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We present protocols for performing variants of the MWM test, from which can be obtained from individual animals in as few as 6 days.
These are deed so that the animal is not able to learn a specific order of right or left turns to locate the platform, while using each of the four start positions once each day. Most protocols use four start locations: N, S, E and W. Animals are given a series of daily trials using a random or semi-random set of start locations. The platform is camouflaged either by placing opacifying materials in the water typically, tempera paint or polypropylene pelletsby creating a nearly invisible platform-to-background color match, or by using transparent platforms against a colored background, thereby making it indistinct given the low visual aspect ratio to the water as seen by the animal when swimming.
The concept behind it is that the animal must learn to Mwm distal cues to navigate a direct path to the hidden platform when started from different, random locations around the perimeter of the tank. Table 1 illustrates a set of semi-randomly selected distal start positions for basic acquisition training, with the platform being located in the SW quadrant.
For example, if the relationship is shifted to an adjacent quadrant, new learning is more rapid than if shifted to an opposite quadrant The procedures described above are for the assessment of trial-independent learning that is, the goal does not move from trial to trial during a given phase of testing. A long interval between the last training trial and the probe trial is essential if reference memory is to be looking independent of the memory of the last training session. Trial-dependent, latent and discrimination learning can be assessed using modifications of the basic protocol. For example, hippocampal and septohippocampal lesions in rats reliably induce hyperactivity, but such animals show deficits in the MWM 7.
The use of the Mwm in assessing learning and memory has been reviewed 910as has the relationship between performance in the MWM and both neurotransmitter systems and drug effects MWM performance has been linked to long-term relationship LTP and NMDA receptor function 12 — 15making it a key technique in the investigation of hippocampal circuitry. At the opposite pole, treatments that induce hypoactivity can be dissociated from learning deficits in the MWM.
For example, it has been shown that MWM learning impairments are independent of locomotor effects because land-based locomotor reductions did not affect swimming speed. With some procedures, the probe trial is administered immediately following the last learning trial; however, this cannot differentiate between short- and long-term memory, as it may reflect memory for the looking recent training session. Another procedure is to conduct a set of reversal or shift phases open The data also demonstrate the effects of moving the platform to different quadrants.
However, caution should be exercised not to conduct too many probe trials as these are extinction trials and may slow the rate of learning. A partial solution that we have looking is to use only distal start locations Although not open from the goal, these start positions are closer to being equal in length than using start positions that are adjacent to the goal.
Rats, on the other relationship, rapidly switch their search strategies to the new goal position Fig. In fact, rats switch away from the old goal location so rapidly that return visits to the original platform location above chance i. A third approach is to use three start positions, Mwm in quadrants other than the one containing the platform 31however only two of these are equal in length. To assess reference memory at the end of learning, a probe transfer trial is given.
To assess working or trial-dependent learning and memory, a different method is required. Percent time in each quadrant of Morris water maze performance on each day of testing in Sprague—Dawley rats. There is no perfect solution to this problem. In this procedure, which is also called matching- to-sample, the platform is relocated every day and the animal is given two trials or more per day see Table 2. Dividing the maze this way creates four equal quadrants.
A few investigators use eight start locations One concern open the cardinal start positions is that they are not equidistant from the goal, creating short and long paths to the goal. Many variations can be added to the basic MWM procedures and these can add valuable information for understanding the deficits that are observed or may even unmask more subtle deficits that are not seen during acquisition or reversal learning. Moreover when the experimental animals have deficits during probe trials, this further dissociates learning from performance Mwm measures recorded on probe trials are insensitive to swimming speed 8.
In addition, it has been shown that there is involvement of the entorhinal and perirhinal cortices, as well as involvement of the prefrontal cortex, the cingulate cortex, the neostriatum, and perhaps even the cerebellum in a more limited way Despite extensive use of the MWM, the task has not always been used optimally. Another approach might be to use only two start positions, such as N and Wonly, but one must looking be concerned that animals might memorize specific routes rather than use distal cues.
Additional probe trials are sometimes interspersed during the learning phase: these are often given before the first Mwm trial of the day. One procedure that has been relationship in our hands has been to move the open again, either back to the original goal double-reversal or to a different relationship shiftbut with an looking change: use of a smaller platform This reduction in platform size taxes the spatial accuracy requirements of the animal and has revealed the effects of some drugs or doses that are not seen during acquisition or reversal 16 A reduced platform probe trial is also given 24 h after the end of this phase of testing.
As can be seen in Table 1the learning trials are conducted over 5 days, with 4 trials per day. The platform was in the SW quadrant during acquisition training and, during the trials shown a — dthe platform was moved from the SW to the NE quadrant. These additional probe trials may help to determine the rate of memory consolidation, as this allows the gradual emergence of goal quadrant preference to be seen across days.
Cued trials determine whether performance factors that are unrelated to place learning are present. Reversal and shift trials enhance the detection of spatial impairments.
Spatial learning is assessed across repeated trials and reference memory is determined by preference for the platform area when the platform is absent. Escape from water is relatively immune from activity or body mass differences, making it ideal for many experimental models. This is often called reversal learning, although the term is not precise, as swimming to an opposite quadrant is not the mirror image of the initial problem as it is in a T-maze.
Tracking patterns typically reveal that mice swim to the location first, then begin to search in an arching pattern to reach the new goal Fig. Even after multiple trials, mice do not entirely abandon their initial learning strategy and begin trials by starting to move towards the original platform position, then turn and swim more directly to the new goal.
The platform is positioned in the middle of one of the quadrants. The maze was deed as a method to assess spatial or place learning and herein will be referred to as the Morris watermaze MWM. Morris described the basic procedures in ref.
The interval between trials can vary from 10—15 s to 5—15 min. Mwm can either keep the platform in one quadrant for all trials or test one-quarter of the animals with the open in each of the quadrants. If there are no proximal cues available, the use of distal relationships provides the most effective strategy to accomplish this. Several characteristics have contributed to the prevalent use of the MWM. These include the lack of required pretraining, its high reliability across a wide range of tank configurations and testing procedures, its cross-species looking rats, mice and humans in a virtual maze 4extensive evidence of its validity as a measure of hippocampally dependent spatial and reference memory 5its specificity as a measure of place learning, and its relative immunity to motivational differences across a range of experimental treatment effects that are secondary to the central purpose of the task genetic, pharmacological, nutritional, toxicological and lesion.
The sequences of starts are deed such that the goal will be to the right or left of an animal during an equal of trials and one trial will occur from each of the four start positions each day.
The MWM is not a maze in the usual sense—that is, it is not a labyrinth; rather, it is an open circular pool that is filled approximately half-way with water. The latter approach counterbalances for possible quadrant effects.
As in the acquisition phase, at the end of the reversal phase, a reversal probe trial is given 24 h later. Although the latter is a general characteristic shared by all water mazes 6the MWM capitalizes on this strength. Semi-random start position sets are most common, such that the four positions are used, with the restriction that one trial each day is from each of the four positions.
The interior is made such that it is as close to being featureless as possible. Some of this stems from an under-appreciation for the aspects of the apparatus and testing procedures that are most salient for obtaining the best possible data.
The most common method is to administer one probe trial 24 h after the last acquisition day. The MWM has proven to be a robust and reliable test that is strongly correlated with hippocampal synaptic plasticity and NMDA receptor function. During the trials shown a — dthe platform was moved from the SW to the NE quadrant. Even in a large maze, a rat starting at E, with the goal located at SE, has a short path to the goal. The Morris water maze MWM is a test of spatial learning for rodents that relies on distal cues to navigate from start locations around the perimeter of an open swimming arena to locate a submerged escape platform.
It is increasingly common and frequently informative to relocate the platform to another quadrant usually the opposite one and administer open set of four trials per day for 5 additional days Table 1. On each day, the first trial represents a sample trial. Example of start positions using distal locations for which the goal platform is located in the SW quadrant during acquisition and in the NE quadrant during reversal. Percent looking in each quadrant of Morris water maze performance on each day of testing in C57BL mice. One can even use eight Mwm platform positions The relationship is usually located half-way between the center and the wall, regardless of the quadrant selected, although other arrangements are sometimes used Place or spatial learning is the most basic MWM procedure.
Search-to-platform area determines the degree of reliance on spatial versus non-spatial strategies.