Severity of Spatial Learning Impairment in Aging: Development of a Learning Index for Performance in the Morris Water Maze methods
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Measures Traditionally Used for Behavioral Analysis in the Water Maze
An additional shortcoming of the traditional measures used for training trials is illustrated in. Two training trial records are shown. In the record on the left, the path is widely distributed, with little focused search in any particular region of the maze. In the record on the right, the search is focused more in proximity to the escape platform. The escape latencies are virtually identical for these two trials. Computing the path length to reach the escape platform also does not distinguish performance because path length per se does not contain information about the spatial distribution of the rat's search relative to the goal. Indeed, the path length to locate the escape platform is slightly shorter in A (573 cm) than in B (590 cm). However, performance on the two trials differs quite markedly if the animal's location is used to compute distance from the goal during...
Measures Traditionally Used for Behavioral Analysis in the Water Maze
Using a measure of proximity to the goal can also be advantageous in the analysis of probe trial performance. The measures commonly used to assess performance on probe trials are designed to reflect the spatial bias of an animal's search pattern. These typically include either the percentage of time that is spent or the percentage of the total path length during the search that is traversed within the training quadrant and, as a second measure, so-called platform crossings. A platform crossing occurs when the animal's path traverses the precise location where the escape platform was positioned during training. shows three sample paths from probe trials. The accuracy of the animal's search relative to the escape platform location differs substantially in these three records. However, each of the measures commonly used to analyze probe trials is inadequate to resolve t...
Measures Traditionally Used for Behavioral Analysis in the Water Maze
Our research and similar studies conducted in a number of other laboratories have shown that the water maze task is sensitive to age-related learning impairment but that striking individual differences are apparent in aged animals (;; ). Thus some aged rats appear to learn the spatial task as proficiently as young animals, whereas other aged rats exhibit marked impairment. Because probe trial data are essential for determining whether performance is based on a search strategy similar to that normally used by young rats, much of our previous research used a protocol in which brief probe trials were interpolated during the course of training. In numerous studies we found that young rats typically reached a criterion performance on probe trials interpolated after 10-15 training trials, whereas fewer than half of the aged rats performed in a comparable manner. illustrates this fin...
Method
Behavioral results from 96 young (4-6 months) and 128 aged (25-27 months) pathogen-free Long-Evans male rats are presented. The aged subjects were obtained as retired breeders at 8-9 months of age (Charles River Laboratories, Raleigh, NC). The young rats were obtained at 90 days of age from the same source and entered the experiment at 4-6 months of age. The young and aged rats were behaviorally tested in replications that used 6-12 young rats and 10-18 aged rats during the months spanning January 1, 1991 through April 30, 1992. Routine screening for viral antibodies and necropsies at the time of sacrifice were performed to ensure the health of aged subjects. Prior to data analysis, a number of aged animals (n = 14) were excluded on the basis of necrospy findings (pituitary tumors, etc.). Tests for viral antibodies were routinely negative, which con...
Procedure
For analysis of spatial learning, measures of path length and latency to escape were analyzed using average performance for blocks of five training trials. Probe trial data were assessed for acquisition of criterion performance, that is, at least 35% of the duration of the probe trial spent in the training quadrant and at least two platform crossings. Analysis of cue training performance was based on latency and path length to escape. These analyses thus used traditional measures for behavioral assessment in the water maze task.
Procedure
The proximity of the animal's position with respect to the goal was used to provide several new analyses of training trial and probe trial performance. The proximity measure was obtained by sampling the position of the animal in the maze (10 times per second) to provide a record of its distance from the escape platform in 1-s averages. For both probe trials and training trials, a correction procedure was implemented so that trial performance was relatively unbiased by differences in distance to the goal from the various start locations at the perimeter of the pool. In making this correction we calculated the average swimming speed for each trial (path length/latency). Then the amount of time required to swim to the goal at that speed from the start location used on the trial was removed from the record prior to computing trial performance; that is, cumulative distance was used o...
Behavioral Analysis Using a Proximity Measure
The newly developed proximity measure was used to analyze both place training trials and probe trial performance. shows the performance of young rats during training trials and compares the customary path length measure with a new measure based on proximity to the platform that is referred to as search error. As described in the Method section, search error represents the corrected cumulative distance from the platform. Although numerical values on the two measures are in the same range at the end of training (averaging 181 cm and 225 cm for path length and search error, respectively), the magnitude of change from the initial level of performance is much greater for search error, F(95,59) = 95.6,p <.0001. The amount of change was significantly greater for search error than for path length in all adjacent blocks (p <.005), with the exception of the change from Block 5 to Block 6 (p <....
Behavioral Analysis Using a Proximity Measure
An analysis was also done on these data to determine whether proximity to the target platform location showed evidence for extinction over the 30-s probe trial interval, that is, whether there was a tendency for rats to abandon their search toward the end of the trial. In this analysis, the average proximity during seconds 11-20 of the interval was compared to the mean proximity during the final 10 s for each probe trial. The middle segment of the interval was selected for this comparison because the correction procedure removed variable portions of the initial segment (depending on swim speed) from calculation of the proximity measure. The analysis of probe trial performance indicated no reliable difference between segments of the interval for Probe Trials 1-5. On the final probe trial, rats searched in closer proximity to the target location during the middle segment tha...
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An additional shortcoming of the traditional measures used for training trials is illustrated in. Two training trial records are shown. In the record on the left, the path is widely distributed, with little focused search in any particular region of the maze. In the record on the right, the search is focused more in proximity to the escape platform. The escape latencies are virtually identical for these two trials. Computing the path length to reach the escape platform also does not distinguish performance because path length per se does not contain information about the spatial distribution of the rat's search relative to the goal. Indeed, the path length to locate the escape platform is slightly shorter in A (573 cm) than in B (590 cm). However, performance on the two trials differs quite markedly if the animal's location is used to compute distance from the goal during the trial: By the method described in the caption to, the cumulative distance from the goal during the search in Trial A is 1,835 cm, whereas it is 781 cm for Trial B.
Using a measure of proximity to the goal can also be advantageous in the analysis of probe trial performance. The measures commonly used to assess performance on probe trials are designed to reflect the spatial bias of an animal's search pattern. These typically include either the percentage of time that is spent or the percentage of the total path length during the search that is traversed within the training quadrant and, as a second measure, so-called platform crossings. A platform crossing occurs when the animal's path traverses the precise location where the escape platform was positioned during training. shows three sample paths from probe trials. The accuracy of the animal's search relative to the escape platform location differs substantially in these three records. However, each of the measures commonly used to analyze probe trials is inadequate to resolve the differences in search accuracy that are evident in this illustration. For example, roughly 50% of the total path length was located in the training quadrant in both A and B, which indicates comparable performance by this measure on these trials. The platform-crossing measure distinguishes A from...
Our research and similar studies conducted in a number of other laboratories have shown that the water maze task is sensitive to age-related learning impairment but that striking individual differences are apparent in aged animals (;; ). Thus some aged rats appear to learn the spatial task as proficiently as young animals, whereas other aged rats exhibit marked impairment. Because probe trial data are essential for determining whether performance is based on a search strategy similar to that normally used by young rats, much of our previous research used a protocol in which brief probe trials were interpolated during the course of training. In numerous studies we found that young rats typically reached a criterion performance on probe trials interpolated after 10-15 training trials, whereas fewer than half of the aged rats performed in a comparable manner. illustrates this finding with composite results from a number of previous studies conducted in our laboratory. We used these data to classify aged rats into subgroups designated as unimpaired and impaired in order to examine whether brain aging distinguishes those rats with spatial learning deficits from their aged coh...
Behavioral results from 96 young (4-6 months) and 128 aged (25-27 months) pathogen-free Long-Evans male rats are presented. The aged subjects were obtained as retired breeders at 8-9 months of age (Charles River Laboratories, Raleigh, NC). The young rats were obtained at 90 days of age from the same source and entered the experiment at 4-6 months of age. The young and aged rats were behaviorally tested in replications that used 6-12 young rats and 10-18 aged rats during the months spanning January 1, 1991 through April 30, 1992. Routine screening for viral antibodies and necropsies at the time of sacrifice were performed to ensure the health of aged subjects. Prior to data analysis, a number of aged animals (n = 14) were excluded on the basis of necrospy findings (pituitary tumors, etc.). Tests for viral antibodies were routinely negative, which confirmed the pathogen-free status of the animals used in the study.
For analysis of spatial learning, measures of path length and latency to escape were analyzed using average performance for blocks of five training trials. Probe trial data were assessed for acquisition of criterion performance, that is, at least 35% of the duration of the probe trial spent in the training quadrant and at least two platform crossings. Analysis of cue training performance was based on latency and path length to escape. These analyses thus used traditional measures for behavioral assessment in the water maze task.
The proximity of the animal's position with respect to the goal was used to provide several new analyses of training trial and probe trial performance. The proximity measure was obtained by sampling the position of the animal in the maze (10 times per second) to provide a record of its distance from the escape platform in 1-s averages. For both probe trials and training trials, a correction procedure was implemented so that trial performance was relatively unbiased by differences in distance to the goal from the various start locations at the perimeter of the pool. In making this correction we calculated the average swimming speed for each trial (path length/latency). Then the amount of time required to swim to the goal at that speed from the start location used on the trial was removed from the record prior to computing trial performance; that is, cumulative distance was used on training trials and average distance from the goal was used on probe trials. By this method, scores obtained with the proximity measure are designed to reflect search error; they represent deviations from an optimal search, that is, from a direct path to the goal.
The newly developed proximity measure was used to analyze both place training trials and probe trial performance. shows the performance of young rats during training trials and compares the customary path length measure with a new measure based on proximity to the platform that is referred to as search error. As described in the Method section, search error represents the corrected cumulative distance from the platform. Although numerical values on the two measures are in the same range at the end of training (averaging 181 cm and 225 cm for path length and search error, respectively), the magnitude of change from the initial level of performance is much greater for search error, F(95,59) = 95.6,p <.0001. The amount of change was significantly greater for search error than for path length in all adjacent blocks (p <.005), with the exception of the change from Block 5 to Block 6 (p <.11).
An analysis was also done on these data to determine whether proximity to the target platform location showed evidence for extinction over the 30-s probe trial interval, that is, whether there was a tendency for rats to abandon their search toward the end of the trial. In this analysis, the average proximity during seconds 11-20 of the interval was compared to the mean proximity during the final 10 s for each probe trial. The middle segment of the interval was selected for this comparison because the correction procedure removed variable portions of the initial segment (depending on swim speed) from calculation of the proximity measure. The analysis of probe trial performance indicated no reliable difference between segments of the interval for Probe Trials 1-5. On the final probe trial, rats searched in closer proximity to the target location during the middle segment than in the final 10-s segment (p <.005). Thus, rats did not tend to abandon their search toward the end of the probe trial throughout most of the training protocol: The average proximity measure appears to be representative of the spatial distribution of the search unconfounded by any extinction effe...
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