Pharmacology Biochemistry& Behavior, Vol. 31, pp. 523-531.©PergamonPress plc, 1989. Printedin the U.S.A.
Specific Effects of Punishment
on Amino Acids Turnover in
Discrete Rat Brain Regions
TATSUO MIYAUCHI, 1 STEVEN I. DWORKIN, CONCHITO CO
AND JAMES E. SMITH
Psychiatry Research Unit, Departments of Psychiatry and Pharmacology
Louisiana State University Medical Center, Shreveport, LA 71130
Received 3 April 1987
MIYAUCHI, T., S. I. DWORKIN, C. CO AND J. E. SMITH. Specific effects of punishment on amino acids turnover in
discrete rat brain regions. PHARMACOL BIOCHEM BEHAV 31(3) 523-53 !, 1988.--Specific effects of punishmenton the
turnover rates of aspartate (Asp), glutamate (Glu) and gamma-aminobutyric acid (GABA) in 14 brain regions were investi-
gated in rats exposed to punishment. Two yoked controls were also used in an attempt to separate the nonspecific effects of
response rate, reinforcement density and direct effects of punisher (foot shock). Punished and unpunished littermate rats
had similar response rates, and the reinforcement density was almost identical for both groups. A third group (yoked-
shock rats) received food and shock independent of responding whenever these were given to the punished rats. When
compared to the unpunished rats, the punishment increased the turnover rates of the three amino acids in all brain regions
examined except GABA turnover in the caudate-putamen and preoptic-diagonal band. The majority of these changes by the
punishment were similarto the effects of the yoked-shock (yoked-shock versus unpunished), although the magnitude of increase
by the punishment was mostly larger than that by the yoked-shock. Six changes by the punishment (increase in the turnover rates
of Asp in the thalamus, Glu in the hypothalamus and GABA in the cingulate cortex, entorhinal-subicular cortex, dentate
gyrus and hypothalamus) appeared to be the specific effects of punishment since the yoked-shock did not affect these
parameters. These results suggest that the punishment caused a hyperexcitation of the amino acidergic neurons in the
limbic systems, particularly those in Papez's circuit.
Punishment Conflict Amino acid neurotransmitters
Neurotransmitter turnover rates
Gamma-aminobutyric acid Anxiety
GELLAR-SEIFTER type of conflict-punishment proce-
dures (17) are useful animal behavioral techniques to investi-
gate the effects and mechanisms of action of anxiolytic
drugs. In this procedure, operant responding which is main-
tained by positively reinforcing stimuli (food, water, etc.) is
suppressed (punished) by response-dependent presentation
of noxious stimuli (electric foot shock, etc.), and anxiolytic
drugs increase the rate of punished responding. A large body
of pharmacological evidence exists implicating central
gamma-aminobutyric acid (GABA) [reviewed by Sepinwall
(38), Enna (14) and Sanger (36)] or serotonin [reviewed by
Sepinwall (38) and Iversen (21)] in the antipunishment effect
of anxiolytics in rats. In contrast to the numerous phar-
macological studies using punishment procedures, however,
there has been no report, to our knowledge, assessing the
neurochemical consequence of punishment. This study was
designated to detect punishment-specific changes in turn-
over rates of amino acid neurotransmitters as indexes of
amino acidergic neuronal activities in discrete brain regions
in rats.
Most punishment experiments utilize multiple schedule
procedures in which punished (2-5 min duration) and unpun-
ished (5-15 min duration) periods appear alternately during a
session. However, these schedules are not suitable for the
neurochemical procedure we are using to determine turnover
rates (8, 24, 25, 39, 40). In this procedure, animals are killed
60 and 90 min after intravenous pulse injection of radiolabel-
led precursors of neurotransmitters, which requires animals
being under a single component (punished or unpunished)
during the pulse interval. Also, response rate and reinforce-
ment density during the punished period are usually less than
those during the unpunished period. The difference in motor
activity (response rate) and food intake (reinforcement den-
sity) between the two components may obscure punish-
ment-specific neurochemical effects.
In the present experiments, therefore, a behavioral pro-
cedure which was suitable for the neurochemical examination
of punishment was developed. To separate the neurochemical
effects of nonspecific factors described above, we used a be-
havioral procedure [yoked-box procedure (16)] which resulted
in similar rates of responding and reinforcement density for
punished and unpunished littermate rats. The yoking proce-
1Present address: Fujigotemba Research Laboratories, Chugai Pharmaceutical Co. Ltd. 705-1 Komakado, Gotemba 412, Japan.
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