OPTO30007:

: Visual Cortex 1 -
Cell types and orienta;on selec;vity
Kandel et al., 2000: Chap. 27; Nicholl s et al., Chap. 20.

The Visual Pathway

www.owlnet.rice.edu/ ~psyc351/imagelist.htm

Ventro-medial surface of occipital and temporal lobes (Human)

Calcarine sulcus
Central vision

upper bank

lower bank

Lower Visual Field

Upper Visual Field

V1 damage

Macaque visual areas (Van Essen, 1991)

There are over 30 different visual areas beyond the striate cortex (V1) in the
macaque brain. There is also a reasonable degree of functional localisation
among these areas.

Early visual corAcal areas

Visual areas are disAnguished

from each other on the basis of
cyto- and myelo- architectonics
and physiological studies.

Striate cortex (V1) is easily
idenAed because of its
prominent layer 4b, the stria of
Gennari.

Primary
Visual
cortex
V1, Area
17 or
Striate
cortex

General plan of topographical representa;on in

the cortex:
Ver;cal
meridian

B
B

V2

V1

Horizontal
meridian
V1/V2 border is represented by
the verAcal meridian.

Visual CorAcal anatomy

Layers (laminae) of
macaque primary visual
cortex, also known as
striate, V1 or area 17.
(Hubel & Wiesel, 1972)

Pyramidal and stellate

cells from cat cortex,
stained by the Golgi
technique
(Cajal, 1955)

Cells stained by HRP

aXer physiological
idenAcaAon
(Gilbert & Wiesel,
1979)

Nicholls et al, From Neuron to Brain, 2001, p. 417

Visual CorAcal anatomy

The layering paZern of the cortex is the result of variaAons in the
staining and packing density of these cells.

Layers 4C and 6 are densest and darkest
Layers 1, 4B, and 5 are most loosely packed. Layer 1 contains hardly
any nerve cells but has abundant axons, dendrites, and synapses.
To show that dierent layers contain dierent kinds of cells requires a
stain like that devised by Camillo Golgi in 1900. The Golgi stain reveals
only occasional cells, but when it does reveal a cell, it may show it
completely, including its axons and dendrites. The two major classes of
corAcal cells are the pyramidal cells, which occur in all layers except 1
and 4 and the stellate cells, which are found in all layers.
Pyramidal cells: large; dendrites radiate from the base; major axon
leaves the corAcal area.
Interneurones: local projecAons stellate: spiny (glutamatergic) and
non-spiny or smooth (GABAergic); Chandelier, double-bouquet, basket..

TerminaAon zones of geniculate aerents in

macaque V1

P cells from LGN project mainly to layers 4C

and 4A, with a sparse projecAon to layer 6.

M cells project mainly to layer 4C, with a
sparse projecAon to layer 6.

K cells project to layers 1 and 3.

Casagrande & Kaas, 1994

Birth of modern visual physiology

The discoveries of Hubel
and Wiesel represent a
break-through in research
into the ability of the brain
to interpret the code of
the impulse message from
the eyes. Thanks to their
invesAgaAons we now
have a deeper insight into
informaAon analysis
within the visual system
and into the processes
forming the basis for the
origin of the visual
impression.
- Press Release from the
Nobel Assembly,
announcing the 1981
Nobel Prize for Medicine &
Physiology

Responses of a simple cell

Nicholls et al, From Neuron to Brain, 2001, p. 417

Responses of a Simple cell in the primary visual

cortex

Responses of a complex cell

Nicholls et al, From Neuron to Brain, 2001, p. 420

Simple cell

Complex Cell

Separate On & O subregions Overlapping On & O

in recep;ve eld
subregions
Small recepAve eld

Larger recepAve eld

Linear spaAal summaAon (X

like)

Non-linear properAes (Y like)

Found in layers 4 and 6

Mostly outside layer 4

Diuse light ineecAve as sAmulus

Most are orientaAon selecAve
Some may be direcAon selecAvity
Some exhibit end-inhibiAon
Many are binocularly driven

Hypercomplex cells: Example of end-stopping

INHIBITORY FLANKS

Poor
GoodResponse
Response
End-stopping is either a local phenomenon (iniAated by layer 6 pyramids) or a
property carried over from the dLGN.

Hubel & Wiesel: Model for simple cell orientaAon

selecAvity (1962)

Kandel et al, Principles of Neural science, 2000, p. 535

Hubel & Wiesel: Model for complex cell

Kandel et al, Principles of Neural science, 2000, p. 536

Hubel & Wiesel: Model for hypercomplex cell

Nicholls et al, From Neuron to Brain, 2001, p. 421

Summary of Hubel & Wiesels model of hierarchy

for Primary Visual Cortex:

Hubel & Wiesels model for Simple Cell: Convergent

excitatory connecAons from a number of LGN cells whose

recepAve elds are in a row in visual space.

Hubel & Wiesels model for Complex Cell: Convergent

excitatory input from a number of simple cells, all tuned to the

same orientaAon.

Hubel & Wiesels model for Hypercomplex Cell: Excitatory

input from a complex cell in the centre and inhibitory inputs in

the inhibitrory end zones.

Many of Hubel & Wiesels ideas have stood the test

of >me, but some have not:
1. Model of orientaAon selecAvity - sAll hotly debated.
(Vidyasagar et al., Trends Neurosci., 1996; Sampolinsky &
Shapely, Curr Opin Neurobiol., 1997; Carandini, Neuron,
2007)
E.g. Removal of inhibiAon on a striate cell (by antagonists of the
inhibitory transmiZer, GABA) reduces or abolishes its
orientaAon selecAvity (e.g., Sillito et al., Brain Res., 1980).

2. The hierarchy scheme: only partly true.

E.g., (i) Complex cells oXen respond to sAmuli that simple cells
do not! (Hammond & MacKay, Exp. Brain Res., 1977).
(ii) Both complex cells and hypercomplex cells can be
monosynapAcally excited from the LGN. (Bullier & Henry, J.
Neurophysiol., 1979).

While originally it was believed

that orientaAon selecAvity is an
emergent property at the corAcal
level, now there is a lot of
evidence that varying degrees of
orientaAon selecAvity (referred to
as orientaAon biases ) are
present subcorAcally in the
reAna (Levick & Thibos, Nature,
1980) and dLGN (Vidyasagar &
Urbas, Exp. Brain Res., 1982). The
dLGN cell on the right shows
responses that are dependent on
the orientaAon of the bar
sAmulus. The quesAon is : Are
they important for corAcal
orientaAon selecAvity?

OrientaAon selecAvity
may arise before the
level of striate cortex!

Vidyasagar & Urbas, Exp. Brain Res., 1982

Dierent schemes proposed to explain striate orientaAon selecAvity

Hubel & Wiesel
(1962, 1968)

Creutzfeldt,
Kuhnt &
Benevento, 1974


Heggelund, 1981

Vidyasagar, Pei
& Volgushev,
1996

Learning objectives!
"
By the end of this lecture you should be able to:"
"
describe the basic anatomical characteristics of the primary visual cortex, the
termination zones of LGN afferents and the morphological cell types in area V1. "
"
understand the basic retinotopy of area V1 and describe the visual loss from focal
V1 damage"
"
understand the distinguishing features of simple, complex and hypercomplex
cells"
understand the model of orientation selectivity proposed by Hubel & Wiesel and
describe briefly some alternative schemes."
"