Androgen Metabolism in the Juvenile Oscine
Forebrain: A Cross-Species Analysis at Neural Sites
Implicated in Memory Function
Colin J. Saldanha,
1,2
Nicola S. Clayton,
3
Barney A. Schlinger
1,2
1
Department of Physiological Science, University of California Los Angeles,
621 Charles E. Young Drive South, Los Angeles, California 90095
2
Laboratory of Neuroendocrinology of the Brain Research Institute,
University of California Los Angeles, Los Angeles, California 90095
3
Section of Neurophysiology and Behavior, University of California Davis, Davis, California 95616
Received 14 August 1998; accepted 19 March 1999
ABSTRACT: Juvenile songbirds are useful models
for studying the neural bases of memory. Memory-reli-
ant behaviors demonstrated at this stage include song
learning (most songbirds) and food caching (food-stor-
ing songbirds). Sex steroids are implicated in the mod-
ulation of memory processes in several vertebrates. The
songbird forebrain expresses aromatase, 5-reductase
and 5-reductase, enzymes which convert testosterone
to estradiol, 5-, and 5-dihydrotestosterone, respec-
tively. To explore the role of local androgen metabolism
on memory processes, we documented the activities of
these enzymes in the anterior neostriatum (NAN), cau-
domedial neostriatum (NCM), and hippocampus (HP) of
four species of juvenile songbird, two of which are food
storers. Areas were dissected, homogenized, and pro-
vided with radiolabeled substrate; and formed estro-
gens, and 5- and 5-reduced androgens were mea-
sured. In the NAN, 5-reductase was the predominant
enzyme, suggesting that local inactivation of testosterone
may preserve the sensitive period of song acquisition. In
the NCM, estrogens were formed in abundance despite
high 5-reductase, suggesting that locally high estrogen
synthesis may play a role in processes of song percep-
tion. In the HP, both estrogens and 5 reduced andro-
gens were formed, suggesting that HP function may be
modulated by both estrogens and androgens. Finally, a
derived measure of steroid-differential reveals that food-
storing songbirds differ from nonstorers in the steroidal
milleiu within the HP, but not in the NAN or NCM.
Thus, distinct loci within the juvenile songbird forebrain
are exposed to different patterns of androgen metabo-
lites. This local conversion may play a role in the neu-
roendocrine modulation of memory in these
birds. © 1999 John Wiley & Sons, Inc. J Neurobiol 40: 397– 406, 1999
Gonadal steroids have been implicated in aspects of
memory function in several vertebrates (McEwen et
al., 1995). In humans, testosterone (T) (Janowsky et
al., 1994) and estrogen (E) (Sherwin, 1988; Kampen
and Sherwin, 1996) are associated with increased
memory performance. In rodents, a corresponding
enhancement is seen in response to T (Gouchie and
Kimura, 1991; Roof and Havens, 1992), or E admin-
istration (Singh et al., 1994; O’Neal et al., 1996). The
data suggest the involvement of T or its aromatized
metabolite E in aspects of vertebrate memory.
Several oscine passerines (hereafter: songbirds) in-
cluding the zebra finch (Taenopygia guttata) (Vockel
et al., 1990; Schlinger and Arnold, 1991), white-
crowned sparrow (Zonotrichia leucophrys) (Schlinger
Correspondence to: C. J. Saldanha
Contract grant sponsor: JD French Alzheimer’s Foundation
Contract grant sponsor: NIH NS 35465
Contract grant sponsor: Whitehall Foundation
Contract grant sponsor: NSF; contract grant number: IBN
9120776
© 1999 John Wiley & Sons, Inc. CCC 0022-3034/99/030397-10
397