments involve taking measurements of physical quantities. No measurement of any physical quantity is ever perfectly accurate, except
More informationActivity 5: The Action Potential: Measuring Its Absolute and Relative Refractory Periods. 250 20 Yes. 125 20 Yes. 60 20 No. 60 25 No.
3: Neurophysiology of Nerve Impulses (Part 2) Activity 5: The Action Potential: Measuring Its Absolute and Relative Refractory Periods Interval between stimuli Stimulus voltage (mv) Second action potential?
More informationA wave lab inside a coaxial cable
INSTITUTE OF PHYSICS PUBLISHING Eur. J. Phys. 25 (2004) 581 591 EUROPEAN JOURNAL OF PHYSICS PII: S0143-0807(04)76273-X A wave lab inside a coaxial cable JoãoMSerra,MiguelCBrito,JMaiaAlves and A M Vallera
More informationExperience-Dependent Plasticity in S1 Caused by Noncoincident Inputs
J Neurophysiol 94: 2239 2250, 2005; doi:10.1152/jn.00172.2005. Experience-Dependent Plasticity in S1 Caused by Noncoincident Inputs David T. Blake, Fabrizio Strata, Richard Kempter, and Michael M. Merzenich
More informationAS COMPETITION PAPER 2007 SOLUTIONS
AS COMPETITION PAPER 2007 Total Mark/50 SOLUTIONS Section A: Multiple Choice 1. C 2. D 3. B 4. B 5. B 6. A 7. A 8. C 1 Section B: Written Answer Question 9. A mass M is attached to the end of a horizontal
More informationConvention Paper Presented at the 112th Convention 2002 May 10 13 Munich, Germany
Audio Engineering Society Convention Paper Presented at the 112th Convention 2002 May 10 13 Munich, Germany This convention paper has been reproduced from the author's advance manuscript, without editing,
More informationReflex Physiology. Dr. Ali Ebneshahidi. 2009 Ebneshahidi
Reflex Physiology Dr. Ali Ebneshahidi Reflex Physiology Reflexes are automatic, subconscious response to changes within or outside the body. a. Reflexes maintain homeostasis (autonomic reflexes) heart
More informationHow To Understand The Distributed Potential Of A Dendritic Tree
Systems Biology II: Neural Systems (580.422) Lecture 8, Linear cable theory Eric Young 5-3164 eyoung@jhu.edu Reading: D. Johnston and S.M. Wu Foundations of Cellular Neurophysiology (MIT Press, 1995).
More informationSecond Quarterly Progress Report NO1-DC-6-2111 The Neurophysiological Effects of Simulated Auditory Prosthesis Stimulation
Second Quarterly Progress Report NO1-DC-6-2111 The Neurophysiological Effects of Simulated Auditory Prosthesis Stimulation J.T. Rubinstein, A.J. Matsuoka, P.J. Abbas, and C.A. Miller Department of Otolaryngology
More informationNeuron. Neurostimulation 2/8/2011
Direct Current Stimulation Promotes BDNF Dependent Synaptic Plasticity: Potential Implications for Motor Learning B Fritsch, J Reis, K Martinowich, HM Schambra, Yuanyuan Ji, LG Cohen & B Lu Leonardo G.
More informationAssembly and use of an economical, optical fiber-based uncaging system
Kandler et al. 1 Assembly and use of an economical, optical fiber-based uncaging system Karl Kandler 1, Tuan Nguyen 1, Jihyun Noh 1, Richard S. Givens 2 1 Department of Otolaryngology University of Pittsburgh
More informationRX-AM4SF Receiver. Pin-out. Connections
RX-AM4SF Receiver The super-heterodyne receiver RX-AM4SF can provide a RSSI output indicating the amplitude of the received signal: this output can be used to create a field-strength meter capable to indicate
More informationHuman vs. Robotic Tactile Sensing: Detecting Lumps in Soft Tissue
Human vs. Robotic Tactile Sensing: Detecting Lumps in Soft Tissue James C. Gwilliam1,2, Zachary Pezzementi 1, Erica Jantho 1, Allison M. Okamura 1, and Steven Hsiao 2 1 Laboratory for Computational Sensing
More informationDiagrams and Graphs of Statistical Data
Diagrams and Graphs of Statistical Data One of the most effective and interesting alternative way in which a statistical data may be presented is through diagrams and graphs. There are several ways in
More informationIntroduction to Machine Learning and Data Mining. Prof. Dr. Igor Trajkovski trajkovski@nyus.edu.mk
Introduction to Machine Learning and Data Mining Prof. Dr. Igor Trakovski trakovski@nyus.edu.mk Neural Networks 2 Neural Networks Analogy to biological neural systems, the most robust learning systems
More informationAnatomy Review Graphics are used with permission of: adam.com (http://www.adam.com/) Benjamin Cummings Publishing Co (http://www.awl.com/bc).
Page 1. Introduction The structure of neurons reflects their function. One part of the cell receives incoming signals. Another part generates outgoing signals. Anatomy Review Graphics are used with permission
More informationPRODUCT SHEET OUT1 SPECIFICATIONS
OUT SERIES Headphones OUT2 BNC Output Adapter OUT1 High Fidelity Headphones OUT1A Ultra-Wide Frequency Response Headphones OUT3 see Stimulators OUT100 Monaural Headphone 40HP Monaural Headphones OUT101
More informationThree Channel Optical Incremental Encoder Modules Technical Data
Three Channel Optical Incremental Encoder Modules Technical Data HEDS-9040 HEDS-9140 Features Two Channel Quadrature Output with Index Pulse Resolution Up to 2000 CPR Counts Per Revolution Low Cost Easy
More informationClassical conditioning of the Aplysia siphon-withdrawal reflex exhibits response specificity
Proc. Nad. Acad. Sci. USA Vol. 86, pp. 762-7624, October 1989 Neurobiology Classical conditioning of the Aplysia siphon-withdrawal reflex exhibits response specificity (conditioned response/beta conditioning/activity-dependent
More informationPhysics 9e/Cutnell. correlated to the. College Board AP Physics 1 Course Objectives
Physics 9e/Cutnell correlated to the College Board AP Physics 1 Course Objectives Big Idea 1: Objects and systems have properties such as mass and charge. Systems may have internal structure. Enduring
More informationBiological Neurons and Neural Networks, Artificial Neurons
Biological Neurons and Neural Networks, Artificial Neurons Neural Computation : Lecture 2 John A. Bullinaria, 2015 1. Organization of the Nervous System and Brain 2. Brains versus Computers: Some Numbers
More informationHearing and Deafness 1. Anatomy & physiology
Hearing and Deafness 1. Anatomy & physiology Chris Darwin Web site for lectures, lecture notes and filtering lab: http://www.lifesci.susx.ac.uk/home/chris_darwin/ safari 1 Outer, middle & inner ear Capture;
More informationAS COMPETITION PAPER 2008
AS COMPETITION PAPER 28 Name School Town & County Total Mark/5 Time Allowed: One hour Attempt as many questions as you can. Write your answers on this question paper. Marks allocated for each question
More informationMITOCW MIT9_14S09_lec33-mp3
MITOCW MIT9_14S09_lec33-mp3 The following content is provided under a Creative Commons License. Your support will help MIT OpenCourseWare continue to offer high quality educational resources for free.
More informationhttp://abcnews.go.com/politics/video/obama-says-brain-initiative-will-be-transformative-18861944
http://abcnews.go.com/politics/video/obama-says-brain-initiative-will-be-transformative-18861944 What are the nervous system s functions? The nervous system organizes and controls an individual s appropriate
More informationBY S. S. STEVENS. Harvard University
A SCALE FOR THE MEASUREMENT OF A PSYCHOLOGICAL MAGNITUDE: LOUDNESS BY S. S. STEVENS Harvard University A scale adequate to the measurement of the subjective magnitude of sensation has long been sought
More informationStand Alone POTS Fiber Optic System. P31372 Station (Subscriber) Unit P31379 Remote (Exchanger) Unit. Description & Installation
Stand Alone POTS Fiber Optic System P31372 Station (Subscriber) Unit P31379 Remote (Exchanger) Unit Description & Installation Printed in USA 09/11 TO466 Rev. A Table of Contents Page 1.0 SCOPE 2 2.0 PRODUCT
More information2 Neuro-Anatomical Organisation of The Cortex. 3 A Generic Computational Model of a Patch of Cortex
Contents 1 Introduction 1.1 On the Cortex in General 2 1.2 Research Questions. 2 1.3 Thesis Outline. 3 2 Neuro-Anatomical Organisation of The Cortex 2.1 The Thalamus and the Spatio-Temporal Code... 4 2.2
More informationBrain Computer Interfaces (BCI) Communication Training of brain activity
Brain Computer Interfaces (BCI) Communication Training of brain activity Brain Computer Interfaces (BCI) picture rights: Gerwin Schalk, Wadsworth Center, NY Components of a Brain Computer Interface Applications
More informationIt s All in the Brain!
It s All in the Brain! Presented by: Mari Hubig, M.Ed. 0-3 Outreach Coordinator Educational Resource Center on Deafness What is the Brain? The brain is a muscle In order to grow and flourish, the brain
More informationElectrical Resonance
Electrical Resonance (R-L-C series circuit) APPARATUS 1. R-L-C Circuit board 2. Signal generator 3. Oscilloscope Tektronix TDS1002 with two sets of leads (see Introduction to the Oscilloscope ) INTRODUCTION
More informationNeurobiology of Learning and Memory
Neurobiology of Learning and Memory 94 (2010) 127 144 Contents lists available at ScienceDirect Neurobiology of Learning and Memory journal homepage: www.elsevier.com/locate/ynlme Remodeling the cortex
More informationName: Teacher: Olsen Hour:
Name: Teacher: Olsen Hour: The Nervous System: Part 1 Textbook p216-225 41 In all exercises, quizzes and tests in this class, always answer in your own words. That is the only way that you can show that
More informationChapter 3.8 & 6 Solutions
Chapter 3.8 & 6 Solutions P3.37. Prepare: We are asked to find period, speed and acceleration. Period and frequency are inverses according to Equation 3.26. To find speed we need to know the distance traveled
More informationTransverse Sections of the Spinal Cord
Transverse Sections of the Spinal Cord The spinal cord is perhaps the most simply arranged part of the CNS. Its basic structure, indicated in a schematic drawing of the eighth cervical segment (Figure
More informationNeurophysiology. 2.1 Equilibrium Potential
2 Neurophysiology 2.1 Equilibrium Potential An understanding of the concepts of electrical and chemical forces that act on ions, electrochemical equilibrium, and equilibrium potential is a powerful tool
More informationFunctional neuroimaging. Imaging brain function in real time (not just the structure of the brain).
Functional neuroimaging Imaging brain function in real time (not just the structure of the brain). The brain is bloody & electric Blood increase in neuronal activity increase in metabolic demand for glucose
More informationAuditory memory and cerebral reorganization in post-linguistically deaf adults
Auditory memory and cerebral reorganization in post-linguistically deaf adults Implications for cochlear implantation outcome D Lazard, HJ Lee, E Truy, AL Giraud Ecole Normale Supérieure, Inserm U960,
More informationSimulation Model of Mating Behavior in Flies
Simulation Model of Mating Behavior in Flies MEHMET KAYIM & AYKUT Ecological and Evolutionary Genetics Lab. Department of Biology, Middle East Technical University International Workshop on Hybrid Systems
More informationControlling the brain
out this leads to a very exciting research field with many challenges which need to be solved; both in the medical as well as in the electrical domain. Controlling the brain In this article an introduction
More informationWhat role does the nucleolus have in cell functioning? Glial cells
Nervous System Lab The nervous system of vertebrates can be divided into the central nervous system, which consists of the brain and spinal cord, and the peripheral nervous system, which contains nerves,
More informationStandards Alignment Minnesota Science Standards Alignment Matrix www.brainu.org/resources/mnstds
Lesson Summary: Neurons transfer information by releasing neurotransmitters across the synapse or space between neurons. Students model the chemical communication between pre-synaptic and post-synaptic
More informationModule 13 : Measurements on Fiber Optic Systems
Module 13 : Measurements on Fiber Optic Systems Lecture : Measurements on Fiber Optic Systems Objectives In this lecture you will learn the following Measurements on Fiber Optic Systems Attenuation (Loss)
More informationReading Materials Required readings: Psychology 3FA3 courseware (available in Campus Bookstore) Background readings Supplementary readings
Psych 3FA3: Neurobiology of Learning and Memory (2002-2003, Term 2) Instructor: Dr. Hongjin Sun, email: 3fa3@psychology.mcmaster.ca Office: Room 415, Psychology Building, Tel 905-525-9140, ext 24367 Lab:
More informationA model of memory, learning and recognition
A model of memory, learning and recognition Bruce Hoeneisen Universidad San Francisco de Quito 6 May 2002 Abstract We propose a simple model of recognition, short-term memory, longterm memory and learning.
More informationPassive Conduction - Cable Theory
Passive Conduction - Cable Theory October 7, 2013 Biological Structure Theoretical models describing propagation of synaptic potentials have evolved significantly over the past century. Synaptic potentials
More informationIntroduction to Psychology, 7th Edition, Rod Plotnik Module 3: Brain s Building Blocks. Module 3. Brain s Building Blocks
Module 3 Brain s Building Blocks Structure of the Brain Genes chains of chemicals that are arranged like rungs on a twisting ladder there are about 100,000 genes that contain chemical instructions that
More informationSimulation Infrastructure for Modeling Large Scale Neural Systems
Simulation Infrastructure for Modeling Large Scale Neural Systems Charles C. Peck, James Kozloski, A. Ravishankar Rao, and Guillermo A. Cecchi IBM T.J. Watson Research Center P.O. Box 218 Yorktown Heights,
More informationCLINICAL NEUROPHYSIOLOGY
CLINICAL NEUROPHYSIOLOGY Barry S. Oken, MD, Carter D. Wray MD Objectives: 1. Know the role of EMG/NCS in evaluating nerve and muscle function 2. Recognize common EEG findings and their significance 3.
More informationFDTD Analysis of Site Free Space VSWR in Test Site Used for Disturbance Measurement above 1 GHz
九 州 工 業 大 学 学 術 機 関 リポジトリ Title FDTD analysis of site free space VS disturbance measurement above 1 GHz Author(s) Kuwabara N; Midori M; Kawabata M Issue Date 9-8 URL http://hdl.handle.net/8/368 9 IEEE.
More informationComputation by Ensemble Synchronization in Recurrent Networks with Synaptic Depression
Journal of Computational euroscience 13, 111 124, 2002 c 2002 Kluwer Academic Publishers. Manufactured in The etherlands. Computation by Ensemble Synchronization in Recurrent etworks with Synaptic Depression
More informationMUSIC RECOGNITION DIANA DEUTSCH. Psychological Review, 1969, 76, 300 307. University of California, San Diego
Psychological Review, 1969, 76, 300 307 MUSIC RECOGNITION DIANA DEUTSCH University of California, San Diego Music recognition is discussed, and it is argued that this involves certain specific processes
More informationWhat is the basic component of the brain and spinal cord communication system?
EXPLORING PSYCHOLOGY David Myers The Biology of Mind Chapter 2 Neural Communication Neurons How Neurons Communicate How Neurotransmitters Influence Us The Nervous System The Peripheral Nervous System The
More informationPyramidal neurons: dendritic structure and synaptic integration
Pyramidal neurons: dendritic structure and synaptic integration Nelson Spruston Abstract Pyramidal neurons are characterized by their distinct apical and basal dendritic trees and the pyramidal shape of
More information