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The Journal of Neuroscience, December 1, 2000, 20(23):8897-8901
Thalamic Reticular Nucleus Activation Reflects Attentional Gating
during Classical Conditioning
Kerry
McAlonan,
Verity J.
Brown, and
Eric M.
Bowman
School of Psychology, University of St. Andrews, St. Andrews KY16
9JU, United Kingdom
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ABSTRACT |
All senses, except olfaction, are routed through the thalamus to
cerebral cortex. Thus, the thalamus is often referred to as the sensory
gateway to cortex. Located between thalamus and cortex is a thin lamina
of neurons called the thalamic reticular nucleus, which may function as
an attentional gate. The phenomenon of blocking in classical
conditioning provides an opportunity to test whether an attended
stimulus activates the thalamic reticular nucleus more than an
unattended stimulus: when a second stimulus is presented together with
a previously conditioned stimulus, conditioned responding to the second
stimulus is inhibited.
Different groups of rats were given conditioning sessions with a single
stimulus, a light or a tone, and then given conditioning sessions with
compound (light and tone) stimuli. Blocking was confirmed using probe
trials of single stimulus presentations. After a final test session of
compound stimulus presentations, the brains were processed for the
presence of Fos protein. Here we show that Fos-positive neurons were
more numerous in the sector of the thalamic reticular nucleus
associated with the attended conditioned stimulus than in the sector
associated with the unattended stimulus. Thus, we provide evidence for
an involvement of the thalamic reticular nucleus in selective attention.
Key words:
attention; thalamus; thalamic reticular nucleus; classical conditioning; rat; blocking
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INTRODUCTION |
The thalamic reticular nucleus
receives collaterals from both thalamocortical and corticothalamic
fibers (Jones, 1975 ; Ohara and Lieberman, 1985), and therefore it is in
a position to gate the flow of information between thalamus and cortex.
By virtue of its anatomy and physiological properties, the thalamic
reticular nucleus has been implicated in attention (Skinner and
Yingling, 1977; Yingling and Skinner, 1977; for review, see Guillery et al., 1998). There is also behavioral evidence to support this view
(Friedberg and Ross, 1993; Montero, 1997; Weese et al., 1999). Crick
(1984) said that if the thalamus is the gateway to cortex, then the
thalamic reticular nucleus is the "guardian of the gateway."
Functional sectors of the thalamic reticular nucleus are defined by the
origin of the cortical and thalamic collaterals they receive and by
physiology. In the rat, a visual sector is located caudodorsally in the
nucleus (Sumitomo et al., 1976; Hale et al., 1982; Coleman and
Mitrofanis, 1996; Lozsadi et al., 1996), whereas an auditory sector is
ventral to this (Shosaku and Sumitomo, 1983). These subsectors of the
reticular nucleus have extensive inhibitory interconnectivity and
therefore lateral inhibition could enhance the relay of relevant
(attended) information while attenuating the relay of irrelevant
(unattended) information. Because the organization of the thalamic
reticular nucleus suggests that it may function as an attentional gate,
we tested this idea by using a marker of neuronal activation, the
presence of Fos protein, during a behavioral task in which attention is
directed to one stimulus and not another, Kamin's (1969) blocking
procedure. In this procedure, a stimulus that reliably predicts reward
evokes a conditioned response. A second stimulus, introduced after
conditioning but presented simultaneously with the first, is redundant
and therefore results in no conditioned response. This second stimulus is referred to as the "blocked stimulus."
Using cytochrome oxidase as a metabolic marker, it has been
demonstrated that secondary auditory cortex responds to an auditory conditioned stimulus predictive of footshock, but not if the stimulus is blocked by previous conditioning to a visual stimulus (Poremba et
al., 1997). Metabolic differences between the two conditions were
observed in structures in which the convergence of auditory and
somatosensory input is modulated by visual input. Thus, the authors
concluded that blocking was not attributable to inattention to the
blocked stimulus, supporting instead an associative explanation of
blocking (Rescorla and Wagner, 1972). Nevertheless, their data do not
preclude the possibility that blocking has an attentional component.
Attentional theories of blocking (Mackintosh, 1975; Solomon, 1977;
Pearce and Hall, 1980; Holland and Gallagher, 1993) propose that
limited attentional resources are directed to the conditioned stimulus.
Because the blocked stimulus conveys no additional relevant
information, it is unattended. If the thalamic reticular nucleus
mediates selective attention, labeling of Fos protein will be
restricted to the sector of the thalamic reticular nucleus associated
with the attended conditioned stimulus and not in the sector of
thalamic reticular nucleus associated with the blocked, unattended, stimulus.
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MATERIALS AND METHODS |
Subjects and apparatus. Fifteen experimentally
naïve male Lister hooded rats (Charles River, Margate, UK) were
pair-housed. The colony was maintained on a 12 hr light/dark cycle
(lights on 7:00 A.M.) with water available ad libitum in the
home cage and food restricted to 15-20 gm/d, provided after testing.
Procedures were licensed under the UK Animals (Scientific Procedures)
Act, 1986. Testing was conducted in standard Skinner boxes, with a food
hopper between two fixed levers (Campden Instruments, Sileby, UK).
Procedure. An appetitive blocking procedure, with food
reward, was used (Table 1). After
conditioning to a single stimulus and exposure to the compound
stimulus, rats were given probe trials of alternating presentations of
single stimuli under extinction, to verify blocking. Thus, there were
12 d of conditioning sessions, comprising 120 stimulus-reward
pairings per day. The conditioned stimulus, presented for 1 sec, was a
light in the food hopper (light, n = 5) or an 800 Hz,
70 decibel tone (tone, n = 6) from a speaker located in
the ceiling of the chamber, or the light and tone together (both,
n = 4). The reward was a 50% sucrose pellet (45 mg;
BioServ, Frenchtown, NJ), delivered into the hopper by automatic
dispenser coincident with the offset of the conditioned stimulus.
Interstimulus intervals were randomized between 1 and 30 sec. On day 13 of training, there was a single session of 70 presentations of the
compound stimulus followed by reward. Blocking was assessed the
following day by 30 alternating presentations of the light and tone
alone, with no reward. Responses at the food hopper were recorded, and
responses in the 1 sec preceding the stimulus were subtracted from
responses during the stimulus, so that positive values would indicate
greater responding during the stimulus, and negative values would
indicate greater responding before the stimulus. The final test
session, which lasted 105 min, consisted of presentations of the
compound stimulus followed by reward.
Immunohistochemistry. Immediately after the final session,
the brains were processed for the presence of Fos protein. To prevent nonspecific Fos elevation, for the 12 hr preceding the final test, rats
were held in darkness with minimal exposure to auditory and visual
stimuli. Rats were carried to the darkened testing room in covered
cages and immediately placed in the operant chambers. Fos protein
synthesis follows mRNA expression 30-45 min after stimulation and has
a half-life of 2 hr. Therefore, the rats were anesthetized immediately
after the final test session and transcardially perfused with saline
followed by 4% paraformaldehyde.
One in four 50 µm sections were cut and put into PBS. Sections
were treated as follows: 30 min wash in 20% sucrose solution (in PBS),
two rinses in PBS followed by a 60 min wash in blocking solution (100 ml of PBS, 20 ml of goat serum, and 1 ml of Triton X-100), two PBS
rinses, 48 hr incubation in primary c-fos antibody (Ab-5; 1:20,000;
Oncogene Research Products, Cambridge, MA), five PBS rinses, 45 min
incubation period on a shaker in Vector Laboratories (Peterborough, UK)
IgG solution (5 µl/ml; Vectastain rabbit ABC kit), five PBS rinses,
45 min incubation in ABC complex (antibody diluting solution, substrate
A and substrate B, each 20 µl/ml) on a shaker, and five PBS rinses,
incubated for 2-10 min in diaminobenzidine peroxidase substrate (Sigma
Fast 3-3'-diaminobenzidine tetrahydrochloride with metal enhancer
tablet sets; Sigma-Aldrich, Poole, UK).
The regions of interest were the visual sector and the auditory sector
of the thalamic reticular nucleus. Based on electrophysiology (Shosaku
and Sumitomo, 1983) and anatomy (Pinault and Deschenes, 1998), these
regions are located approximately between bregma  3.1 and 4.1 mm,
with the visual sector located dorsal to the auditory sector.
Therefore, we counted all neurons stained positively for Fos protein in
the dorsal 50% and ventral 50% of the thalamic reticular nucleus on
15 sections between 2.7 and 4.1 mm, but used counts from the
sections between approximately bregma 3.1 and 4.1 mm to obtain means
for visual (dorsal 50%) and auditory (ventral 50%) sectors.
| |
RESULTS |
The probe trials confirmed that the procedure resulted in a
conditioned response to the stimulus presented during the conditioning sessions and not to the second stimulus. Rats conditioned to the compound stimulus showed intermediate levels of conditioned responding to either stimulus alone (Fig. 1;
interaction of group and stimulus, F(2,12) = 24.7, p < 0.01).
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Figure 1.
Conditioned responding was assessed in probe
trials in which the stimuli light or toneswere presented alone and
not followed by food reward. Responses in the 1 sec before stimulus
onset were subtracted from responses during the 1 sec stimulus. The
left panel shows responding to the light; the
right panel shows responding to the tone. The rats were
preconditioned, either with the light (white bars), the
tone (black bars), or the compound (gray
bars) stimulus. The number of hopper entries was highest when
the preconditioned stimulus was presented and lowest when the blocked
stimulus was presented. Intermediate values were observed for rats
trained with the compound stimulus.
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Modality-specific subsectors of the thalamic reticular nucleus were
activated according to the attended stimuli. Neurons stained positively
for Fos protein were more numerous in the sector of the thalamic
reticular nucleus associated with the conditioned, attended, stimulus
(Figs. 2,
3). Thus, rats conditioned to light showed activation of visual thalamic reticular nucleus, whereas rats
conditioned to tone showed activation of the auditory thalamic reticular nucleus. Rats who had received previous conditioning to the
compound stimulus showed activation of both visual and auditory
thalamic reticular nucleus (interaction of group and thalamic reticular
sector, F(2,12) = 4.0, p < 0.05).
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Figure 2.
Mean counts of Fos-positive neurons in the dorsal
50% and ventral 50% of the thalamic reticular nucleus on sections
between bregma 2.7 and 4.1 mm for each group.
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Figure 3.
Coronal sections through the thalamic reticular
nucleus of the same hemisphere of one rat from each group, showing
Fos-positive neurons in different sectors of the thalamic reticular
nucleus according to the group. Fos staining was seen in the dorsal
(visual) portion of the thalamic reticular nucleus in the
light-conditioned, but not in the tone-conditioned, group. The converse
pattern of activity was found in the ventral and more posterior
(auditory) portion of the thalamic reticular nucleus. In the compound
stimulus-conditioned rats, Fos-positive neurons were found in both
regions. On the test day, all rats received the same, compound,
stimulus presentations. Abbreviations from Paxinos and Watson (1997):
Rt, thalamic reticular nucleus; fi,
fimbria; st, stria terminalis; ic,
internal capsule; eml, external medullary lamina;
LDVL, laterodorsal nucleus ventrolateral division;
VPM, ventral posteromedial thalamic nucleus;
VPL, ventral posterolateral thalamic nucleus.
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Across rats in all groups, there was a positive correlation between
magnitude of conditioned responding to the light and the number of
Fos-labeled neurons in the visual thalamic reticular nucleus (Fig.
4; partial correlation, removing
variability caused by auditory conditioned responding,
r = 0.45, df = 12, p = 0.053, one-tailed). There was a stronger positive correlation between the
magnitude of auditory conditioned responses and the number of
Fos-positive neurons in the auditory thalamic reticular nucleus (partial correlation, removing variability caused by visual conditioned responding, r = 0.62, df = 12, p < 0.01, one-tailed). Thus, in addition to the mean differences between
groups, the number of Fos-positive neurons in each sensory sector
predicts the magnitude of behavioral effects in individual animals.
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Figure 4.
Plots showing the residual conditioned response
against the residual Fos count. The visual responses (visual
conditioned response and the mean Fos count in the visual portion of
the thalamic reticular nucleus) were predicted from the auditory
responses and vice versa. The observed responses were then subtracted
from predicted responses, to obtain the residual. Thus, variability in
the residuals of the visual (or auditory) responses is independent of
variability in the auditory (or visual) responses.
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DISCUSSION |
Activation of the thalamic reticular nucleus was observed in rats
attending to conditioned stimuli in a blocking procedure. The
activation was specific to the sector of the thalamic reticular nucleus
associated with the sensory modality of the conditioned stimulus. There
was less activation of the sector of the thalamic reticular nucleus
associated with the sensory modality of a blocked stimulus, which did
not elicit a conditioned response. The degree of Fos labeling in the
thalamic reticular nucleus correlated with the magnitude of the
conditioned response to the conditioned stimulus. This result supports
the view that the thalamic reticular nucleus is involved in the
processing of sensory information (Hartings et al., 2000) but
demonstrates that this processing is modulated by attention.
The three groups of ratslight, tone, and both light and
tonediffered only according to the stimuli to which they had received before conditioning. On the test day, all rats were treated the same
and received the same repeated presentations of the compound light-tone stimulus. The possibility that the rats, in attending and
responding to the conditioned stimulus, might fail to orient to the
second stimulus was considered. To avoid this objection, the light
stimulus was presented within the food hopper, to which all the rats
oriented and approached when the compound stimulus was presented. The
tone was presented from an omnidirectional speaker, located above the
rat in the chamber. Thus, the stimuli impinging on the sensorium of the
rats would not differ as a function of the rats orienting behavior, so
that neural differences can reasonably be expected to reflect
differences in attention and not sensation. It should also be noted
that no attempt was made to restrict other stimuli within the chamber
during the test session: a ventilation fan was on throughout and
neither the dispenser nor the hopper panel was silent.
Montero (1997) demonstrated selective c-fos activation in the visual
sector of the thalamic reticular nucleus in normal rats exploring a
complex novel environment and in the somatosensory sector of the
thalamic reticular nucleus in functionally blind rats, dependent on
tactile cues to explore. Montero (1997) suggested that sectors of the
thalamic reticular nucleus actively compete for limited attentional
resources. It is also plausible to suggest that the sectors of the
thalamic reticular nucleus might compete to provide limited attentional
resources and thus the thalamic reticular nucleus subserves selective
attention. Montero (2000) has recently shown that Fos labeling in the
thalamic reticular nucleus, but not the geniculate nucleus, is
dependent on primary visual cortical input.
Rhythmic burst firing in the thalamus, including the reticular nucleus,
is prevalent during periods of neural synchronization, such as during
slow wave sleep, deep anesthesia, or absence seizures (Steriade et al.,
1986, 1993a,b) and is thought to represent a temporary detachment of
relay cells from their sensory inputs. In periods of cortical
desynchrony, the thalamic reticular nucleus fires in a tonic mode.
Guido and Weyand (1995) demonstrated that arrhythmic burst firing in
thalamic relay cells in awake animals may support better signal
detection by elevating the signal-to-noise ratio and suggested that the
functional implications of burst versus tonic firing in the awake
animal may relate to orienting versus focal attention. This suggestion
is given further support by recent work of Ramcharan et al. (2000), who
suggest that burst firing in the awake animal provides an attentional
"wake-up call." The switch from burst to tonic firing in thalamic
relay cells may be modulated by descending cortical projections and by
the cholinergic innervation of the thalamus and thalamic reticular nucleus (McCormick and Prince, 1986; Marks and Roffwarg, 1991; Kim and
McCormick, 1998). The thalamic reticular nucleus inhibits thalamic
relay cells (French et al., 1985; Cox et al., 1997; Kim et al., 1997),
which would be consistent with the suggestion that sensory input
activates the thalamic reticular nucleus, which in turn inhibits and
changes the firing mode of the thalamic relay cells. Therefore, it is
parsimonious to suggest the Fos-labeling seen here reflects neuronal
excitation in the sector of the thalamic reticular nucleus associated
with the attended modality.
The specificity of the labeling of thalamic reticular nucleus supports
the view that this nucleus is more than a component of the sensory
relay but rather acts as an attentional gate or filter. These data do
not demonstrate that the mechanism of blocking is attentional, however
they do provide support for the suggestion that blocking has an
attentional component, changing the processing of stimuli according to
the associative value of the stimulus.
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FOOTNOTES |
Received May 31, 2000; revised Aug. 22, 2000; accepted Sept. 12, 2000.
This work was supported by the Medical Research Council (UK) project
grant number G9531294 (E.M.B.) K.M. received a Biotechnology and
Biological Sciences Research Council (UK) Special Studentship. We thank
Mary Latimer for advice and assistance with immunohistochemistry, Dr.
Patrick Pallier for helpful discussion, and the staff of the School of
Psychology Animal House and Workshop.
Correspondence should be addressed to Dr. Verity J. Brown, School of
Psychology, University of St. Andrews, St. Andrews KY16 9JU, Scotland,
UK. E-mail: vjb{at}st-and.ac.uk.
| |
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TOP
ABSTRACT
INTRODUCTION
