Object exploration and reactions to spatial and nonspatial changes in hooded rats following damage to parietal cortex or hippocampal formation.

Hooded rats with bilateral lesions of the anterior part of the hippocampal formation (HIP), anterior region of the posterior parietal cortex (APC), or posterior region of the posterior parietal cortex (PPC) were compared with controls for their exploration of 5 objects in an open field, habituation of locomotion and object investigation, and response to spatial and nonspatial change. First, all groups displayed habituation of both locomotor and exploratory activity. Second, controls selectively reexplored displaced objects, and APC-lesioned rats reexplored all objects, whereas PPC- and HIP-lesioned rats failed to react to the spatial change. Third, a novel object induced reexploration in all groups. Results are consistent with the roles of the HIP and PPC in spatial information processing. Moreover, the APC and PPC are involved in attentional effortful processing and visuospatial information processing necessary for spatial representation, respectively. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

How do room and apparatus cues control navigation in the Morris water task? Evidence for distinct contributions to a movement vector.

The present study compared the relative influence of location and direction on navigation in the Morris water task. Rats were trained with a fixed hidden or cued platform, and probe trials were conducted with the pool repositioned such that the absolute spatial location of the platform was centered in the opposite quadrant of the pool. Rather than swimming to the platform location, rats swam in the direction that was reinforced during training, resulting in navigation to the relative location of the platform in the pool and search at the appropriate distance from the pool wall. Pool relocation tests revealed disruptions in cued navigation if the cued platform remained at the absolute location, whereas no disruption was observed if the platform remained at the relative location (same direction). The results indicate that direction holds greater influence than does location and further demonstrate that this observation is not altered by the amount of training or time on the platform. The authors propose that navigation in the water task involves a movement vector in which the distal cues and apparatus provide direction and distance information, respectively. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential effects of selective immunotoxic lesions of medial septal cholinergic cells on spatial working and reference memory.

The effect of injection into the medial septum of a toxin selective for cholinergic neurons, 192 IgG-saporin, was examined in rats trained to perform 2 versions of the radial 8-arm maze task. Rats were first trained to perform a task with varying delays (0, 1, 2 min) imposed between the 4th correct arm choice and access to all 8 arms. Lesioned rats made significantly more errors in the first 4 choices compared with controls and significantly more errors after delays; however, this effect was not delay dependent. Rats were then trained on a different version of this 8-arm maze task in which they learned to avoid 2 arms that were never baited. There was no treatment effect on acquisition of this task. These data are consistent with the hypothesis that the cholinergic projection to the hippocampus facilitates the acquisition of information into the system responsible for short-term memory for locations visited (spatial working memory) but is not involved in retention of this information. It also appears to play no role in either the acquisition or retention of place-nonreward associations (spatial reference memory). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Preweanling rats solve the Morris water task via directional navigation.

Adult rats show a preference for directional navigation over place navigation in the Morris water task. Here, the authors investigated whether preweanling rats with a newly developed ability to perform the water task also solve the task via directional navigation. After 24-day-old rats were trained to find a hidden platform in a fixed spatial location, a no-platform probe trial was conducted with the pool either in the same position as that used during training (no shift group) or shifted to a new position in the room (shift group). The authors found that rats in the shift group did not search for the platform at its absolute spatial location but rather navigated in the same direction that the platform was located during training and searched at the correct distance from the pool wall, resulting in a search at a location that was never trained. This pattern of results suggests that young rats learn to solve the water task by navigating in a particular direction rather than navigating to a precise place--a finding that may have implications for understanding hippocampal development. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Cholinergic receptor blockade in the rat impairs locale but not taxon strategies for place navigation in a swimming pool.

Normal Long-Evans hooded rats and rats drugged with atropine sulfate (10–200 mg/kg, intraperitoneally), a cholinergic muscarinic blocker, were evaluated in the Morris water task for their use of spatial navigation strategies. Atropine-treated Ss were impaired on a place response of swimming to a platform hidden in a pool filled with opaque water. With extended training, they did learn the place response, though not with the precision of controls. Acquisition could not be accounted for by habituation to the drug. In contrast with the acquisition deficit, pretrained Ss were relatively unimpaired by the drug. Atropine-treated Ss were not impaired in acquisition or retention of a cue task (swimming to a visible platform) or a position response task (turning to locate a platform). Atropine-treated Ss were unable to acquire a place learning set or to perform a learning-set response that they had acquired when undrugged. The impairments following atropine were characteristic of a deficit in the use of a locale strategy (rapid use of relational properties of distal cues), whereas their successes were characteristic of the use of taxon strategies (cue or position responses). Results suggest that locale systems of navigation are more importantly dependent on cholinergic brain mechanisms than taxon strategies. The resistance of preacquired place responses to atropine suggests that normally rats acquire place responses by using a locale strategy but effect rehearsed responses by using taxon strategies. (46 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential effect of ketamine on the reference and working memory versions of the Morris water maze task.

The assumption that blockade of long-term potentiation by N-methyl-D-aspartate antagonists interferes with spatial memory was supported by experiments showing that 15 mg/kg ketamine impairs acquisition of navigation to a hidden platform but not to a visible platform. Higher doses were required to impair retrieval of overtrained place navigation. In a working memory version of the task, retrieval latencies were shorter than acquisition latencies with 4- to 15-min but not with 30- and 60-min delays. Latent learning was only effective with the 4-min delay. Ketamine prolonged the initial search of the hidden platform at 3 mg/kg and impaired latent learning but not active acquisition at 1.5–20 mg/kg. Comparison of behavioral and synaptic effects of ketamine suggests that long-term potentiation is a necessary condition, but not a sufficient condition, for acquisition of place navigation, because search strategy and latent place learning are impaired by ketamine doses not interfering with this synaptic phenomenon. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Working memory, response selection, and effortful processing in rats with medial prefrontal lesions.

Examined the effects of lesions of the prelimbic area of the prefrontal cortex on acquisition and retention of nonmatching (NMTS) and matching-to-sample (MTS) tasks. 64 male rats participated. Both tasks involved a reference and a working memory component, but only working memory was impaired by the lesions. A comparison of the 2 tasks revealed quantitatively similar deficits in postoperatively trained rats. In preoperatively trained rats, however, the deficits were more important in the MTS task than in the NMTS task. In addition, an effect of interference between successive trials was observed in the NMTS task but not in the MTS task. Perseverative tendencies were observed in the MTS task only. Results suggest that prefrontal lesions induce working memory deficits as a result of poor temporal encoding and increased susceptibility to interference, and impair effortful processing, such as that engaged in response selection mechanisms. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Cholinergic receptor blockade produces impairments in a sensorimotor subsystem for place navigation in the rat: Evidence from sensory, motor, and acquisition tests in a swimming pool.

In Experiment 1, control rats and rats treated with atropine sulfate or atropine methyl nitrate (50 mg/kg) were trained to escape to a visible platform from different starting points in a swimming pool. All groups learned the task by concomitantly developing position responses, by orienting according to room cues, and by orienting to the platform, but probe trials showed that the atropine-sulfate group made more use of the platform as a local cue and made less use of distal cues than did the other groups. The atropine-sulfate group also made fewer searches during acquisition, made fewer searches when the platform was removed on probe trials, and were less responsive to novel cues placed above and around the pool. Swim speed, as estimated by the distance swum on probe trials, was also greater in atropine-sulfate-treated rats. The postulate that rats treated with atropine sulfate preferentially guide swimming by using position responses and local cues was tested in Experiment 2 by comparing their performance with that of control rats in a place task in which the target platform was hidden (no local cues present) and in a similar place task in which the target platform was visible, as was a second incorrect platform that sank when climbed upon (two competing local cues present). Although both tasks were acquired by the control and drugged rats, the two-platform task, as predicted, was comparatively more difficult for the atropine-treated rats. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Medial prefrontal lesions in the rat and spatial navigation: Evidence for impaired planning.

Rats with medial prefrontal cortical lesions were tested in a modified water maze navigation task. In Stage 1, the rats were trained to locate a hidden platform from a single start location. They were then subjected to a series of trials during which a second start position was used (Stage 2). In Stage 3, the rats had to navigate to a new goal location from the 2 experienced start positions. Stage 4 required the rats to navigate to the same goal as in Stage 3, starting from 4 distinct positions. Finally, a single probe trial with no platform was conducted. Rats with prefrontal lesions were impaired only during Stage 4. This deficit was specific to the 2 start positions newly introduced during this stage, suggesting a dysfunction of planning processes. This impairment might result from a working memory deficit, precluding the animal from forming an adequate representation of the whole course of movements required to reach the platform. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sex differences in performance in the Morris water maze and the effects of initial nonstationary hidden platform training.

Sex differences in rats' performance on a stationary hidden-platform task (spatial task) in the Morris water maze and the effects of initial nonstationary hidden platform training (NSP training) were examined. The NSP training was designed to familiarize rats with the general requirements of the water-maze task without providing spatial information. NSP training led to faster acquisition and improved retention of the subsequent spatial task in both males and females. There was a sex difference favoring males on acquisition and retention of the spatial task only in rats that had not received previous NSP training. Moreover, there was an apparent reversed sex difference favoring females on some measures of spatial performance in NSP-trained rats. These results suggest that performance on the water-maze task, including the expression of sex differences, can be altered by previous familiarization with nonspatial aspects of the task. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Comparison of spatial learning in woodmice (Apodemus sylvaticus) and hooded rats (Rattus norvegicus).

The spatial learning of woodmice and rats was examined in two different experiments. In the Morris place navigation task, the woodmice were slower to escape and took more circuitous routes than did the rats. However, in a special probe trial their accuracy in swimming repetitively through the position of the absent training platform was comparable to that of the rats. In the second experiment, subjects were allowed to escape from a large arena through a hole connected with their home cage. Rats and woodmice learned this task in few trials. In a special trial with no connected hole, subjects of both species showed accurate searching behavior in and around the hole of the training location. However, only the rats developed direct approaches in a systematic manner. These differences are discussed in the contexts of the species specific constraints that might affect the acquisition of direct approaches in a wild species. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Place navigation by rats in a swimming pool.

Describes characteristics of place navigation by rats in the Morris water task. Ss learned to find a small invisible platform in a large, circular swimming pool to escape from cool milk. A variety of spatial localization strategies, including spatial mapping, response sequencing, and distal cue navigational strategies, were demonstrated. Using variants of this task, the following was demonstrated: (a) Rats very readily learned true mapping strategies, being able to swim directly to the invisible platform from anywhere in the pool after only a few trials; this ability was not dependent on navigating by specific distal cues nor on starting from a familiar place. (b) Ss acquired information that facilitated subsequent place navigation merely by viewing the room from the location of the invisible goal. (c) The location of the invisible platform was remembered extremely well for several weeks. (d) The availability of only distal auditory beacons permitted acquisition of an accurate spatial mapping strategy. (e) A single S could acquire both mapping and nonmapping strategies in the swimming pool and apply them when required by the situation. Results emphasize the utility of the Morris water task and demonstrate some of the basic features of mapping and nonmapping strategies in solving a variety of spatial problems. (French abstract) (18 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Transfer of Spatial Behavior Between Different Environments: Implications for Theories of Spatial Learning and for the Role of the Hippocampus in Spatial Learning.

In 3 experiments, rats were required to find a submerged platform located in 1 corner of an arena that had 2 long and 2 short sides; they were then trained to find the platform in a new arena that also had 2 long and 2 short sides but a different overall shape. The platform in the new arena was easier to find if it was in a corner that was geometrically equivalent, rather than the mirror image, of the corner where it had previously been located. The final experiment revealed that hippocampal lesions impaired rats' ability to find the platform in these arenas. The results suggest that rats did not use the overall shape of the arena to locate the platform but relied on more local cues and that the hippocampus plays a role in navigation based on these cues. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Differentiating visual and kinesthetic imagery in mental practice.

Through two experiments, the study sought to emphasize the usefulness of the visual and kinesthetic imagery in mental practice. In Experiment 1, it was hypothesized that when the task to be learned through mental practice necessitates the reproduction of a form by drawing, the visual image, which provides a wide span of apprehension, is more suitable than the kinesthetic image. On the other hand, the kinesthetic image that supplies inputs from the muscles' positions and movements should be more appropriate for the acquisition of the duration of the drawing. In Experiment 2, it was hypothesized that the task, transformed into a motor task necessitating minute coordination of the two hands, would benefit more from kinesthetic imagery. To have optimal control over what was actually experienced during mental practice, the participants' imagery skills were measured. The participants also benefited from prior imagery training. The results demonstrate that when using mental practice to initially acquire a task, visual imagery is better for tasks that emphasize form while kinesthetic imagery is better for those tasks that emphasize timing or minute coordination of the two hands. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The relative influence of place and direction in the Morris water task.

Previous work from our laboratory has demonstrated that rats display a preference for directional responding over true place navigation in the Morris water task. The present study evaluated the range of situations in which this preference is observed and attempted to identify methods that favor navigation to the precise location of the escape platform in the room. A preference for directional responding over place navigation was observed in a wide range of procedures that included providing extensive training (Experiment 1), providing only platform placement experience in the absence of active swim training (Experiment 2), training navigation to multiple platform locations in a moving platform variant of the task (Experiment 3), and explicitly training navigation to a precise location in the room, versus navigation in a particular direction, regardless of the pool's position in the room (Experiments 4-5). A modest preference for navigation to the precise spatial location of the platform was observed when the pool wall was virtually eliminated as a source of control by filling it to the top with water (Experiment 6). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Absence of snapshot memory of the target view interferes with place navigation learning by rats in the water maze.

Contribution of visual and nonvisual mechanisms to spatial behavior of rats in the Morris water maze was studied with a computerized infrared tracking system, which switched off the room lights when the S entered the inner circular area of the pool with an escape platform. 10 naive male rats trained under light–dark (LD) conditions found the escape platform more slowly than 10 male rats trained in permanent light (LT). After group members were swapped, the LT-pretrained rats found under LD conditions the same target faster and eventually approached latencies attained during LT navigation. Performance of LD-trained rats deteriorated in permanent darkness (PD) but improved with continued PD training. Thus LD navigation improves gradually by procedural learning (extrapolation of the start-target azimuth into the zero-visibility zone) but remains impaired by lack of immediate visual feedback rather than by absence of the snapshot memory of the target view. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Visits to starts, routes, and places by rats (Rattus norvegicus) in swimming pool navigation tasks.

In 2 studies, 40 young adult female Sprague-Dawley rats were trained to escape to visible or to hidden platforms in a swimming pool and then given probe trials, which required that they searched for a platform that had been removed or repositioned. Results indicate that to solve the tasks, Ss simultaneously used a number of behavioral strategies including position responses, cue responses, and place responses. On the probe trials, they not only displayed behaviors that were reinforced during training but also displayed novel behaviors. Ss trained on the place task (hidden platform) made more swims across the platform's previous location, whereas Ss trained on the cue task (visible platform) made more returns to previously used start points. Increases in the number of start points produced more returns to start points, whereas increases in the number of platform locations produced more searches for platforms. It is concluded that rats make coextensive use of all relevant strategies to solving spatial navigation tasks and that their search patterns on probe trials reflect both previously reinforced behaviors and novel unconditioned search behaviors. (34 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Evidence for a shift from place navigation to directional responding in one variant of the Morris water task.

Previous work from our laboratory has demonstrated that rats display a preference for directional responding over place navigation in a wide range of procedural variants of the Morris water task (Hamilton, Akers, Weisend, & Sutherland, 2007; Hamilton et al., 2008). A preference for place navigation has only been observed when the pool is reduced as a cue by filling it with water. Studies using dry land mazes have suggested that rats place navigate early in training and later switch to other forms of responding (e.g., motor). The present study evaluated whether rats switch from place navigation to directional responding in the “full-pool” variant of the water task. Rats were given 12, 24, or 36 hidden platform training trials. Probe trials with the pool repositioned in the room revealed a preference for place navigation in rats given 12 trials, an equal division of response preferences in rats given 24 trials, and a preference for directional responding in rats given 36 trials. These results indicate that the early preference for place navigation in the full-pool water task is transient and yields to a preference for directional responding with continued training. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Path integration and lesions within the head direction cell circuit: Comparison between the roles of the anterodorsal thalamus and dorsal tegmental nucleus.

Experiments were designed to determine whether 2 regions of the head direction cell circuit, the anterodorsal thalamic nucleus (ADN) and the dorsal tegmental nucleus (DTN), contribute to navigation. Rats were trained to perform a food-carrying task with and without blindfolds prior to receiving sham lesions or bilateral lesions of the ADN or DTN. ADN-lesioned rats were mildly impaired in both versions of the task. DTN-lesioned rats, however, were severely impaired and showed reduced heading accuracy in both task versions. These findings suggest that although both the DTN and ADN contribute to navigation based on path integration and landmarks, disruption of the head direction cell circuit at the level of the DTN has a significantly greater effect on spatial behavior than lesions of the ADN. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Unilateral injury of posterior parietal cortex and spatial learning in hooded rats

The influences of bilateral or unilateral injuries within the posterior parietal cortex (PPC) upon spatial learning in a water maze were examined in three experiments. Place-learning and response-learning were investigated in a four-alley 'Greek-cross' shaped water maze with extra-maze visual cues available. No differences were detected on any of several measures sensitive to learning between the lesion groups on the place-learning task. Microanalysis of behavior within trials revealed that animals with either bilateral or right unilateral PPC injuries committed significantly more total errors, initial alley entrance ('reference memory') errors, and re-entry ('working memory') errors in the response-learning paradigm than did either the control or left PPC-injured rats. No differences were detected between the latter two groups on these measures. Unilateral lesions resulted in asymmetrical placing responses ipsilateral to the injury 10 days after surgery whereas bilateral injuries resulted in asymmetrical placing with mixed directionality. The acquisition of the response-learning problem in the absence of visual cues was studied on animals prepared with unilateral lesions and housed post-operatively either in isolation or in a 'complex environment.' In the absence of visual cues both right and left PPC-injured rats committed more errors than sham controls, and differential post-surgical housing did not attenuate these impairments. These same animals were trained on the landmark navigation task. Although no differences appeared between the lesion groups, a generalized but transient facilitation of learning was observed in animals housed in the 'complex' environment. Unilateral injuries placed in sham controls failed to disturb retention of the landmark navigation strategy. Because none of the PPC-injured animals were deficient in the landmark task, a result which is contrary to observations in other laboratories, the influence of post-surgical recovery interval upon acquisition of the landmark navigation strategy was explored. Animals were prepared with right PPC injuries and trained following either a 5 or 35 day recovery interval. Only those animals limited to the short recovery interval proved to have a spatial deficit in the landmark task. It is concluded that injuries in the PPC of either hemisphere disturb egocentric spatial functions. However, animals with left PPC injuries are able to compensate by using allocentric visual cues if they are available. It is due to the special role played by the right PPC in complex visuospatial functions that animals with this injury are unable to compensate.