340 R.M. SHAPLEY AND C.

ENROTH-CUGELL

CBo1

~a 2 ~ ~ All

IPL
b

~F~ G ON-Beta G

Ill,oN
FIG. 63. Diagram of connections in the inner plexiform layer of the cat retina from serial EM reconstructions. Chemical
synapses are indicated by the arrows: gap junctions are indicated by the symbol -I . CB indicates a cone bipolar cell; four
subtypes are labeled. RBP stands for rod bipolar cell. A l l is a subclass of amacrine cells. The Beta ganglion cells are presumed
to be the anatomical equivalent of the X cells. The inner plexiform layer is subdivided into sublaminae-a and -b, as indicated
in the figure. On ganglion cells receive input in sublamina-b, while off ganglion cells receive input in sublamina-a. From
Sterling (1983).
minate type. Then the interplexiform cell makes inputs, is offered in Fig. 63, from a recent review
synapses on all types of bipolar cells at the level of article (Sterling, 1983).
the IPL but more especially at the level of the outer
plexiform layer (OPL). It is the perfect example of
a feedback neuron. Moreover, it could affect all A P P E N D I X 2 - - R E C E P T I V E FIELDS
bipolars which is a property one would want from
a gain control since on- and off-pathways must be The neural signal which leaves the retina consists
adapted in the same way in order to keep contrast of trains of impulses carried by the axons of retinal
sensitivity constant. However, the wide terminal ganglion cells. The response of these cells to a visual
branching of the interplexiform cell in the OPL is stimulus may be defined as a change in the rate of
a problem if this cell is to play a role in regulating firing of impulses. An adequate stimulus to cause
gain on steady backgrounds, because, at least for such a change in firing is some change in the
X cells, the area of the retina over which adaptive illumination on the retina. The work of Hartline
signals are summed for the center is less than 1 deg 2, (1940), Barlow (1953), and Kuffler (1952, 1953)
i.e. less than 200 /a by 200/a. However, there is showed that each retinal ganglion cell generated
evidence for a different kind of gain control which responses, as defined above, to stimulation over a
depends on stimulus contrast rather than average limited area of the retina, and this area was defined
flux (see Section 3.8; Shapley and Victor, 1978; cf. as the receptive field of that ganglion cell. Working
Werblin and Copenhagen, 1974) and perhaps the in the cat retina, Kuffler (1953) found that ganglion
interplexiform cell's properties may match those of cell receptive fields consisted of two concentric
this other gain control. zones which he called the center and surround. The
A schematic diagram of the neuronal circuitry in center and surround were mutually antagonistic. In
the cat's IPL, where ganglion cells receive their on-center cells in which the center caused excitatory