Info

?

BDNF expression

Multiple

Contextual conditioning (136)

Yes

?

Increased glu release

PP/dentate

Operant conditioning (137-139)

?

?

Increased glu uptake

SC/CA1

Contextual conditioning (140)

?

?

hypothesis that LTP occurs in vivo with memory formation. There also have been quite a number of other experiments of exactly the same sort, demonstrating that LTP-associated molecular events occur in the hippocampus and other parts of the CNS with learning (see Table 2). I focused on ERK activation as a specific example of this general category of experiment because it is the system with which I am most familiar.

Thus, while it has not been possible so far to directly demonstrate LTP physiologically in association with spatial learning, biochemical "markers" for LTP induction such as ERK activation, CaMKII activation, PKC activation, glutamate release, and altered gene expression (see Table 2) have been demonstrated to occur with spatial learning. That such a broad spectrum of molecular changes occurs with both LTP in vitro and spatial learning in vivo strongly suggests that LTP is indeed occurring in the hippocampus with hippocampus-dependent memory formation.

Molecular Events in LTP Versus Memory Formation—Some Dissociation in Time Courses

One additional comment is worth making in the context of biochemical markers for LTP. The data are consistent with the hypothesis that LTP is occurring in vivo, as various investigations have found that biochemical mechanisms occurring in LTP are valid predictors of biochemical changes associated with and necessary for associative learning. In a number of studies, however, different time courses for LTP-associated molecular changes versus learning-associated changes have been observed. For example, in our studies of protein kinase activation with LTP and contextual fear conditioning, we found different time courses for MAPK, PKC, and a-CaMKII activation during LTP versus fear conditioning (Figure 6). MAPK is activated within 2 minutes of LTP induction and returns to baseline levels of activation within 1 hour. PKC undergoes a similar time course of activation as a-CaMKII, both

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