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The reaching reaction in the rat: a part of the digging pattern?

The relationship between acquired lateralized reaching and analogous innate movement patterns was examined in adult rats. In Expt. I, 8 naive rats were trained to reach into a narrow trough-shaped feeder. Analysis of the videorecorded movement showed that the forelimb extension starts 120 ms before the forepaw enters the feeder, lasts 190 ms and can be repeated at 280-ms intervals. Before reaching, the rat assumes a stereotyped posture and inserts its nose into the feeder entrance. Twelve rats used in Expt. II were raised from weaning with limited opportunity to practice manipulative skills. Six of these animals were then trained as in Expt. I. In spite of the lack of experience, they learned the reaching task as efficiently as normal rats. The remaining 6 rats were offered food placed into a wide tunnel-shaped feeder. The animals did not take advantage of the possibility of using both forelimbs, but developed typical lateralized reaching. In Expt. III, 4 rats were trained to retrieve food pellets buried under a layer of sand. The videorecorded digging movement essentially resembled reaching but the two forepaws alternated, the extension amplitude was shorter and the movement was never terminated by grasping. It is concluded that the lateralized reaching develops from modified digging and that the kinship of the two movements accounts for the stereotypy and limited modifiability of the acquired reaction.

Animals↗

Septohippocampal system and the prelimbic sector of frontal cortex: a neuropsychological battery analysis in the rat.

Rats with lesions in the posterodorsal septal area (aimed at transecting the precommissural fornix) and rats with lesions in the prelimbic sector of the medial frontal cortex were tested postoperatively on a neuropsychological test battery comprised of the following tasks: time-spent-eating in two adaptation boxes, time-to-emerge and ambulation in an open field, general activity, contingently-reinforced (continuous) and schedule-specific (delayed non-matching-to-sample) T-maze alternation, visual and olfactory discrimination in a T-maze, temporal alternation (response patterning) and tactile Go/No-Go discrimination in a runway, approach-avoidance conflict in the runway, step-through inhibitory avoidance, one-way active avoidance, two-way active avoidance, and conditioned taste aversion. It was found that: (1) rats with septal (SEP) lesions spent more time eating than control (CON) rats and rats with lesions in prelimbic cortex (PRE). PRE rats did not differ from CON rats; (2) PRE rats emerged into an open field faster, and spent less time in home cage than CON and SEP rats. SEP rats stayed in home cage less than CON rats. PRE and SEP rats crossed more squares in the open field than CON rats; (3) SEP and PRE rats were more active than CON rats; (4) SEP rats performed the contigently-reinforced and schedule-specific T-maze alternation tasks worse than PRE rats, and PRE rats performed these tasks worse than CON rats. PRE, but not SEP, rats showed improvement with continued practice at brief intertrial and interrun intervals.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Light input and the reversal of functional lateralization in the chicken brain.

During its later stages of development, the chicken embryo is oriented in the egg so that it occludes its left eye with its body and the right eye is positioned so that it can receive light input. This lateralized light input has been shown to play a decisive role in determining the direction of brain lateralization for two behavioural functions, copulation and performance of a visual discrimination task known as the 'pebble floor test', since the direction of lateralization for these functions can be reversed by occluding the right eye of the embryo on day 19/20 of incubation and at the same time exposing the left eye to light. The sensitive period during which this role of lateralized light input influences the lateralization for copulation extends to day 1 posthatching if the eggs are incubated and hatched in darkness, but it is over by day 3. For the pebble floor test the sensitive period is already over by day 1 posthatching. By exposing eggs to light for various times on day 19 of incubation, it was possible to determine that between 2.5 and 6 h of lateralized light exposure is necessary to stabilise the normal direction of lateralization so that it can no longer be reversed by occlusion of the right eye. Thus, in the developing chicken embryo substantial neural reorganization must occur in response to a brief period of lateralized light input.

Animals↗

Effects of dopamine depletion from the caudate-putamen and nucleus accumbens septi on the acquisition and performance of a conditional discrimination task.

Three experiments compared the effects of dopamine depletion from the caudate-putamen (CAUD; dorsal striatum) or nucleus accumbens septi (NAS; ventral striatum), or a systemically administered dopamine receptor antagonist (alpha-flupenthixol) on the acquisition and performance of a conditional discrimination task involving temporal frequency. In Expt. 1, rats receiving 6-hydroxydopamine (6-OHDA) lesions of the CAUD were impaired in the acquisition of a visual version of the task, and rats with 6-OHDA lesions of the NAS were not reliably impaired. Even when the rats with CAUD lesions had acquired the discrimination, they were still significantly slower to collect earned food pellets. Both CAUD and NAS lesions reduced a bias to respond to the faster of the two discriminative stimuli. In Expt. 2, rats with 6-OHDA lesions of CAUD were markedly impaired in their accuracy and speed of responding when they had been trained to criterion preoperatively. These effects could not be mimicked in controls by prefeeding (which had only minor effects on performance). Rats with 6-OHDA-induced lesions of the NAS were unimpaired in either visual or auditory discrimination performance, but were slower to extinguish responding than controls. In Expt. 3, alpha-flupenthixol (0.1-0.56 mg/kg, i.p.) produced dose-dependent impairments in both latency to respond and choice accuracy in visual and auditory versions of the task. In conjunction with other results, these data suggest that (1) dopamine receptor blockade and central dopamine depletion can impair discrimination performance under certain conditions (2) dopamine depletion from the ventral and dorsal striatum, respectively, have dissociable effects on behaviour controlled by conditioned reinforcers and discriminative stimuli and (3) the disruption of discrimination performance by dorsal striatal dopamine depletion is probably attributable to several factors.

Animals↗

Effects of partial lesion of dorsal hippocampal afferent and GM1 ganglioside treatment on conditioned emotional response and hippocampal afferent markers in rats.

Acquisition of the conditioned emotional response (CER) in 32 male hooded rats previously learned to press a bar for food and divided into four groups was studied. Two groups received electrolytic lesions of the dorsal hippocampal afferent and were thereafter injected either with GM1 ganglioside (30 mg/kg daily) or with buffer. Two remaining groups were sham operated and similarly injected. The partial hippocampal deafferentation evoked immediate enhancement of bar presses rate which persisted during the 2-week period of testing. CER training undertaken 2 days after surgical procedures appeared unsuccessful, whereas similar training with a cue of different modality initiated a week later resulted in acquisition of conditioned suppression of bar presses in all groups. Toward the end of training the conditioned suppression was more pronounced in lesioned than in control rats. The GM1 injections attenuated the conditioned suppression in control rats, presumably due to an antinociceptive role of ganglioside treatment. Behavioural training did not change the normal distribution pattern in cholinergic and serotonergic hippocampal afferent markers showing dorso-ventral gradient along longitudinal axis. The lesion-induced decrease pattern was also not affected. However, in contrast to previous findings in non-trained animals, the GM1 treatment was not effective in protecting against degenerative changes in the hippocampus of trained rats.

Afferent Pathways↗

Differences in the recovery rate of a learned forelimb movement after ablation of the motor cortex in right and left hemisphere in white rats.

After ablation of the motor cortex contralateral to the preferred limb, rats were forced to use this limb in grasping food out of a horizontal tube by restraining the non-preferred foreleg with a bracelet. In a right-handed rat the motor behaviour is less impaired and the recovery is better after a left hemisphere lesion in comparison with a left-handed rat after a right hemisphere lesion. This difference is primarily the time needed to use the limb in reaching for seed (testperiod 3) and the total time needed for 10 successive seizings of seed (testperiod 5). Surprisingly, the differences in the course of recovery do not correlate with the degree of preoperative limb preference: the initially ambidextrous rate also show the same differences in results of the motor cortex ablation in left and right hemisphere. However, in the group of initially consistent ambidextrous rats, after 10 weeks of tests with a restrained forelimb, the testing under unrestrained free conditions shows a gradual decrease in the induced limb preference and a shift to the use of the foreleg contralateral to the intact hemisphere, while in contrast the initially consistent left- or right-handed rats preserve the limb preference under the unrestrained testing conditions. Therefore the degree of initial preference still influences the choice of limb after motor cortex ablation and intensive training to use the 'damaged' limb.

Animals↗

The effects of hippocampectomy on performance by rats of a running recognition task using long lists of non-spatial items.

Unoperated rats, or rats with cortical control lesions or hippocampal aspiration lesions, were trained on a non-spatial delayed non-matching-to-sample task. Once the non-matching rule had been learned, the task was modified so as to assess list memory, with list lengths of up to 32 items. Hippocampectomy induced a significant drop in choice accuracy, that was independent of list length; nonetheless, hippocampectomised rats still performed at better than chance levels. The cortical control lesion appeared to induce an intermediate (though generally non-significant) deficit. It was concluded that hippocampectomy leaves rats still capable of choosing with considerable accuracy on non-spatial working memory tasks that use trial-unique or 'pseudo-trial-unique' stimuli, even under experimental conditions in which a heavy memory load is imposed.

Animals↗

Chronic infusions of GABA into the medial prefrontal cortex induce spatial alternation deficits in aged rats.

It has been proposed that functions associated with the prefrontal cortex could change as a consequence of aging. Previous experiments in young rats have demonstrated that anatomical lesions or chronic GABA infusions into this area produce deficits in spatial delayed alternation tasks. The present study examines the effect of chronic (7 days) GABA or saline infusion into the prefrontal cortex on the performance of delayed alternation task in old rats (24 months). The results suggested that aged rats needed more sessions to acquire the delayed alternation task. GABA infusions into the prefrontal cortex produced deficits in spatial alternation tasks similar to those previously observed in young rats. Performance rapidly recovered after the infusion period. Histological analysis showed similar lesion size in both groups. The results suggest that aged prefrontal cortex and/or related areas participating in the acquisition of the delayed alternation task are more sensitive to aging processes. Furthermore, the prefrontal cortex is important for the retention of a previously learned spatial delayed alternation task. The structures involved in functional recovery from these deficits appear to be fully functional in aged rats.

Aging↗

Role of the tectal and posterior commissures in lateralization of the avian brain.

We report a role for the tectal and posterior commissural system (TC/PC) in functional lateralization of the chicken brain. Unlike the corpus callosum of mammals which is thought to generate lateralization, the TC/PC in the chicken brain is shown to suppress lateralization. The TC/PC was sectioned on day 2 post-hatching. The performance of this TC/PC-sectioned group on a battery of monocular, behavioural tests was compared to a sham-lesioned group and an untreated group. The first of these tests measured pecking and fear responses to a small bead. In both the sham-lesioned group and the untreated group there was no change in the pecking response over repeated presentations of the bead, nor was there a left eye/right eye difference in either of these groups. In contrast, the TC/PC-sectioned group showed increased pecking of the bead each time it was presented. This increase in the pecking response was significantly greater for those chicks tested using the right eye than for those tested using the left eye. That is, sectioning the TC/PC caused a lateralized response which was not seen in either of the control groups, demonstrating that the intact TC/PC suppresses lateralization at the tectal level. In the other tasks (latency to peck at a mealworm and a visual discrimination learning task) the untreated controls showed clear left eye/right eye differences while the sham-lesioned group and the TC/PC-sectioned group showed no lateralization. It is suggested that for these two tasks the loss of lateralization is due to lesioning an area in the midbrain but not to sectioning the TC/PC.

Animals↗

Effects of a long-interval retention on the hypoxia-induced impairment of performance in a fixed-interval schedule.

Retention of the fixed-interval (FI) responding was studied in rats after a chronic hypoxic treatment. The rats were trained over a 25-day period on a FI 60 s schedule. The rats were then given a normoxic or hypoxic (3.5% O2 for 10 min) treatment once a day for 4 days after which they were maintained in the animal quarters for 6 supplementary days. A retention test consisting of 4 daily FI 60 session followed the 10-day interval. The results showed that the performance during the retention test was very similar for both groups of rats. These data contrast with the deficit of retention observed when the retention FI 60 session were undertaken 24 h after the same hypoxic treatment. It is suggested that the results are due to a recovery from hypoxia-induced retrieval deficits rather than to the progressive attenuation of non-specific effects of hypoxia, such as anhedonia, hypoactivity, motivational changes or structural brain damage.

Amnesia, Retrograde↗

Microinjections of flupenthixol into the caudate-putamen but not the nucleus accumbens, amygdala or frontal cortex of rats produce intra-session declines in food-rewarded operant responding.

Results of recent studies suggest that dopamine (DA) transmission in the caudate-putamen may be involved in food reward-related learning. The purpose of the present study was to evaluate this hypothesis by injecting the DA antagonist cis-flupenthixol (25 micrograms in 0.5 microliters) into the dorsal caudate-putamen of rats (n = 19) trained to lever press for food presented according to a variable interval 30-s schedule. Additional groups received non-reward (n = 8), systemic cis-flupenthixol (0.01, 0.1 mg/kg i.p.; ns = 8), dorsal caudate-putamen injections of the inactive isomer trans-flupenthixol (n = 10), frontal cortical (dorsal to the caudate-putamen site) injections of cis-flupenthixol (n = 6), or cis- or trans-flupenthixol injected into the nucleus accumbens (ns = 9, 8) or amygdala (ns = 6, 5). Rats were tested in 30-min sessions and response rates were recorded every 5 min. As expected, non-reward produced a gradual decline in responding. A similar pattern was seen in the groups receiving systemic (0.1 mg/kg) or dorsal caudate-putamen injections of cis-flupenthixol. No significant effect was seen following systemic (0.01 mg/kg), cortical or amygdala cis-flupenthixol or dorsal caudate-putamen or amygdala trans-flupenthixol. Accumbens cis-flupenthixol reduced rates but did not produce a gradual decline in responding; however, accumbens trans-flupenthixol led to a time-dependent elevation in response rates making interpretation of the accumbens results difficult. It was concluded that dopaminergic projections to the dorsal caudate-putamen may play a critical role in mediating the effects of food-reward on operant responding.

Amygdala↗

Neuronal responses in the ventral striatum of the behaving macaque.

To analyse the functioning of the ventral striatum, the responses of more than 1,000 single neurons were recorded in a region which included the nucleus accumbens and olfactory tubercle in 5 macaque monkeys. While the monkeys performed visual discrimination and related feeding tasks, the different populations of neurons found included neurons which responded to novel visual stimuli; to reinforcement-related visual stimuli such as (for different neurons) food-related stimuli, aversive stimuli, or faces; to other visual stimuli; in relation to somatosensory stimulation and movement; or to cues which signalled the start of a task. The neurons with responses to reinforcing or novel visual stimuli may reflect the inputs to the ventral striatum from the amygdala and hippocampus, and are consistent with the hypothesis that the ventral striatum provides a route for learned reinforcing and novel visual stimuli to influence behaviour.

Amygdala↗

Morphological changes associated with stages of memory formation in the chick following passive avoidance training.

Memory formation following learning is presumed to result from modification in the efficacy of neural circuitry, either through strengthening of pre-existing synapses, or formation of new contacts. An ideal paradigm to investigate memory formation is one-trial passive avoidance training of day-old chicks, in which the birds learn to avoid pecking a bead coated with an aversive substance, methylanthranilate. Following training, a sequence of biochemical, electro-physiological, pharmacological and morphological events takes place within two loci in the forebrain, the intermediate and medial hyperstriatum ventrale (IMHV), and part of the paleostriatal complex, the lobus parolfactorius (LPO). Our data reviewed here suggest that the initial acquisition of memory involves population changes in the fine spatial organization of synaptic vesicles and active zones in synapses in the IMHV whereas longer-term changes are more prominent in the LPO and involve, primarily, a bilateral increase in the density of synapses and dendritic spines. The short-term synaptic changes are as dynamic as the molecular changes which have hitherto been considered the preserve of short-term correlates of memory formation.

Animals↗

Ventral pallidum self-stimulation: a moveable electrode mapping study.

The distribution of electrical self-stimulation (ESS) foci within the ventral pallidum (VP) was mapped using moveable electrodes in rats. The function relating ESS bar-pressing rate to the frequency of cathodal rectangular pulses (0.4 mA and 0.1 ms) was obtained for several positions of a moveable electrode in the VP and in the various adjacent to VP nuclei. The rate-frequency functions were fitted to a sigmoid model to obtain the asymptotic rate and threshold frequency. ESS was found in almost all (98%) VP sites tested and to a lesser degree (66%) in the surrounding areas (namely globus pallidus and caudate). Depending on the VP site, maximum rates varied from 14 to 85 bar presses/min, whereas threshold frequencies varied from 10.2 to 36.4 pulses/train; no correlation between these two aspects of ESS was found. Extra-pallidal areas contained less low-frequency threshold sites compared to VP. The lowest threshold found in the VP was slightly higher than that usually obtained for the most rewarding brain areas (VTA, dorsal raphé, LH, amygdala), which suggests that the VP represents an important structure for reward. Furthermore the threshold frequencies were found to decline along the rostrocaudal axis of the VP which supports the view that the VP is heterogeneous in regard to reward related functions.

Amygdala↗

The effects of lesions of the entorhinal cortex and the horizontal nucleus of the diagonal band of broca upon performance of a spatial location recognition task.

The effects of lesions of the horizontal nucleus of the diagonal band of Broca (HNDB) and the entorhinal cortex (ENTO) upon performance of a spatial location recognition task were assessed. Two hypotheses of entorhinal cortex function were considered in interpretation of the data. The first hypothesis assumes that the anatomical proximity and interconnections between the hippocampus and the entorhinal cortex are indicative of a functional similarity. The second hypothesis recognizes the interconnections of the hippocampus and the entorhinal cortex, but posits that the two structures have distinct functions. The hippocampus receives input primarily from the medial septum (MS), whereas the entorhinal cortex receives input from the HNDB and the MS. The study attempted to assess the function of the ENTO and the HNDB in a spatial location recognition task with the intent of comparing the findings to previous research of the hippocampus and MS. Performance of animals with HNDB lesions and the ENTO lesions was at chance levels for all serial positions. Results indicated that there are functional differences between the HNDB and other basal forebrain structures. Further, the performance deficit of the ENTO group does not differ from that observed following lesions of the hippocampus.

Animals↗

The nucleus accumbens in monkeys (Macaca fascicularis): I. The organization of behaviour.

A behavioural comparison was made between six unoperated control monkeys and six monkeys which received bilateral ibotenic acid lesions of the nucleus accumbens. Two of the control monkeys were subsequently given bilateral lesions of the anterior cingulate and medial frontal cortex (areas 24, 25 and 32) and were retested on the behavioural tasks. The NA lesioned monkeys, but not the anterior cingulate lesioned monkeys, were significantly impaired on a hoarding task in which they were required to remove 18 peanuts from their shells and store them in their cheek pouches. These same monkeys were not impaired when the nuts were presented without shells. Evidence is provided which suggests that this deficit is not motivational or due to gross motor impairments. A second task in which the animals were required to search through four boxes to retrieve food revealed a decrease in the tendency for the NA and cingulate lesioned animals to use an organized pattern of searching. Both groups were found to return to a previously opened box more often than controls. However, neither group showed signs of perseverative behaviour. Data from a ten-box version of this task suggest that these return errors were not due to a decrease in working memory. Together these studies suggest that both the NA and the anterior cingulate cortex contribute to the ability to organize behaviour temporally and spatially.

Animals↗

Development of hippocampal specialisation in two species of tit (Parus spp.).

Food storing birds have been shown to have a larger hippocampus, relative to the rest of the telencephalon, than do non-storers. A previous study reported that this difference in relative hippocampal volume is not apparent in a comparison of nestling birds, but emerges after birds have fledged. This conclusion was based on a comparison of a storing and a non-storing species in the corvid family. The present study compared another storer/non-storer pair of species in order to test whether the results of the previous study can be replicated in another family of birds. The volumes of the hippocampal region and remainder of the telencephalon were measured and estimates of neuron size, density and total number in the hippocampal region were made for nestlings and adults of the food-storing marsh tit Parus palustris and non-storing blue tit Parus caeruleus. Relative hippocampal volume did not differ between nestlings of the two species, whilst the relative hippocampal volume of adult marsh tits was greater than that of blue tits. The difference between adults arose because in marsh tits but not blue tits, adults had a significantly larger relative hippocampal volume than did nestlings. Neuron density was significantly higher in both species in nestlings than in adults and adult blue tits had fewer neurons than did adult marsh tits. The results of this study are largely consistent with the earlier study comparing a storing and non-storing species of corvid, suggesting that the observed patterns may reflect a general difference between storers and non-storers in the development of the hippocampal region.

Aging↗

Making two movements at once: impairments of movement, posture, and their integration underlie the adult skilled reaching deficit of neonatally dopamine-depleted rats.

Adult rats depleted bilaterally of dopamine in infancy display a profound impairment in skilled forelimb use in reaching for food. This impairment was investigated using end-point measures of reaching success, movement analysis, and kinematic measures. The rats made few successful reaches in either an easy or a difficult reaching test. Their reaches were characterized by many attempts in which trajectories of the limb were irregular and the movements were slow. Their lack of success was related in part to an impairment in making component movements of the reach, including aiming, pronating, grasping, and supinating the paw and in releasing the food pellet. It was also related to an inability to adjust posture as the limb was voluntarily moved toward the food. The results are consistent with the hypotheses that the basal ganglia, including its dopamine innervation, is important for enabling voluntary movements and postural adjustments and perhaps also the simultaneous performance of two movements at the same time.

Aging↗