Jourmd ~4Neuro,wienceMethod~, 11 (1984) 47-60 47

Elsevier

NSM 00384

Developments of a water-maze procedure for

studying spatial learning in the rat
Richard Morris
M R C Cognitive Neuroscience Group, P.wchological Laboratory, Unit~ersiO' of St. Andrews, St. Andrews, Fif~,
(U.K.)
(Received December 22nd, 1983)
(Revised March 28th, 1984)
(Accepted April 3rd, 1984)

Key words: water-maze--spatial memory

Developments of an open-field water-maze procedure in which rats learn to escape from opaque water
onto a hidden platform are described. These include a procedure (A) for automatically tracking the spatial
location of a hooded rat without the use of attached light-emitting diodes; (B) for studying different
aspects of spatial memory (e.g. working memory); and (C) for studying non-spatial discrimination
learning. The speed with which rats learn these tasks suggests that they may lend themselves to a variety
of behavioural investigations, including pharmacological work and studies of cerebral function.

Introduction

The purpose of this paper is to describe developments of a new behavioural

procedure designed initially to study spatial localisation in the rat (Morris, 1981).
The procedure was devised in response to theoretical controversies about the neural
basis of spatial and working memory (O'Keefe and Nadel, 1978; Olton et al., 1979)
and ideas that learning about spatial relations may be fundamentally different from
other forms of associative learning.
The essential feature of the technique is as follows: rats are placed into a large
circular pool of water from which they can escape onto a hidden platform. The
platform is hidden, first by arranging that its top surface is just beneath the water
surface, and second by rendering the water opaque so that the platform is invisible.
Thus the platform offers n o local cues to guide escape behaviour. In principle, rats
could escape from the wafer by swimming randomly, or in unsystematic search paths
throughout the pool; but in practice, normal rats very quickly learn to swim directly

Corre~j~ondence: R. Morris, MRC Cognitive Neuroscience Group, Psychological Laboratory, University

of St. Andrews, St. Andrews, Fife, Scotland, U.K.

0165-0270/84/$03,00 <~ 1984 Elsevier Science Publishers B.V.

48

towards the platform from any starting position at the circumference of the pool.
The accurate directionality of their escape behaviour (and other measures of
performance) provide evidence that the rats escape by learning the spatial position
of the platform relative to distal cues. Comparison of the performance of normal
and brain-damaged rats (Morris et al., 1982; Sutherland et al., 1983), and of rats
given drugs (Sutherland et al., 1982) or cerebral neurotoxins (Hagan et al., 1983)
offers a new way of examining the neurobiology of spatial learning.
The technique of using escape from water to motivate learning has been used for
many years (Glaser, 1910; Wever, 1932; Waller et al., 1960). Water temperature, and
increases in water temperature when rats swim from one compartment into another
at a higher temperature (Woods et al., 1964) affect learning rate in simple escape
tasks. Swimming 'ability' is also related to learning rate (Woods and Holland, 1961).
In these and the present procedures, the animals do not need to be deprived of food
to motivate learning, or given extensive pretraining. The use of electric shock to
motivate escape behaviour is obviated. Rats are natural swimmers and 'take to'
water-escape tasks easily, even if they have sustained telencephalic brain lesions of
various kinds. With respect to the present task, the water provides an intramaze
environment which controls for olfactory cues and, as will be described, allows for a
variety of training procedures to dissociate learning and performance effects.
This paper describes several new developments of the 'water-maze' procedure
including: (1) a technique for tracking the paths taken by rats during swimming; (2)
variations in the training schedules to examine different aspects of spatial memory;
and (3) procedures for examining non-spatial discrimination learning.

Materials and Methods

Subjects
The results of several experiments using both male and female rats will be
described. The rats are maintained on ad libitum food and water, and housed singly
or in groups of two rats per cage.

Apparatus
Two different pools have been constructed (Fig. 1). The first was 1.32 m in
diameter x 0.60 m high (as used in Morris, 1981). The second, currently in use, is
2.14 m in diameter x 0.40 m height. Both were constructed from hardboard, at-
tached to a baseboard, and coated with fibreglass and resin. The first was then
painted white, but this coating 'flaked' and had to be frequently repainted; the
larger pool is coated with permanent white gelcoat. Each pool is filled with water
(26 + I ° C ) to depths of 0.40 m and 0.25 m, respectively, and rendered opaque by the
addition of a small quantity of milk (pure or powdered). The second pool has been
fully plumbed into the laboratory for ease of filling and draining, and mounted on a
frame so that the water is at waist height. Four points around the circumference of
the pool are arbitrarily designated North, South, East, or West and, on this basis, the
pool area divided into 4 quadrants (NW, SW, etc).
 4g

-- TV Camera

Lamps powered via rheosfof

I / ~ 1. .~
TransLucent perspex disc
Ceiling !
/ \
/ \

!
/
\
Curtains - usuoLtyopen
i ,
i

Perspex hidden Hardboard sides, coated with

platform fibregtass and white getcoat

I'~] ~ ~ Baseboard -[J [(~]

Bright lamps

- - 2.1z, m ,-
Fig. I . Cross-sectional diagram of pool, and its position in the laboratory. Note support frame to ensure

water level is at roughly waist height, and perspex disc attached to laboratory ceiling to limit field of v i e w
of TV camera.

Escape from the water is provided by means of circular rigid platforms (9 cm

diameter). Two platforms were made from sections of perspex tubing filled with
stones, they were sealed at the top to provide the platform and at the bottom to form
a baseboard. The height of these platforms is arranged to be 1 cm below, or 1 cm
above the water level; thus one platform is hidden, the other visible. A third
platform was made from a short (3 cm high) section of perspex tubing filled with
expanded polystyrene. This platform is anchored by means of nylon threads to a
metal base (section of heavy galvanised caging) which sits on the bottom of the pool.
Each platform can be moved to any position in the pool, generally at the centre of
one of the 4 arbitrary quadrants. These platforms are painted white, grey or with
black and white stripes and used in different procedures designed as 'hidden
platform', 'cueing procedure' and 'two-platform task' (Fig. 2) as described below.

A B E

Hidden ploff0rrn Cueing procedure Two platform rusk

Fig. 2. Cross-sectional diagram of pool showing 3 different arrangements of escape platforms as described
in text.
50

Tracking techniques
The simplest measure of performance in the pool is the latency to escape from the
water. As rats swim at a fairly constant speed for a given water temperature, latency
is quite sensitive to the amount that the rat has learned about the task, The original
procedure was to monitor the rat's behaviour via a video-camera placed above the
pool, relaying the output to a video-recorder and monitor. The paths taken in
escaping were then transcribed onto charts, slowing down the replay of the tapes
when necessary to achieve accuracy of transcription. From these charts, path-length
is measured using a map-measuring device. Path-directionality, defined as deviation
from the correct heading angle towards the platform, is measured at various
distances along the path with a protractor. These transcribing and measuring
procedures have been adequate but suffer from inevitable inaccuracies. A tracking
system has therefore been adopted to allow for objective analysis.
The tracking system consists of an image-analyser (HVS Ltd, VP110) coupled to
a small BBC Model B Microcomputer. The image-analyser transforms the conven-
tional video-image into an image of high-contrast edges. A threshold adjustment
allows the operator to monitor various degrees of contrast and the image-analyser
provides, by way of output, the x and y coordinate of the first high-contrast edge
found in tracking down the horizontal lines of the video picture. The x and y
coordinate is displayed and also expressed as two analogue voltage signals for relay
to the computer. These signals are sampled at the rate of 10 Hz.
With suitable reflected lighting, this system can be used to track the position of a
black hooded rat by adjusting the contrast sensitivity until the black head of the rat
is picked out against the white surfaces of the water and of the remainder of the
animal's body. Thus the image-analyser monitors the exact x and y coordinate of the
rat's head. No light-emitting diodes attached to the animal are necessary.
The image-analyser will also operate from the composite video output of a
video-recorder such as the SONY C9 Betamax recorder which provides a high signal
to noise ratio, particularly with CrO 2 tape. The animals may then either be tracked
'on-line' or from videotape. The SONY C9 also has remote infrared controls so that
a single experimenter may start and stop the videotape while handling the rat.
One complication is that the HVS image-analyser samples from an operator-de-
fined 'sampling space' shaped in the form of a square. As the pool is round, this
sampling space must be equal on both sides to the diameter of the pool leaving 4
'sectors' outside its perimeter, at the corners of the sampling square. To ensure that
these do not result in high-contrast edges 'trapping' the x, y coordinate monitoring
system, a perspex sheet which a circular disc cut from it is placed high above the
pool beneath the video-camera, positioned to delimit the view of the camera to the
full water surface only. This sheet is back-illuminated by lights powered via a
rheostat which can be adjusted until any contrast edge between the disc and the
perimeter of the opaque water surface is eliminated. The only high-contrast edge
remaining is then around the black head of the rat (Fig, 3).

Computing programs
A suite of computer programs has been written for (a) sampling, (b) processing
 51

1 2

Fig. 3. Video pictures on Monitors 1 and 2. Monitor 2 shows the high-contrast transformation of the
video image and position of cursor-line pointing to the x. y coordinate of a rat's head.

and (c) displaying the paths taken by the animal, taking advantage of the full colour
graphics of the BBC Model B computer. The x, y coordinates are sampled at 10 Hz
between start and stop signals indicating entry into the pool and arrival at the
platform, respectively. The data coordinates are stored on diskettes. The first stage
of data-processing involves a 'smoothing' algorithm which rejects false x, y coordi-
nates. If the lighting arrangements are set properly, the level of artifact is very low,
generally less than 1%. However, false coordinates do arise due to occasional
'flashes' of high-contrast on the water-surface, or due to artifacts during replay from
the video-recorder. The algorithm operates according to the principle that the rat can
only swim at certain speeds and thus any x, y coordinate pair greater than a certain
distance from a preceding coordinate pair must be false. False coordinates are
replaced with the average values of preceding and succeeding permissible coordi-
nates. The 'smoothed' data are then used to calculate path-lengths, directionality,
position in pool and other measures of performance as described below. All
programs were written in BBC Basic software, in consultation with a professional
software house (Computer One Ltd.).

Behavioural procedures

1. Acquisition and reversal

The simplest experimental procedure involves training rats to find the hidden
platform at a fixed location in space over a series of trials. Fig. 4A shows the results
of an experiment in which 16 male rats were trained to find the platform in the small
pool over 20 trials spread across 3 consecutive days (from Morris, 1981). For one
group, the platform was placed in one quadrant for all 20 trials (different quadrants
for different rats) and then moved on trial 21 to the opposite quadrant (e.g. to SE
from NW). For a second group, platform position varied from trial to trial
52

A. B.
120'
Acquisition Reversal

100"
o
: ~ PLace
',,, ~ o~ Random

b' °' o c? o- ~o o

; . . . . . . . ; . . . . . . . . ,'7' ';0;,' ';,

Trials
Fig. 4. A: latency to escape from the water over 20 trials of acquisition by Groups Place and Random.
The rapid escape by rats of Group Place cannot be due to local cues from the platform, reflected waves
etc. Group Place were then given 4 trials of 'reversal'. Note increased latency on trial 21. B: path taken on
trial 21 by one typical rat. Note initial searching in former platform location in NE. prior to finding
platform now in SW.

randomly. The initial decline in latency (e.g. trials 1 8) is due to two factors: (1) the
rats learning to swim away from the side walls and thus increasing their likelihood of
contacting the platform by chance; (2) true spatial learning about the location of the
platform. For Group Place, escape latency stabilised between 5-8 s as the rats
learned to take relatively direct paths to the hidden platform; Group Random, on
the other hand, shows only the initial decline in latency associated with overcoming
thigmotaxis. These findings show that the effective performance of Group Place is
not due to any local visual cue associated with the platform, reflected waves or other
non-spatial cues.
On trial 21, the escape latency increased for Group Place because, initially, the rat
searched near its former platform location (Fig. 4A, B). Relearning is generally very
rapid. Although moving the platform to the opposite quadrant is most easily
described as a 'reversal' procedure, this description is inexact. The reason is that the
allocentric spatial relationship of the distal room cues remains invariant when the
platform is moved. The rat needs only to learn the new position of the platform, not
that of the entire array of cues upon which effective escape performance is reliant
(cf. transposition procedures of Suzuki et al., 1980). Work by Sutherland et al.,
(1983) has shown that moving the platform to a new place is accompanied by a
disinhibition of exploratory rearing on the platform which habituates over the course
of initial training.

2. Path-directionality
Rats which have learned the position of the platform head towards it from a
distance (Fig. 5A). Thus the angle subtended between a tangent to the rat's
swim-path and a line drawn directly towards the platform declines as accuracy
improves. The angles recorded for different rats may be collated using circular
statistics (Gaile and Burt, 1980) and plotted on polar diagrams. Fig. 5B shows
 53

A. Rat path B. Polar diagrams

Hippocampal Corfical confrot

Fig. 5. A: accurate path taken by normal rat from its starting position at South to hidden platform at
NW. Tangent to path is drawn at position 0.5 m along path; angle x (0-360 o ) measures deviation of the
path from a line drawn direct to the centre of the platform. B: polar diagrams showing angles ( o ) for rats
given bilateral aspiration lesions of hippocampus (n = 10) or small control lesions of cortex (approx 1
mm2) overlying dorsal hippocampus (n = 13). The angles are plotted according to the convention x = 0 o
points East. The resultant vectors, R, are calculated using directional statistics. Only the angles of
approach of the cortical control animals show a significant deviation from uniformity, namely in the
direction of the platform.

p a t h - d i r e c t i o n a l i t y d a t a on trial 28 of training from a group of h i p p o c a m p a l a n d

cortical lesioned rats (from Morris et ai., 1982). The length of the vector d r a w n from
the centre of the p o l a r d i a g r a m s is a measure of the t e n d e n c y for a g r o u p of rats to
be h e a d i n g in the ' c o r r e c t ' direction.

3. Use of cueing procedure

I m p a i r m e n t s in p e r f o r m a n c e in the h i d d e n p l a t f o r m task after b r a i n lesions or
drugs m a y arise b e c a u s e these lesions affect the m o t i v a t i o n to escape, or sensorimo-
tor factors rather than spatial learning per se. The cueing procedure, in which the
escape p l a t f o r m p r o t r u d e s a b o v e the water surface provides a control for this. In one
e x p e r i m e n t ( M o r r i s a n d Schenk, 1983) rats with e n t o r h i n a l cortex lesions were given
28 trials of h i d d e n p l a t f o r m training, d u r i n g which a lesion deficit was observed,
followed by 20 trials of the cueing procedure. T h e y were then returned to the h i d d e n
p l a t f o r m c o n d i t i o n for 4 further trials. Fig. 6 shows the m e a n p e r f o r m a n c e over the
last 4 trials of the cueing p r o c e d u r e a n d the succeeding 4 trials. T h e r e are t w o - m a i n
findings in these results: (1) the cueing p r o c e d u r e a b o l i s h e d the lesion-induced
deficit in escape latency; but (2) the succeeding h i d d e n p l a t f o r m p r o c e d u r e caused
the latency deficit to reappear. Thus, it seems that the e n t o r h i n a l cortex lesion rats
are suffering a genuine spatial deficit, rather than an i m p a i r m e n t i n d u c e d by
m o t i v a t i o n a l or s e n s o r i m o t o r factors.

4. Transfer tests
T h e s p a t i a l l y d i s t r i b u t e d s w i m m i n g p a t t e r n on ' r e v e r s a l ' trials suggested an
a l t e r n a t i v e type of test; namely, removing the p l a t f o r m from the a p p a r a t u s entirely.
T h e rat is p l a c e d into the water for a limited p e r i o d (e.g. 60 s) a n d his ' s p a t i a l bias'
54

15 • E[
o SH+C

>,

I I

Cuen
ig Hidden
procedure p[Qfform

Fig. 6. Mean escape latencies of a group of rats given radiofrequency lesions of entorhinal cortex (n = 1 l)
and a combined group of sham lesion and unoperated control rats (n = 12). Note increased latency on
reintroduction of hidden platform procedure is restricted to EC group showing that their impairment is
not due to motivational or reinforcement aspects of the task.

60sec TrQnsfer test 7s1

so ~

- - C L

2£ hour t e s t

sot
0

Fig. 7. Paths taken during successive 60 s and 24 h transfer tests of a single typical rat given only limited
training, traced from the graphics printer output from the image analysis program. Right-hand columns
show time spent and annulus crossings for Adjacent-left quadrant (in this case NW), Training, Adjacent-
right, and Opposite quadrants.
 55

measured. One measure records the time spent searching in each of the 4 quadrants
of the pool. A second measure records the number of crossings of regions of the pool
marking the exact position and surface area of the former platform. The rationale
behind these measures is that rats trained to (say) NE should spend relatively more
time in NE than in the other 3 quadrants~ and that to the extent that their searching
for the now absent platform is accurate, they will cross a small circular area inside
the NE quadrant (where the platform used to be) more often than the corresponding
areas of the other 3 quadrants.
Originally, both measures were recorded by hand with the aid of electronic timers
and counters: lines were marked on the video-monitor to indicate the 4 quadrants
and 4 'annuli' marked to note the positions of the platform within each quadrant.
However, the tracking analysis program is now used to compute both measures
automatically. Fig. 7 shows an example from an experiment in which rats were given
only very limited training (13 trials) prior to transfer tests 60 s and 24 h after the
final trial of training. The results from one typical rat are presented. Note that he
spends over 50% of his time in the training quadrant during the first transfer test,
but only 36% during the second test. Furthermore, he crosses the former platform
position relatively more often during the first than the second test.

5. Effects of repeated transfer tests

Unpublished pilot experiments have shown that the absolute magnitude and rate
of decline of the 'spatial bias' revealed in transfer tests is dependent upon two main
factors: (1) the total number of training trials; and (2) the spacing of trials. In one
experiment in which only one trial was given per day (Heatlie, 1983) excellent
transfer test performance was obtained after only 10 training trials (60% time spent
in training quadrant). The decline in spatial bias with repeated testing may be due to
(1) true forgetting; or (2) extinction effects (the platform no longer being present). A
study was therefore conducted to compare these factors. Three groups of 7 rats were
trained for 20 trials and then given transfer tests as follows: Group 1 was given
transfer tests 4 h, 48 h and 336 h after training; Group 2 at 48 h and 336 h; and
Group 3 at 336 h only. To provide a single index of performance for each group at
each retention interval, the number of crossings of the training annulus position has
been plotted as a percentage of the total number of platform annulus crossings. This
'spatial bias' measure would be 25% in the event of no spatial bias. The results
plotted in Fig. 8 show (1) that at each interval, the group which was repeatedly
tested (Group 1) performs at least as well (if not better) than the groups only given
one or two transfer tests; and therefore (2) that the function relating spatial bias to
retention interval is relatively insensitive to extinction effects. More thorough studies
concerning the relationship between training parameters and retention are planned.

6. Spatial "matching to sample' procedure

A series of experiments are currently underway to compare spatial and working-
memory theories of hippocampal function. Full details will be reported elsewhere,
but the necessary prerequisite of these experiments was the development of 'match-
ing to sample' procedures in the pool. The simplest strategy is to run two trials per
56

07

0 Oroup 1

0"6 • 2

c
[] 3

0'5

0~-

L
el_

-- Chanca

0.2 I I
/~ ~8 336

Refenfion infervat - hours

Fig. 8. Retention functions based on p r o p o r t i o n of training annulus crossings relative to total crossings at
3 p o s t - t r a i n i n g intervals, These are pilot data based on n = 7 rats per group only.

day, per rat. On trial 1, the platform is placed in any of several positions in the pool
(e.g. SW, N W , N E or SE). The rat is allowed to swim to find the platform and then
left on it for 3 0 - 6 0 s. On trial 2, the platform is again hidden in the same position.
The question at issue is whether the rats can benefit on trial 2 from information
about the platform's position gained during trial 1. If so, the escape latency on trial
2, should be faster than on trial 1. The results of one experiment using this
procedure in the small pool (Morris, 1983) showed significantly faster escape on trial
2. However, before accepting this improvement as evidence of spatial ' m a t c h i n g to
sample', an alternative possibility must be considered. Trials 1 and 2 differ in other
respects, e.g. on trial 1, the rat is dry; on trial 2, he is wet. Thus, the improvement in
latency may be a non-specific or motivational difference. Accordingly, a further
group of rats was trained in which platform position on trial 2 was also r a n d o m with
respect to position, as on trial 1. The results for this group showed the following
pattern of results: on that subset of trials for which the platform was in a different
position on trial 2 than it had been on trial 1, escape latency on trial 2 was no
different than that shown during trial 1. However, for the 25% of trials for which the
platform was in the same position on trial 2 as it had been on trial 1 (required by the
r a n d o m schedule of four possible platform positions), escape latency on trial 2 was
faster than on trial 1. These findings imply that the animals are showing a 'win-stay'
tendency in the pool irrespective of training schedule. More extensive results are
presented in Morris (1983). A replication of this experiment using a ' t w o - p l a t f o r m '
procedure (see b e l o w - - s e c t i o n 7) is currently being tested to provide a measure of
choice accuracy of spatial ' m a t c h i n g to sample'.
 57

7. Spatial and non-spatial discrimination learning

The hidden platform procedures do not provide any index of spatial discrimina-
bility. A 'two-platform' procedure has therefore been developed (Fig. 2), which can
be used to study both spatial and non-spatial discrimination learning. Both plat-
forms are visible, 1 cm above the water. One platform is rigid; the other is the
floating platform as described previously. As the floating platform sinks beneath the
rat when he attempts to climb onto it, the animal soon learns to avoid it and to
approach the rigid one preferentially. Training can be in two forms: spatial dis-
crimination training, in which the rigid platform is defined by its position in space:
or t'.isual discrimination training, in which it is defined by its brightness or pattern
(e.g. grey positive; black and white stripes negative). In the latter case, heavy black
curtains suspended from rails attached to the laboratory ceiling are drawn around
the pool to exclude extra-maze cues. In one experiment in which these procedures
have been used, rats learned the spatial task to a criterion of 17/20 correct in
49 + 10 trials (inclusive of criterion); the visual discrimination task took 120 + 13
trials (n = 9). A 'correct' response was defined as approach to the rigid platform
without touching the floating platform. This procedure is presently being used to
examine the effects of hippocampal lesions (with modifications to examine reference
and working-memory, Olton et al., 1979) and the effects of anticholinergic drugs
(Hagan and Tweedie, in prep).
The surrounding black curtains can also be used to exclude extra-maze cues in
rats trained with the curtains open to localise a platform at a fixed location. As
shown in Fig. 9, pulling the curtains around the training quadrant section of the
pool caused a partial drop in performance; pulling them around the entire pool
caused performance to drop to chance. These findings imply that extra-maze cues
are essential for effective spatial localisation performance. However, they also imply
that the cues at or nearest the training quadrant are not the only cues of importance.

100

oa
t..a

50

r i i

Open 11~ Art

£ur'l'ains
Fig. 9. Percent correct rigid platform approaches in 10 trial test blocks with curtains open, drawn around
training quadrant only, or completely enclosing pool. See text for discussion of implications.
58

That performance drops to chance when all extra-maze cues are excluded implies, as
an incidental but not unimportant finding, that the rats are unable to distinguish the
floating and rigid platforms on the basis of local cues alone.

Discussion

The primary purpose of this paper has been to describe several developments of
the water-maze procedure. The automatic tracking provides a level of objectivity
usually only possible in fully automated experiments. The matching-to-sample
technique shows that contrary to the situation in foraging tasks, rats may show
'win-stay' strategies in escape behaviour irrespective of the schedule imposed upon
them. And the two-platform technique shows how non-spatial learning theory may
also be studied using the same general paradigm.
One conceptual and several practical aspects of the task deserve further comment.
The use of the hidden platform requires that the rat approach a position in space
which it can neither hear, see or smell. Extra-maze cues must be used for accurate
localization. Thus the absolute levels of performance reflect the 'acuity' of spatial
memory and its disruption by drugs or lesions, In T-mazes, by comparison, rats
make only a binary decision between going left or right, to one part of the apparatus
or to another. Once that decision is made, the design of the maze arms leads the
animal to reward; no further demands are placed upon the spatial localisation
system. In the water-maze, on the other hand, accurate directionality requires
continual monitoring of the animal's position in relation to extra-maze cues. This
may involve 'cognitive mapping', as proposed by O'Keefe and Nadel (1978): or
some more parsimonious 'snapshot' procedure for homing in on the correct location
(Cartwright and Collett, 1982).
From a practical point of view, the general procedures may prove useful for
several different kinds of neurobiological study. Already, the technique has been
exploited in studies of cortical function (Kolb et al., 1983a, b) sex differences and
hemispheric asymmetry (Therrien et al., 1982; Dennenberg, pets. comm.) superior
collicular lesions (Milner and Lines, 1983), nutritional deficiency (Heatlie, 1983) and
in studies of catecholamine depletion (Hagan et al., 1983). The latter study revealed
intact spatial learning in animals that were ataxic after striatal dopamine depletion,
the water serving to 'activate' the animals. Studies of 'latent learning' (Sutherland
and Linggard, 1982) and 'overshadowing' and 'blocking' of spatial and non-spatial
learning (Morris and Heatlie, in prep.) are also underway, using controlled cues
inside the curtaining, as well as pharmacological studies, some using osmotic
minipumps. Gage (pets. comm.) is currently using the procedure to investigate
recovery of function after septo-hippocampal transplants.
The main virtue of the procedure is the speed of training. Few of the experiments
conducted in St. Andrews have lasted more than two weeks, although there is no
reason why more extensive training could not be examined. The disadvantages of the
technique are (1) an inability to vary motivational level or reinforcement magnitude:
(2) that experiments must be run by hand, which is more tedious than using fully
 59

automated equipment; (3) that immersion into water may cause endocrinological or
other stress effects which, while not of primary interest, may nevertheless interact
with lesion or pharmacological manipulations in uncontrolled ways.

Equipmenl

The image analysing equipment may be obtained from HVS IMAGE ANALYS-
ING, 22 Cromwell Road, Kingston KT2 6RE, UK. The BBC Model B Computer is
widely available, but we have found that T W I L L S T A R C O M P U T E R S LTD., 17
Regina Road, Southall, Middlesex, UB2 5BL, UK. can usually supply within 48
hours. Fibreglass and resin are available from yacht chandlers.

Acknowledgements

I am grateful to Christopher Barman and Jim Hagan who helped to build both
water mazes: to Paul Heatlie for helping with installation of the tracking system, to
Joram Feldon and Nick Rawlins for first suggesting the floating platform procedure,
and to Robin Fowler and Peter Frew for technical advice. The initial work was
supported by MRC Grant G 9 7 9 / 2 6 5 / N . The new pool was built with the aid of
SERC Grant GR/C/39071. Further enquiries should be directed in the first
instance to Richard Morris at the address above.

References

Cartwright, B.A. and Collett, T.S. (1982) How honey bees use landmarks to guide their return to a food
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