T. Klingberg et Pe. Roland, RIGHT PREFRONTAL ACTIVATION DURING ENCODING, BUT NOT DURING RETRIEVAL, IN A NONVERBAL PAIRED-ASSOCIATES TASK, Cerebral cortex, 8(1), 1998, pp. 73-79
Brain imaging studies have shown that episodic encoding into long-term
memory preferentially activates the left prefrontal cortex and retrie
val activates the right prefrontal cortex. However, it is unclear to w
hat degree verbal analysis contributes to the left prefrontal activati
on during encoding. The present study was designed to avoid verbal ana
lysis during encoding by using abstract pictures and computer-generate
d sounds which were difficult to code verbally. Sounds and pictures we
re grouped into six stimulus-stimulus pairs. When the sound from a pai
r was presented, the subjects were instructed to recall and visualize
the associated picture. After 2.0 s the associated picture and another
picture appeared on the screen and the subjects were required to iden
tify the associated picture. Feedback about the choice was then given.
Regional cerebral blood flow (rCBF) was measured with [O-15]butanol a
nd positron emission tomography (PET) in 10 subjects during initial tr
aining on the paired-associates task (encoding scan) and after 35 min
of training (retrieval scan). Performance during the encoding scan was
59% correct and during the retrieval scan 98% correct, with a mean re
action time of 709 ms during retrieval. The rCBF was also measured dur
ing a control condition without any instruction to encode or retrieve.
Compared with retrieval, encoding showed significant activation of th
e posterior part of the right middle frontal gyrus, the right inferior
parietal cortex, the cingulate cortex, the left inferior parietal cor
tex and the left inferior and middle temporal gyri. The rCBF increase
during encoding was strongly correlated with the rate of encoding. Ret
rieval was compared with both encoding and control. In none of these c
omparisons was there any prefrontal activation. The lack of prefrontal
activation during near-perfect performance of the retrieval task sugg
ests that the prefrontal cortex is not necessarily active when retriev
al is fast and accurate, or what might be called automatic. Encoding w
as not associated with more activation of the left than the right pref
rontal cortex. This result presents a limitation to the generality of
left prefrontal activation during episodic encoding, which has been fo
und in several previous brain imaging studies. Differences between stu
dies in the relative activation of left and right prefrontal cortex du
ring encoding and retrieval might he due to differences in paradigms,
the type of stimulus used, and the demand for working memory and verba
l analysis.