64 65

CHAPTER THREE Neural Transmission

To oscilloscope To oscilloscope

1, Two simultaneous EPSP's sum to produce a greater EPSP

A Stimulated „„ i B Stimulated „ A + B Stimulated
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-65h
-70 -70

2. Two simultaneous IPSP'ssum to produce a greater IPSP -70

C Stimulated D Stimulated C + D Stimulated
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-70 A A A
o
a -75 1. Two EPSP's elicited in rapid succession sum to
produce a larger EPSR

3. A simultaneous IPSP and EPSP cancel each other out

A Stimulated C Stimulated A + C Stimulated
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B B B
FIGURES.? The time
2. Two IPSP's elicited in rapid succession sum to
combinations of spatial
produce a larger IPSR FIGURE 3.8 The two
summation.
combinations of temporal
summation.

twice in rapid succession can produce a greater IPSP than that produced by
a single stimulation at the same intensity.
It is important not to be misled by the way in which neural integration is
studied in the laboratory. Spatial summation is studied independently of
temporal summation by administering two discrete stimuli to different
sites at exactly the same time, and temporal summation is studied indepen-
dently of spatial summation by stimulating one site twice in rapid succes-
sion. However, the normal operating conditions of most neurons are quite
different; most neurons continuously integrate signals over both time and
space as the neuron is continually bombarded with stimuli through the
thousands of synapses covering its dendrites and cell body. Remember
that although schematic diagrams1 of neural circuitry rarely include neu-
rons with more than a few representative synaptic contacts, the real situa-
tion is much more complex. For example, each mammalian motor neuron
receives about 10,000 synaptic contacts.
The location of a synapse on a. receptive membrane is an important
factor in determining its potential to influence neural firing. Because EPSPs
and IPSPs are transmitted decrementally, those synapses near the hillock
trigger zone have the most influence on the firing of the neuron. This may
explain why synaptic contacts tend to be segregated, that is, why synaptic
contacts from a particular source or from a particular class of neurons tend
to be located on a particular part of the target neuron. For example, axons
from one part of the brain might all synapse on the cell body of the target
neuron, whereas those from another area might favor dendritic target
sites.
In some ways the firing of a neuron is like the firing of a gun. Both are
all-or-none reactions triggered by their own graded responses to stimula-
tion. As a trigger is squeezed, it responds by gradually moving back until it
causes the gun to fire. Similarly, as depolarizing stimulation of gradually
increasing intensity is applied to a neuron, the neuron becomes less and
less polarized until the threshold of excitation is reached, and the neuron
fires. And like the firing of a gun, neural firing is an all-or-none event; just