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lobe learning task is the paired-associates task, in which certain
objects are arbitrarily paired with other objects, for example an
airplane with a soccer ball. When the monkeys are shown the airplane
as the sample stimulus, they must select the soccer ball from among
other possible test objects. Several studies have shown that neu-
ronal activity in the temporal lobe correlates with paired-associate
learning ( 221-223 ).
7.5. Frontal Cortex:
Primary Motor Cortex
Numerous tests of basic motor control or motor learning can be
used to assess damage to motor cortex and the resulting effects on
motor responses, such as reaching, grasping, and locomotion.
These tests include reaching to different locations with or without
applied force fi elds and attempting to trace different patterns on a
display ( 224-227 ). Grasping can be assessed by having subjects
attempt to grab and retrieve various items in order to receive
reward; test items are created that require different types of grips,
such as precision and power grips; see ref. ( 228 ). Tests of locomo-
tion include the assessment of motor responses coordination in
freely moving monkeys or those moving on a treadmill; see refs.
( 206, 229, 230 ).
Premotor cortical damage can give rise to defi cits in basic motor
control as outlined in the previous section. However, there are
other tasks that have been shown to engage premotor cortex in
particular, such as the conditional motor learning task ( 231 ). In this
task, subjects must learn to associate a particular arbitrary stimulus,
e.g., a red square, with a particular response, e.g., left response.
The stimuli are typically displayed in the center of the display, one
per trial, and thus provide no spatial information themselves.
Instead, an arbitrary association must be made between a symbolic
cue and a response. It has been shown that neurons in premotor
cortex underlie this ability ( 231, 232 ). Thus, damage to premotor
cortex should cause defi cits in conditional motor learning, which
can be characterized using this task.
7.6. Frontal Cortex:
Premotor Cortex
Supplementary motor cortex appears to be particularly involved in
cases in which sequences of motor responses are required, for
instance when people are playing the piano. Monkeys can be trained
to perform sequences of responses, such as touching a series of
stimuli in a specifi ed sequence or making a series of responses, such
as turning a knob, opening a door, and retrieving a food reward
( 206, 233, 234 ). Potential damage to supplementary motor cortex
can then be assessed by determining the resulting defi cits on these
sequencing tasks.
7.7. Frontal Cortex:
Supplementary Motor
7.8. Prefrontal Cortex
Although controversy remains over how best to characterize
prefrontal cortical function, converging evidence points to certain
key functions of this reagion. For example, a great deal of work
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