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Volume 17, Number 23,
Issue of December 1, 1997
pp. 8997-9009
Development of Intrinsic and Synaptic Properties in a Forebrain
Nucleus Essential to Avian Song Learning
Received July 25, 1997; revised Sept. 12, 1997; accepted Sept. 17, 1997.
Frederick S. Livingston and
Richard Mooney
Department of Neurobiology, Duke University Medical Center, Durham,
North Carolina 27710
In male zebra finches, the lateral magnocellular nucleus of the
anterior neostriatum (LMAN) is necessary for the development of learned
song but is not required for the production of acoustically stereotyped
(crystallized) adult song. One hypothesis is that the physiological
properties of LMAN neurons change over development and thus limit the
ability of LMAN to affect song. To test this idea, we used in
vitro intracellular recordings to characterize the intrinsic
and synaptic properties of LMAN neurons in fledgling [posthatch days
(PHD) 22-32] and juvenile zebra finches (PHD 40-51) when LMAN
lesions disrupt normal song development, and in adults (>PHD 90) when
LMAN lesions are without effect. In fledglings, depolarizing currents
caused LMAN projection neurons to fire bursts of action potentials
because of a putative low-threshold calcium spike (LTS). In contrast,
juvenile and adult LMAN projection neurons fired accommodating trains
of action potentials when depolarized but did not exhibit the burst
mode of firing. Electrical stimulation of thalamic afferents elicited
both monosynaptic EPSPs mediated by AMPA and NMDA receptors and
polysynaptic IPSPs mediated by GABAA receptors from LMAN
neurons at all ages studied here. In whole-cell voltage-clamp
recordings, the EPSCs (NMDA-EPSCs) consisted of fast and slow
components. Unlike juvenile and adult NMDA-EPSCs, those in fledglings
were dominated by the slower component. Thus, both the intrinsic and
synaptic properties of LMAN neurons change markedly during early song
development (PHD 22-40) and achieve several adult-like properties
during early sensorimotor learning and well before the time when LMAN
lesions no longer disrupt song development.
Key words:
LMAN;
DLM;
song nuclei;
zebra finch;
whole cell;
intracellular recordings;
NMDA receptors;
AMPA receptors;
low-threshold
calcium spike;
LTS;
vocal plasticity;
critical periods;
song
learning
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