THE FIRING OF

PLACE CELLS IN
THE DARK

Anatomical location of Brocas area.

Paul Broca

Brains of Leborgne(top)
and Lelong(bottom).

There are in the human mind a group of faculties and in the brain groups of convolutions, and
the facts assembled by science so far allow to state, as I said before, that the great regions of the
mind correspond to the great regions of the brain.

Egaz Moniz

Prefrontal Lobotomy

Procedure(top),changes(bottom)

Normal psychic life depends upon the good functioning of brain synapses,
and mental disorders appear as a result of synaptic derangements.
Synaptic adjustments will then modify the corresponding ideas and force
them into different channels. Using this approach we obtain cures and

Penfield opines Cerebral Cortex to be the

site of Implicit Memory.

Wilder Penfield

William Beecher Scoville

Milner

Brenda

H.M, the amnesiac we shall never forget

sheds clues on memory, their distinction and location .

Hippocampus- Anatomy

Rat Brain-Lateral View

Arrangement of Neurons In
Hippocampus

The left and Right Hippocampi are

dissimilar behaviorally.

Left
Right

Single-cell Recording in the Hippocampal

Formation Reveals Two Major Classes of Units:
Principal Cells and Theta Cells.

Okeefe and Ranck (1973) let rats explore a certain environment while they recorded
activity of individual neurons in CA1 region known as Single Units.
They noticed that there were two major classes of cells :complex-spike and theta cells.
Ranck had trained his animals to approach one location to obtain food and another to get water,
and emphasized the relation of the complex- Spike cell firing pattern to the behavioral approach
to reward. OKeefe was more impressed by the spatial correlate and named the cells place cells.
The second class of neurons, the theta cells, has less specific behavioral correlates. As the name
implies, their behavioral correlates are closely related to those of the gross EEG waves and in
particular theta.
It is highly likely that in the rat the complex spike cells are pyramidal cells and the theta cells
are one or more types of interneuron. Intracellular staining of neurons that display complex
spikes in brain slices reveals they have the morphology of pyramidal cells, whereas those without
complex spikes are interneurons.

Complex-spike cells and theta cells have different physiological properties.

A. Theta cells have a steady firing rate and constant size amplitude spikes,
whereas C-S cells sometimes emit a complex-spike burst in which the later
action potentials in the burst are lower in amplitude and broader.

*Firing field of a CA1 place cell.

A. Raw data from a rat foraging
for food in a square box for 10
minutes. Gray line traces the
animals path through the
environment; black boxes show
firing of place cell. B. Place firing
field as a grayscale map where
darker colors represent higher
firing rates. Inset number in

 Firing fields of 32 place cells simultaneously recorded while a rat foraged for food in a 62-cm2
box. The place field maps are arranged topographically so fields in the northwest of the box
are located at the upper left, fields in the southwest are located at the lower left, and so on.
In reality there is no topographical relation between the location of cells in the hippocampus
and the location of their fields in an environment. ( Source: The Hippocampus Book)

A place is defined as a location marked by no

locally
perceptible cues and is therefore only recognizable
relative to
other perceptible landmarks or orienting gradients.

Long-Lasting Working Memories of Obstacles Established by Foreleg

Stepping in
Walking Cats Require Area 5 of the Posterior Parietal Cortex.
David A. McVea,Andrew J. Taylor, andKeir G. Pearson.

ttttt.mp4

tttt2.mp4

Grid Cells and companions.

Discovered in 2005 by May- Britt Moser and Edvard Moser when they ut it
was unclear whether those place signals originated in the hippocampus
itself or came from the outside. To address this question, we made intrahippocampal lesions that disconnected the output stage of the circuit CA1
from the earlier stages. To our surprise, this did not abolish place coding
in CA1.
At first they noticed that many entorhinal cells spiked every time a rat
went to a particular spot, like the place cells in the hippocampus. However,
each cell had multiple firing locations and those firing locations formed a
peculiarly regular pattern a hexagonal grid much like the arrangement
of marbles in a Chinese checker board. Every cell did it this way, with
actual firing locations differing between cells. The cells were organized
topographically in the sense that the size of and distance between grid
fields increased from dorsal to ventral.

a) Stepwise increase of grid spacing at successive dorsoventral levels of medial entorhinal cortex. Spatial
autocorrelograms for four example cells (one per dorsoventral module). (b) Remapping of hippocampal place
cells in two environments. Top panels show responses of three grid cells to a change in the environment.
Independent responses are illustrated by different degrees of rotation and translation. Bottom panels show
inputs from the grid cells, each from a different module (purple, green and orange), at different locations in the
environments. The three grid cells provide input to a particular CA3 cell (white spot at left) sufficient to cause it
to fire when, and only when, the nodes of the three grids overlap. This occurs only at one location in this
example. In the second environment, the altered coactivity of the grid cells activates a different subset of place
cells at each location, and global remapping is observed in hippocampal place ensembles.

Source:Memory, navigation and theta rhythm in the hippocampal-entorhinal

system.
Buzsaki and E. Moser, 2013

Entorhinal cortex grid cells.

Raw data (left) and firing fields (right)
for three grid cells. Note the regularity of
the firing, which forms a triangular grid
pattern in each cell. Numbers indicate the
peak rate. Peaks of the three cells (top
in grayscale, bottom as numbers 13)
are slightly offset from each other so when
superimposed they tend to tesselate the
space. (Source: After Hafting et al., 2005,
by permission.

Four spatial cell types in the hippocampal formation.

A. Place cell has single localized
field and no directional selectivity.
B. Place by directional cell has
single localized field and strong directional selectivity.
C. Head
direction cell does not have localized firing but has strong directional
selectivity.
D. Grid cell has multiple place fields and no directional selectivity.
Directional grid cells also exist.
(Source: Edvard and May-Britt Moser.)

Border cells express proximity to

boundaries
in a number of environmental configurations

The Firing of Hippocampal Place

Cells in the Dark Depends on the
Rats Recent Experience.
Gregory J. Quirk, Robert U. Muller, and John L.Kubie,
1998.

THE HERO

THE
SETUP

Arena

81cm

81cm

51cm

51cm

Four hippocampal place cells tested in the light-dark-light sequence. The 3 firing rate maps in each row
correspond to the 3 segments
of the L-D-L sequence. Cells A and B show strong persistence of spatial firing in the dark in the square and cylinder
chambers, respectively. Cell
C weakly persisted, and cell D failed to persist. The persistence scores between the maps are as follows: cell A, Al:A2
= 0.57, Al:A3 = 0.68, A2:

Histograms showing the distributions of the persistence scores

between the initial lighted segment and (A) the dark segment of the
L-D-L sequence(B, ) the light segmenotf the L-D-L sequencea,n d( c)
the dark segment of the D-L sequence. The triangle at 0.14 on the
abscissa marks the cut-off for significance at the 0.001 level.

Three examples of cells whose firing patterns did not persist when the rat was placed directly into the
darkened chamber.

 Firing rate map for a lighted, 8 min recording session that was run subsequent to
the D-L sequences for the cell shown in Figure 3C. This cell, which showed a firing
field at 4 oclock in the cylinder in the L-D-L sequence, stopped firing when the rat
was put into the darkened chamber (Fig. 3). The map above shows that the firingfield returned in a subsequent lighted session. Median rates of pixels are as
follows (order: yellow, orange, red, green, blue, purple, in Hz.): 0.0, 1 .0, 3.3, 6.2,
10.0, 13.8.

Spatial Firing of Hippocampal Place Cells in Blind Rats

Etienne Save, Arnaud Cressant, Catherine Thinus-Blanc, and Bruno Poucet

Contrary to place cells in sighted rats, no cell in blind rats was observed to fire
in the firing field if the rat had not made physical contact with an object
previously.
In many cells recorded from blind rats, knowledge of one landmark position
was enough to activate firing in the place field. This confirms that, to produce
coherent firing, the hippocampal place cell system needs information about the
location of objects. This result additionally suggests that the system is able to use
the intrinsic (e.g., olfactory, tactile) properties of objects to recognize
which object the animal has encountered.
Once landmark positions are known, the place cell system relies on the dynamic use
of internally generated, motion-related information to update the position of the
system throughout the environment.
Such information includes kinesthetic signals from the vestibular system,
proprioceptive cues, and motor reference copy signals. Although motion-related
signals are known to accumulate errors across successive movements in space, such
errors can be corrected at each contact with an object by using the fixed
object locations as a means for recalibrating a calculated position.
At any rate, our study suggests that the spatial impairments of blind animals, if any,
are not the consequence of a decreased ability of the hippocampal place cell system