5. Long-Term Memory

Most modern theories of semipermanent structural change (or _engrams_,

as they are sometimes called) look either to the molecular level or to

the cellular level. Various specific locales for the engram have been

suggested, including (1) modifications of RNA molecular structure,

(2) changes of cell size, synapse area or dendrite extensions, (3)

neuropile modification, and (4) local changes in the cell membrane.

There is, in fact, rather direct evidence of the growth of neurons or

their dendrites with use and the diminution or atrophy of dendrites

with disuse. The apical dendrite of pyramidal neurones becomes thicker

and more twisted with continuing activity, nerve fibers swell when

active, sprout additional branches (at least in the spinal cord) and

presumably increase the size and number of their terminal knobs.

As pointed out by Konorski (11), the morphological conception of

plasticity according to which plastic changes would be related to the

formation and multiplication of new synaptic junctions goes back at

least as far as Ramon y Cajal in 1904. Whatever the substrate of the

memory trace, it is, at least in adults, remarkably immune to extensive

brain damage and as Young (24) has said: “ ... this question of the

nature of the memory trace is one of the most obscure and disputed in

the whole of biology.”

Self-Organizing Systems, 1963 · 第 26 / 92 题