Psychology 832: Physiological Psychology - Neural models of cortical memory function.

Spring 2000, Tuesday, 12:30 — 2:30 pm •

Dr. Michael Hasselmo

2 Cummington St. Rm. 105E

Voice: 353-1397


Office Hours: Mon.& Wed, 4:00 — 5:00 pm




How Much


Class Participation


5% of grade



May 14

30% of grade




30% of grade



March 10

5% of grade



April 21

30% of grade


• The EXAM will contain questions on the relation between neural network models and real neural systems, and on the basic computations involved in associative memories.

• The PRESENTATION will involve preparing for the topic presented during one week of the term. Students will provide a highly simplified overview of one of the modeling papers listed on the syllabus.

• The PROJECT OUTLINE will give an overview of the final project, with a list of references and an introduction to a specific research question.

• The FINAL PROJECT will allow individual research on a particular subject. The nature of the project will depend upon the student’s background. Those with programming experience can do a programming project; those without can explore available software or do an extensive literature review.


This course covers computational models of neurobiological mechanisms for memory function and spatial navigation, with a particular emphasis on cellular and circuit models of the hippocampus and related cortical structures.


Hippocampus Volume 6, number 6 Special Issue: Computational models of hippocampal function in memory. Wiley-Liss: New York, 1996.

Koch, C. and Segev, I. (eds.) (1998) Methods in Neuronal Modeling: From Ions to Networks Cambridge, MA: MIT Press.

Supplemental reading

Bower, J.M. & Beeman, D. (1997). The Book of GENESIS: Exploring Realistic Neural Models with the GEneral NEural SImulation System. 2nd edition. New York: TELOS/Springer-Verlag.

O'Reilly, R.C., Munakata, Y. and McClelland, J.L. (eds.) (1999) Cognitive Neuroscience: A computational exploration. Cambridge, MA: MIT Press. (In press)


Jan. 11 Introduction

Course logistics. History and overview of research on models of cortical circuits involved in memory, including hippocampus and related structures.

Models of Cellular Physiology Involved in Memory

Jan. 18 Modification of intrinsic properties in hippocampal pyramidal cells.

Covers modifiable properties of pyramidal cells important for memory function in local circuits, including spike generation, afterhyperpolarization potentials and bursting properties.

Traub RD, Wong RK, Miles R, Michelson H (1991) A model of a CA3 hippocampal pyramidal neuron incorporating voltage-clamp data on intrinsic conductances. J. Neurophysiol. 66: 635-650.

Blackwell KT, Vogl TP, Alkon DL (1998) Pattern matching in a model of dendritic spines. Network 9:107-121.

Book of GENESIS, Chapter 7.

Jan. 25 Models of short term synaptic plasticity.

Covers models of paired pulse facilitation, depression and post-tetanic potentiation.

Tsodyks M, Pawelzik K, Markram H (1998) Neural networks with dynamic synapses. Neural Comput. 10:821-835.

Abbott LF, Varela JA, Sen K, Nelson SB (1997) Synaptic depression and cortical gain control. Science 275: 220-224.

Grossberg S (1984) Some normal and abnormal behavioral syndromes due to transmitter gating of opponent processes. Biol. Psychiatry 19: 1075-1118.

Feb. 1 Models of long-term potentation and the NMDA receptor

Role of NMDA receptor in mediating long-term synaptic modification.

Brown TH, Kairiss EW, Keenan CL (1990) Hebbian synapses: biophysical mechanisms and algorithms. Annu Rev Neurosci 1990;13:475-511

Levy WB, Steward O (1983) Temporal contiguity requirements for long-term associative

potentiation/depression in the hippocampus. Neuroscience 1983 Apr;8(4):791-7

Models of Hippocampal Episodic Memory Function

Feb. 8 LTP and memory storage in region CA3.

McNaughton, B.L. & Morris, R.G.M. (1987). Hippocampal synaptic enhancement and information storage within a distributed memory system. Trends in Neuroscience, 10, pp. 408—415.

Hasselmo, M.E., Schnell, E., & Barkai, E. (1995). Dynamics of learning and recall at excitatory recurrent synapses and cholinergic modulation in hippocampal region CA3. Journal of Neuroscience, 15, pp. 6259—6274.

Rolls, E.T. (1996) A theory of hippocampal function in memory. Hippocampus 6: 601-620.



Feb. 15 Cholinergic innervation from the septum and encoding.

Hasselmo, M.E. & Bower, J.M. (1993). Acetylcholine and memory. Trends inNeurosciences, 16, pp. 218—222.

Hasselmo, M.E., Wyble, B.P. and Wallenstein, G.V. (1996) Encoding and retrieval of episodic memories: Role of cholinergic and GABAergic modulation in the hippocampus. Hippocampus 6: 693-708.

Models of Spatial Navigation

Feb. 22 Pathways stored as sequences of activity.

Levy, W.B. (1996) A sequence predicting CA3 is a flexible associator that learns and uses context to solve hippocampal-like tasks. Hippocampus 6: 579-590.

Wallenstein, G.V. and Hasselmo, M.E. (1997) GABAergic modulation of hippocampal activity: Sequence learning, place field development, and the phase precession effect. J. Neurophysiol. 78: 393-408.

March 1 Navigation guided by two-dimensional representations.

Redish AD, Touretzky DS The role of the hippocampus in solving the Morris water maze. Neural Comp. 1998 527 10, 73-111.

Gerstner W, Abbott LF Learning navigational maps through potentiation and modulation of hippocampal place cells. J Comput Neurosci 1997, 4, 79-94

Muller, R.U. and Stead, M. (1996) Hippocampal place cells connected by Hebbian synapses can solve spatial problems. Hippocampus 6: 709-719.

Recce, M. and Harris, K.D. (1996) Memory for places: A navigational model in support of Marr's theory of hippocampal function. Hippocampus 6: 735-748.

March 8 Integration of place cells and head direction cells.

Burgess, H. and O'Keefe, J. (1996) Neuronal computations underlying the firing of place cells and their role in navigation. Hippocampus 6: 749-762.

Sharp, P.E., Blair, H.T. and Brown, M. (1996) Neural network modeling of the hippocampal formation spatial signals and their possible role in navigation: a modular approach. Hippocampus 6: 720-734.

March 10 Integration of place cells and head direction cells.

EEG Phenomena in vivo and in vitro

March 15 Oscillatory dynamics of neuronal membranes and local circuits.

Fransen, E., Wallenstein, G.V., Alonso, A., Dickson, C.T. and Hasselmo, M.E. (1998) A biophysical simulation of intrinsic and network properties of entorhinal cortex. In: J.M. Bower (ed.) Neurocomputing: Proceedings of the Computational Neuroscience Conference, Elsevier,

Rinzel, J. and Ermentrout, B. (1998) Analysis of neural excitability and oscillations. In: Koch, C. and Segev, I. (eds.) (1998) Methods in Neuronal Modeling: From Ions to Networks Cambridge, MA: MIT Press, pp. 251-292.

Abbott, L. and Marder, E. (1998) Modeling small networks. In: Koch, C. and Segev, I. (eds.) (1998) Methods in Neuronal Modeling: From Ions to Networks Cambridge, MA: MIT Press. pp. 361-410.

March 22 Oscillatory dynamics in slice preparations of hippocampus - Nancy Kopell.

J. White, C. Chow, J. Ritt, C. Soto-Trevino and N. Kopell, (1998) Synchronization and oscillatory dynamics in heterogeneous, mutually inhibited neurons,'' J. Comput. Neurosci. 5: 5-16

G.B. Ermentrout and N. Kopell, (1998) Fine structure of neural spiking and synchronization in the presence of conduction delays, Proc. Nat. Acad. Sci., USA 95: 1259-1264.

C. Chow, J. White, J. Ritt, C. Soto-Trevino and N. Kopell, (1998) Frequency control in synchronous networks of inhibitory neurons", J. Comput. Neurosci., 5: 407-420.

March 29 Theta rhythm, phase precession and sequence retrieval

Jensen O. and Lisman JE Hippocampal CA3 region predict memory sequences: accounting for the phase advance of place cells. Learning and Memory, 1996, 3, 433 279-287.

Wallenstein, G.V. and Hasselmo, M.E. (1997) GABAergic modulation of hippocampal activity: Sequence learning, place field development, and the phase precession effect. J. Neurophysiol. 78: 393-408.

Models of Conditioning

April 5 Models of adaptive timing phenomena.

Grossberg, S. and Merrill, J.W.L. (1996) The hippocampus and cerebellum in adaptively timed learning, recognition and movement. J. Cog. Neurosci. 8: 257-277.

Grossberg, S. and Merrill, J.W.L. (1992) A neural network model of adaptively timed reinforcement learning and hippocampal dynamics. Cog. Brain Res. 1:3-38.

Grossberg, S. and Schmajuk, N.A. (1987) Neural dynamics of attentionally modulated Pavlovian conditioning: Conditioned reinforcement, inhibition and opponent processing. Psychobiol. 15: 195-249.

Fiala JC, Grossberg S, Bullock D (1996) Metabotropic glutamate receptor activation in cerebellar Purkinje cells as substrate for adaptive timing of the classically conditioned eye-blink response. J. Neurosci. 16: 3760-3774.

April 12 Other classical conditioning models.

Gluck, M.E. and Myers, C.E. (1996) Integrating behavioral and physiological models of hippocampal function. Hippocampus 6: 643-653.

Buhusi, C.V. and Schmajuk, N.A. (1996) Attention, configuration, and hippocampal function. Hippocampus 6: 621-642.

April 21 Research project due.

Interaction with neocortical structures

April 19 Hippocampal regulation of neocortical representations.

Shen, B. and McNaughton, B.L. (1996) Modeling the spontaneous reactivation of experience-specific hippocampal cell assemblies during sleep. Hippocampus 6: 685-692.

McClelland, J.L. and Goddard, N.H. (1996) Considerations arising from a complementary learning systems perspective on hippocampus and neocortex. Hippocampus 6: 654-665

Carpenter, G. & Grossberg, S. (1993). Normal and amnesic learning, recognition and memory by a neural model of cortico-hippocampal interactions. Trends inNeuroscience, 16, pp. 131-137.

April 26 Models of parietal cortex spatial representations

Zipser, D. & Anderson, R.A. (1988). A back-propagation programmed network that simulates response properties of a subset of posterior parietal neurons. Nature, 331, pp. 679—684.

Pouget A, Sejnowski TJ (1994) A neural model of the cortical representation of egocentric distance. Cereb. Cortex 4: 314-329.


McNaughton BL, Mizumori SJ, Barnes CA, Leonard BJ, Marquis M, Green EJ (1994) Cortical representation of motion during unrestrained spatial navigation in the rat. Cereb. Cortex 4: 27-39.

May 5 Models of prefrontal cortex working memory function.

Lisman, J., Fellous, J.-M. and Wang, X.-J. (1998) A role for NMDA channels in working memory. Nature Neurosci. 1: 273-275.

Moody SL, Wise SP, di Pellegrino G, Zipser D (1998) A model that accounts for activity in primate frontal cortex during a delayed matching-to-sample task. J. Neurosci. 18: 399-410.

Durstewitz D, Kelc M, Gunturkun O (1999) A neurocomputational theory of the dopaminergic modulation of working memory functions. J. Neurosci. 19: 2807-2822.