Behavioural Processes, Elsevier
12 (1986) 287-297
28-l
SOUE FACTORS AFFECTING PIGEONS’ VISUAL TRACKING BEHAVIOR DONALDK. UILKIE Department of Psychology, Vancouver Canada VET lY7 (Accepted
22 August
The University
of
British
Columbia,
1985)
ABSTRACT Wilkie, D.R., 1985. Some factors affecting behavior. Behav. Processes 12: 287-297.
pigeons’
visual
tracking
A novel delayed conditional discrimination procedure was used successfully to investigate some factors affecting pigeons’ tracking Trials began with the brief illumination of one of visual stimuli. of the keys (randomly selected) from the leftor right-hand column of a three by three matrix of pecking keys. Then the next key in Finally, the remaining key in the row the same row was briefly lit. was lit along with one (randomly chosen) of the remaining five keys A peck to the former but not from the leftor right-hand columns. the latter key was designated as correct and was rewarded. In Experiment 1 pigeons made correct choices on between 70 and 80 percent of trials, thereby demonstrating an ability to visually track objects. In Experiment 2 tracking accuracy was: a) reduced when either the first or second key in the sequence was omitted, b) improved when the sequence was repeated three times, c) reduced when duration of key illumination was reduced, and d) reduced by delays imposed between keys early but not late in the sequence. It was also found that tracking of vertical and horizontal sequences was approximately equal.
INTRODUCTION Most
investigators
laboratory of
the
have used
relatively
stimuli,
Berkeley,
Warmath,
Hodos,
Smith,
1983;
Mello,
1968;
1970,
studies), consistent latter
type
Morgan’s (1925) only
with
gerbils film
that
study
study
of
to
0376-6357/86/$03.50
behavior
for
with (1979.
a review
early
that
of
a moving
for ball
require
3)
cue.
seed. of
the
this
example,
basket
successfully
of
and Kijhler’s
subjects
sunflower pursuit
earlier
importance
[see,
a moving
Experiment
movement during
of
a moving
that
& Bloch, 1980;
in some manner
a thrown
intercepting exist
the
keys
6 Carpenter,
6 Merigan,
movements
all
dynamic
to
Rivard.
1936,
a dog catching
and consume
detailed
Carpenter
Martinoya,
Although
in nearly used
respond
their
was recognized of
have
to
Pasternak
cue.
experiments
and Dizio
pursue
records,
the
a chimpanzee of
their
Cheal,
of
1978;
1978;
Kennedy,
that
simply
1976;
coordinate
behavior
a handful
coordinate Ingle,
of
see
than
(1930)
6 Tunkl,
psychological
Furthermore,
studies
required
Rulvanny,
in the
stimuli.
6. Bonbright,
1971;
rather
behavior
number of
have been
1958;
Siegel,
animal
stationary
small
subjects
(e.g.,
of
of
fruit.1
to
For example, trained By using target
was
0 1986 Elsevier Science Publishers B.V. (Biomedical Division)
288
measured. gerbils pursue lark
One of were
to
a collision (1975)
involve
that
trajectory,
pecking
actually
of
other
keys
which
the white
conditions,
pigeons
key and pecked
the
every
target
pecking
the white of
field
the
trajectories,
a paradigm
of
variables
strategies
Pisacreta’s
procedure
the
research
present
its
discrimination by three
left-
lit,
implemented
observed
of
two keys one of Choice
trial, not
Because to the
first
experiment
was concerned
learn
task.
Experiment
such
the “sit
with
five
the
the
study the
to
predict
of
the
subject
to
purpose
of
in the
a paradigm
box
and
tracking
behavior.
containing
with
brief.
The third
key
former
sequence or left
in this question
a three
the
remaining
and wait”
the
and
conditional
start
of
right
ability
of
in a row. the
be effective
could
this
Trials
field to
key.
occurred
delayed
the white
keys
allowing
in a Skinner
keys.
rewarded.
will
the
with
may be either
on a particular
sometimes
of
columns.
and is
illuminations
investigation
key
and responded
was therefore
to develop
pecking
along
right-hand
as correct
rows
of
without
for
the
of
systematic
as sometimes
to avoid
the white
animals’
attempt
a variant
illumination
then or
is
task,
matrix
successive row is
to
the
position
on that
It
key of
in the
keys
in the
is
designated
key
and in any of strategy
paradigm.
The
of
pigeons
whether
1
Subjects King pigeons
(designated
three
Pisacreta
WETHOD
Four Silver
a
Under these
a few preferred
the
seems needed.
utility
Our procedure
such
and to
Then,
3 s.
tracking
of
not
predict
pigeons
field
“X”.
the
appeared
did
to
key displaying
one of
allows
it
trained
every
stopped
on such behavior
use non-tracking
demonstrate
a white
importance that
and
subjects
a white
tracked
when it
obvious
and
and Lancaster
do when tracking
He first
when the
toward
the
target
although
the
sometimes
was changed
pigeons
by orienting
Because
effect
the
which,
displayed
successfully
.5 s,
press)
required
motion.
However,
was that
moving
work.
displayed
field
it.
(in
animals
keys
findings a quickly
a study
moved,
that
eight
displaying
changed
related
or apparent
whichever
of
Goodale
conducted
something in real
interesting ahead
course.
(1.982)
cues
objects
most
orient
have reported
Pisacreta
peck
their
able
as HO, JA,
SA, and KI)
289
served.
Each subject
experience about
with
90% of
and after
had previous
static
stimuli.
free-feeding
test
delayed
matching
The pigeons
body weight
to
sample
were maintained
by mixed
grain
at
obtained
during
sessions.
AJnaratus The subjects contained pecking
two the
were mounted keys
of
emitting
the
chambers
and a three
chambers
5 cm apart,
(2 cm in diameter)
15 g t.15
diode
(approximately
Data collection
the
1.7
(Gilbert
6 Rice.
which matrix
keys
center; 3.5
of
(3.25
in the
cm apart.
sensed
Behind
key pecks
and a FLV117 red
cm in
other
having
a
light
cm/m2).
and experimental
NOVA 3 computer
to
which
greater,
of
by three
were mounted
N) or
each
Plexiglas
center
a microswitch,
about
Data General language
in four
feeder
In two of
key was mounted
force
tested
a BRS/LVE grain keys.
diameter)
each
were
control
operating
were
under
carried
out
by a
RDOS and the MANE
1979).
Procedure Because lit
keys
each
for
of
grain
start
were
of
of
replacement)
and lit
and the
one
(S2)
also
for
next
immediately. was lit, right of
along
with
columns.
them:
sequence rewarded key led
the
in that .5 s. one of
remained
key that
access
the
the
remaining lit
until
occupied
the
was designated
to mixed
grain.
intertrial
or
(without turned
off
on
key
in the
row (S3)
keys
in the
left
the
pigeon
end point
A peck
one
the
and was
at the
which
or
pecked of
as correct
interval,
session,
left-
was then
last
peck
At the
row was turned the
five
the
selected
(Sl)
particular Finally,
the
keys
illuminations
5-s
in either
key
each
interval.
This
.5 s.
of
to
keys
to
was necessary.
During
for
These
with
six
previously
training
60 sessions.
was randomly
at the
only
of
the matrix
A peck key
trained
by a 10 s intertrial
one of
column
had been
no preliminary
consisted
separated
a trial,
right-hand
subjects
reward,
The experiment 60 trials
the
distractor
was extended
to
15 s.
RESULTS AND DISCUSSION Tracking for
all
subjects
percent. final
accuracy
Figure 20 sessions
These
results
increased
and reached 1 shows of
the
clearly
over
the
asymptotic
choice
course levels
accuracy
during
of of
the
experiment
about
the
first
70 to
80
10 and
experiment. demonstrate
that
in the
present
paradigm
pigeons not
are
appear
able to
to
Fig. 1. Percent of key during the first
Experiment
b)
presentation
imposing
the
a “retention
sequences
also
Tracking
effects
of
accuracy of
choice
d) variation
cues.
the
tracking repetition of
and S2,
choice
stimulus,
although
the
task
does
easy.
trials on which the subjects 10 and last 20 sessions of
experiment
influence
Sl or S2’
Sl
a visual
chose the Experiment
correct 1.
2
In this might
track
be especially
of
the cues, the
interval” accuracy
several
manipulations
were measured:
Sl,
S2, c)
delays
variation between
between for
a)
SS sequence
tbat
omission
of
the
Sl
and S2,
duration
of
e)
S2 and presentation
vertical
of
before
of
the
and horizontal
was compared.
l¶ETHOD Subjects
and Aonaratus
The four
pigeons
from Experiment
1 served.
(CA, QU. PP.. PR, HE, HA, CP, and PO) with
similar
Eight
other
backgrounds
birds as the
291
other
pigeons
that
used
also
participated.
in Experiment
The apparatus
was identical
to
1.
Procedure Most order
subjects
in which
manner.
Sessions
intertrial
determine
consisted
of
if
procedure
procedures.
varied
60 trials,
These
shortening
the
effect
The
in a quasi-random
separated
During
s period
elapsed
procedures
(complete
there
sequence)
sequence
were
by 10 s
from
implemented
three
to
accuracy.
other
procedure
S3 and the were
two alternations
and test
procedures
by S5 and the
were
each
and a .5
presented.
between with
would
first
Sl was presented
distractor
to
two keys
In the
S2 was followed
(i.e.,
the
before
procedures
on tracking
Sl was omitted
distractor).
three
different
administered
of
SL*_.
a deleterious
both
in several
were
intervals.
Omission
have
participated procedures
In
baseline phase
lasting
sessions. Repetition
determine
if
facilitate were
of
sequence.
repetition
tracking
of
repeated also
of
During three
of
sessions
on the
Sl,
to
illuminations
preceding
blocks
the
third
was implemented
key
As in the
three-session
test
times;
procedure
a sequence
accuracy.
two alternations
conditions.
This
S2.
repetition
of
would
procedures baseline
and test
SS sequence the
there
was
distractor
was
presented. duration
Kev
determine
if
accuracy.
the
varied.
This
duration
of
In each
equally
probable.
as usual.
of
eight
sessions
On one,
During
the
procedure
was implemented
key illuminations three
types
Sl and S2 were
other
two,
these
to
affected of
trials
presented
stimuli
tracking
for
lasted
were 500 ms,
250 and 2000
ms. Interval implemented affected
between to
tracking
equal
probability
trial
(short-short)
S2 and SS.
key
within
each
another
the
s then
2 s and 2 s then
of
of
varied.
simultaneous
if
a retention
.5 s,
Delays onset interval
varied. between
types
trials
28 sessions.
the
procedure
was
illuminations occurred During both
delays
and long-short)
with
one type
of
Sl and S2 and were
the
2 8.
delays
were
respectively.
were of
of
This
key
.5 s separated
(long-long)
two (short-long
and the such
delays Four
a delay
During other
if
accuracy.
During
Delav
illuminations
determine
imposed
between
S3 and the affected
the
offset
distractor
accurate
choice.
of
S2
to determine The delays
.5
292
occurred
with
were
2,
.5,
equal
probability
and 4 s.
There
Horizontal sequences
vs vertical
trials
sessions.
Horizontal 1.
trials
that
equal
sequences
Vertical
except
with
were
trials
S1 was selected
12 sessions.
between
and vertical in each
identical
were
from
the
of
either
The delays
S1 and S2.
probability
trials
sequence
of
Horizontal
sequences.
occurred
Experiment
in each
was no delay
to
identical top
of
33
those
to
in
horizontal
or bottom
row of
keys.
RESULTS Omission key
in the
of
S1ps2_.
sequence
had a deleterious
sequence
length
stimulus
had a larger
suggesting important
that to
omission
is
important
contiguity
effective
to
effect,
tracking.
than
effect, of
key
Renetition
incorrect
permitting keys,
of
first
or
second
demonstrating Omission
omitting
the
illuminations
of
first
that
the
second
(Figure
in space
2)
and time
tracking.
Percent of trials on which Fig. 2. key when Sl or Sp was omitted.
before
the
the
subjects
sequence.
Repeating
the
the
to
between
pigeon
facilitated
choose
tracking
(Table
chose
sequence
1).
the
the
correct
more than correct
and
once
is
293
TABLE 1 Results
Subject
of
some of
the manipulations
Number of CA JA KI S,
Subject
JA SA KI
Subject
BA CP FO
to
in Experiment
3 98 85 72
1 a7 70 65
Repetitions
S, Delay
2.
.5 8
2 s
4 s
72 77 79
74 79 a5
71 a2 85
Horizontal
Seouence
Vertical
64 65 69
Percent of trials on which the subjects Fig. 3. key when the duration of Sl and S2 was varied.
Seouence 60 58 71
chose
the
correct
294
Kev duration tracking
accuracy
illuminated was best
for
delays
factorially
short
than
the Delav
delays tracking
of
kev
a long
same delay varied. up to accuracy
Tracking
lit
the
S
When there 4 s between
was poorest
longest
(2
both
4).
when keys
were
tracking
s). Tracking
varied.
1 (Figure
affected
(HE and JA)
and S2 and between Tracking
S
accuracy
and S
2 3 was best when a
Sl and S2 and S2 and S3.
delays
between
imposed
(Table
key illumination
For two subjects
when these delay
of
illuminations
between
separated
was poorest
Furthermore,
were
manipulated
C.5 6) delay
Tracking
3).
250 ms.
keys
between
as the
Duration
(Figure only
when the
Interval varied were
varied.
S
between
1
were
long
(2 8).
and S2 was more disruptive S2 and S3.
were no delays
between
S2 and S3 had little
effect
ST and S2, on
1).
Fig. 4. Percent of trials on which the subjects chose the correct short (S - .5 s) key when the delay between Sl and S2 was either or long CL - 2 8) or a combination of short and long (S then L, L then S).
295
Horizontal horizontal (Table
vs vertical
sequences.
and vertical
sequence
Tracking
trials
accuracy
on
was approximately
equal
1).
DISCUSSION The major develop
purpose
a paradigm
tracking
of
procedure
visual
that
goal:
the
by manipulation
of
non-tracking
strategies
Pisecrete,
in this
(1979).
questions
such
sometimes
with
In such
as the
etc.
used
It
version
is
depending
pursuer
avoided,
or merely for
strategies
cited
in Ingle
Fleck,
1975)
overtake” pursuer
that
observed,
may also et
moving
al.,
to
al.,
1979).
may be used
strike
orient This
used
other
procedures
the
the
strategy the
prey.
a target
strategy “fixate target
the
moving
might
while
be celled
and overtake” moves
faster
being
species. and
1977,
(Lancaster e “fixate
strategy
be rate
pursued,
found
(Ingle,
upon the
6 and
ability
of
the
is
ineffective
end toads
at 3O’/s.
when pursuing target
might end the
been
fish
permit
moving
in different
has found--frogs
by gerbils of
is
have
This
also
object
be called
relies
used
sequences,
strategies
jacket
This
type
paradigm
track
and toads
and leather
as Ingle
the
the
will
object
strategies Frogs
pecking
be studied.
tracked
whether
be even
of
of
how animals
use what might
than
will
of
paradigm
tracking
of
exist.
1979)
ahead
the
could
and be different
behind
when the
ineffective--when
lit
effected
use
vs non-linear
prey
targets
The strategy is
is
stimuli.
to
matrix
version
present
different
to move faster
seed
the
procedure
linear
different
velocity
three
pursuing
consistently
of
arrangement
about
can move,
strategy.
fast
tracking
a much larger
accuracy the
questions
likely
the
Some evidence
of
of
upon the
et which
other
effects
at answering
objects.
for
successively
do with
an elaborated
on tracking
An elaborated attempts
potential
track of
discrimination
when implemeted
size,
discrimination
paradigm
or on e CRT-photocell-on-the-beak
target
to
es duration
unable
as they
have
Accuracy
such
are
conditional
by Blough
for
key.
to able
to
accurate
1982).
The delayed
keys
are
attempt
affecting
conditional
appears
lit
was to
factors
Pigeons
variables
subjects
research
The delayed
lest
Furthermore,
more useful
present
stimuli.
this
end choose
(cf.
the
investigating
was implemented
accomplishing keys
of
for
a moving
moving “predictive strategy than
the
sunflower
forward
(Ingle
tracking”
et end
is pursuer
or when
296
the
distance
between
Human infants catching
rapidly
move their to
evidence moving
it.
the
that
we are
light.
vision, that
the
the
pigeon of
keys
Hofsten.
with
the
visual
feedback
unsure
of
rewarded cues
influencing
great. of
object (1985)
is
strategy
1983).
when
They appear
and toward
it
to
so as
have described
important
when aiming
at
by which
animals
are have
accuracy are
able
pigeon
only
track
the
has a panoramic
a few degrees
proximity
to
the
of
location
field
visual
display
of arc
and
made
difficult.
a delayed choices
how our pigeons
the
occupied
was in close
seems to the
that
any strategy
In conclusion,
visual
type
(van
and Fleury
Hay,
The fact
that
observation
which
a different
objects
is
targets.
At present of
use
and pursuer
simultaneously Bard,
showing
target
seem to moving
hands
intercept
the
conditional contingent
potential of to
tracking perform
discrimination upon tracking
for
the
and for such
study
procedure a sequence
of
in
of
factors
elucidating
the means
behavior.
ACKNOWLEDGMENTS This research was supported by a grant from the Natural Sciences and D. Kennedy, and Engineering Research Council of Canada. R. Takai. K. Arbour also provided K. Arbour assisted with testing the birds. Send reprint requests to Donald helpful comments on the manuscript. I4. Wilkie, Psychology Department, The University of British Columbia, Vancouver, B.C. Canada V6T lY7.
REFER8NCES Bard,
C., Hay. L., 6 Fleury, M. (1985). Role of peripheral vision in the directional control of rapid aiming movements. Canadian Journal of PsvcholoKy, 39, 151-161. Movement Berkeley, hl. A., Warmath, D. S., 6 Tunkl, J. E. (1978). Journal of Comparative discrimination capacities in the cat. and Physioloaical Psychology, 92, 463-473. Effects of number and form of stimuli on Blough, D. S. (1979). Journal of Experimental visual search in the pigeon. PsycholoKy: Animal Behavior Processes, 2, 211-223. (1958). Carpenter, B., 6 Carpenter, J. T. The perception of movement by young chimpanzees and human children. Journal of Comparative and PhvsioloKical PsvcholoKy, 5l, 782-784. Gilbert, S. G., & Rice, D. C. (1979). NOVA SKED II: A behavioral notation language utilizing the Data General Corporation real-time disk-operating system. Behavior Research Rethods 6 Instrumentation. l_&, 71-73. Neural mechanisms of visual orientation Goodale. ?I. A. (in press). In A. Hein 6 t4. Jeannerod in rodents: Targets versus places. (Eds.), Spatially oriented behavior. Berlin: Springer-Verlag. Detection of the Hodos, W., Smith, L., Bonbright, J. C. (1976). Journal of velocity of movement of visual stimuli by pigeons. the Experimental Analysis of Behavior, 25, 143-156.
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Ingle, D.. Cheal, M. D., 6 Dizio, P. (1979). Cine analysis of visual orientation and pursuit by the Mongolian gerbil. Journal of Comparative and Physiolonical Psychology, 93, 919-928. The nature and physiological basis of Kennedy, J. L. (1936). Psycholoxical Review, 43, movement discrimination in animals. 494-521. London: Routledge and KShler, W. (1925). The mentalitv of apes. Kegan Paul. Pursuit and prediction in Lancaster, B. S.. & Mark. R. F. (1975). the tracking of moving food by a teleost fish (Acanthaluteres spilomelanurus). Journal of Experimental BioloKy, 63, 627-645. (1983). Comparing frontal Bartinoya, C., Rivaud, S., & Bloch, S. and lateral viewing in the pigeon. II. Velocity thresholds for movement discrimination. Behavioural Brain Research, 8, 365-385. (1968). Interhemispheric transfer of a discrimination Hello, N. K. of moving patterns in pigeons. Brain Research, 1, 390-398. Borgan. C. L. (1930). The animal mind. New York: Longmans. Vision Bulvanny, P. (1978). Velocity discrimination by pigeons. Research, l8, 531-536. (1980). Movement detection by cats: Pasternak. T., & Merigan, W. H. Journal of Invariance with direction and target configuration. Comparative and PhvsioloKical Psycholoxv, 94, 943-952. Stimulus control of the pigeon’s ability to Pisacreta, R. (1982). peck a moving target. Journal of the Experimental Analvsis of Behavior, 37, 301-309. Siegel, R. K. (1970). Apparent movement detection in the pigeon. Journal of the Experimental Analvsis of Behavior, l4. 93-97. Siegel, R. K. (1971). Apparent movement and real movement detection in the pigeon: Stimulus generalization. Journal of the Experimental Analvsis of Behavior, l6, 189-192. von Hofsten, C. (1983). Catching skills in infancy. Journal of Experimental Psvcholoxy: Human Perception and Performance, 9, 75-85.
12 (1986) 287-297
28-l
SOUE FACTORS AFFECTING PIGEONS’ VISUAL TRACKING BEHAVIOR DONALDK. UILKIE Department of Psychology, Vancouver Canada VET lY7 (Accepted
22 August
The University
of
British
Columbia,
1985)
ABSTRACT Wilkie, D.R., 1985. Some factors affecting behavior. Behav. Processes 12: 287-297.
pigeons’
visual
tracking
A novel delayed conditional discrimination procedure was used successfully to investigate some factors affecting pigeons’ tracking Trials began with the brief illumination of one of visual stimuli. of the keys (randomly selected) from the leftor right-hand column of a three by three matrix of pecking keys. Then the next key in Finally, the remaining key in the row the same row was briefly lit. was lit along with one (randomly chosen) of the remaining five keys A peck to the former but not from the leftor right-hand columns. the latter key was designated as correct and was rewarded. In Experiment 1 pigeons made correct choices on between 70 and 80 percent of trials, thereby demonstrating an ability to visually track objects. In Experiment 2 tracking accuracy was: a) reduced when either the first or second key in the sequence was omitted, b) improved when the sequence was repeated three times, c) reduced when duration of key illumination was reduced, and d) reduced by delays imposed between keys early but not late in the sequence. It was also found that tracking of vertical and horizontal sequences was approximately equal.
INTRODUCTION Most
investigators
laboratory of
the
have used
relatively
stimuli,
Berkeley,
Warmath,
Hodos,
Smith,
1983;
Mello,
1968;
1970,
studies), consistent latter
type
Morgan’s (1925) only
with
gerbils film
that
study
study
of
to
0376-6357/86/$03.50
behavior
for
with (1979.
a review
early
that
of
a moving
for ball
require
3)
cue.
seed. of
the
this
example,
basket
successfully
of
and Kijhler’s
subjects
sunflower pursuit
earlier
importance
[see,
a moving
Experiment
movement during
of
a moving
that
& Bloch, 1980;
in some manner
a thrown
intercepting exist
the
keys
6 Carpenter,
6 Merigan,
movements
all
dynamic
to
Rivard.
1936,
a dog catching
and consume
detailed
Carpenter
Martinoya,
Although
in nearly used
respond
their
was recognized of
have
to
Pasternak
cue.
experiments
and Dizio
pursue
records,
the
a chimpanzee of
their
Cheal,
of
1978;
1978;
Kennedy,
that
simply
1976;
coordinate
behavior
a handful
coordinate Ingle,
of
see
than
(1930)
6 Tunkl,
psychological
Furthermore,
studies
required
Rulvanny,
in the
stimuli.
6. Bonbright,
1971;
rather
behavior
number of
have been
1958;
Siegel,
animal
stationary
small
subjects
(e.g.,
of
of
fruit.1
to
For example, trained By using target
was
0 1986 Elsevier Science Publishers B.V. (Biomedical Division)
288
measured. gerbils pursue lark
One of were
to
a collision (1975)
involve
that
trajectory,
pecking
actually
of
other
keys
which
the white
conditions,
pigeons
key and pecked
the
every
target
pecking
the white of
field
the
trajectories,
a paradigm
of
variables
strategies
Pisacreta’s
procedure
the
research
present
its
discrimination by three
left-
lit,
implemented
observed
of
two keys one of Choice
trial, not
Because to the
first
experiment
was concerned
learn
task.
Experiment
such
the “sit
with
five
the
the
study the
to
predict
of
the
subject
to
purpose
of
in the
a paradigm
box
and
tracking
behavior.
containing
with
brief.
The third
key
former
sequence or left
in this question
a three
the
remaining
and wait”
the
and
conditional
start
of
right
ability
of
in a row. the
be effective
could
this
Trials
field to
key.
occurred
delayed
the white
keys
allowing
in a Skinner
keys.
rewarded.
will
the
with
may be either
on a particular
sometimes
of
columns.
and is
illuminations
investigation
key
and responded
was therefore
to develop
pecking
along
right-hand
as correct
rows
of
without
for
the
of
systematic
as sometimes
to avoid
the white
animals’
attempt
a variant
illumination
then or
is
task,
matrix
successive row is
to
the
position
on that
It
key of
in the
keys
in the
is
designated
key
and in any of strategy
paradigm.
The
of
pigeons
whether
1
Subjects King pigeons
(designated
three
Pisacreta
WETHOD
Four Silver
a
Under these
a few preferred
the
seems needed.
utility
Our procedure
such
and to
Then,
3 s.
tracking
of
not
predict
pigeons
field
“X”.
the
appeared
did
to
key displaying
one of
allows
it
trained
every
stopped
on such behavior
use non-tracking
demonstrate
a white
importance that
and
subjects
a white
tracked
when it
obvious
and
and Lancaster
do when tracking
He first
when the
toward
the
target
although
the
sometimes
was changed
pigeons
by orienting
Because
effect
the
which,
displayed
successfully
.5 s,
press)
required
motion.
However,
was that
moving
work.
displayed
field
it.
(in
animals
keys
findings a quickly
a study
moved,
that
eight
displaying
changed
related
or apparent
whichever
of
Goodale
conducted
something in real
interesting ahead
course.
(1.982)
cues
objects
most
orient
have reported
Pisacreta
peck
their
able
as HO, JA,
SA, and KI)
289
served.
Each subject
experience about
with
90% of
and after
had previous
static
stimuli.
free-feeding
test
delayed
matching
The pigeons
body weight
to
sample
were maintained
by mixed
grain
at
obtained
during
sessions.
AJnaratus The subjects contained pecking
two the
were mounted keys
of
emitting
the
chambers
and a three
chambers
5 cm apart,
(2 cm in diameter)
15 g t.15
diode
(approximately
Data collection
the
1.7
(Gilbert
6 Rice.
which matrix
keys
center; 3.5
of
(3.25
in the
cm apart.
sensed
Behind
key pecks
and a FLV117 red
cm in
other
having
a
light
cm/m2).
and experimental
NOVA 3 computer
to
which
greater,
of
by three
were mounted
N) or
each
Plexiglas
center
a microswitch,
about
Data General language
in four
feeder
In two of
key was mounted
force
tested
a BRS/LVE grain keys.
diameter)
each
were
control
operating
were
under
carried
out
by a
RDOS and the MANE
1979).
Procedure Because lit
keys
each
for
of
grain
start
were
of
of
replacement)
and lit
and the
one
(S2)
also
for
next
immediately. was lit, right of
along
with
columns.
them:
sequence rewarded key led
the
in that .5 s. one of
remained
key that
access
the
the
remaining lit
until
occupied
the
was designated
to mixed
grain.
intertrial
or
(without turned
off
on
key
in the
row (S3)
keys
in the
left
the
pigeon
end point
A peck
one
the
and was
at the
which
or
pecked of
as correct
interval,
session,
left-
was then
last
peck
At the
row was turned the
five
the
selected
(Sl)
particular Finally,
the
keys
illuminations
5-s
in either
key
each
interval.
This
.5 s.
of
to
keys
to
was necessary.
During
for
These
with
six
previously
training
60 sessions.
was randomly
at the
only
of
the matrix
A peck key
trained
by a 10 s intertrial
one of
column
had been
no preliminary
consisted
separated
a trial,
right-hand
subjects
reward,
The experiment 60 trials
the
distractor
was extended
to
15 s.
RESULTS AND DISCUSSION Tracking for
all
subjects
percent. final
accuracy
Figure 20 sessions
These
results
increased
and reached 1 shows of
the
clearly
over
the
asymptotic
choice
course levels
accuracy
during
of of
the
experiment
about
the
first
70 to
80
10 and
experiment. demonstrate
that
in the
present
paradigm
pigeons not
are
appear
able to
to
Fig. 1. Percent of key during the first
Experiment
b)
presentation
imposing
the
a “retention
sequences
also
Tracking
effects
of
accuracy of
choice
d) variation
cues.
the
tracking repetition of
and S2,
choice
stimulus,
although
the
task
does
easy.
trials on which the subjects 10 and last 20 sessions of
experiment
influence
Sl or S2’
Sl
a visual
chose the Experiment
correct 1.
2
In this might
track
be especially
of
the cues, the
interval” accuracy
several
manipulations
were measured:
Sl,
S2, c)
delays
variation between
between for
a)
SS sequence
tbat
omission
of
the
Sl
and S2,
duration
of
e)
S2 and presentation
vertical
of
before
of
the
and horizontal
was compared.
l¶ETHOD Subjects
and Aonaratus
The four
pigeons
from Experiment
1 served.
(CA, QU. PP.. PR, HE, HA, CP, and PO) with
similar
Eight
other
backgrounds
birds as the
291
other
pigeons
that
used
also
participated.
in Experiment
The apparatus
was identical
to
1.
Procedure Most order
subjects
in which
manner.
Sessions
intertrial
determine
consisted
of
if
procedure
procedures.
varied
60 trials,
These
shortening
the
effect
The
in a quasi-random
separated
During
s period
elapsed
procedures
(complete
there
sequence)
sequence
were
by 10 s
from
implemented
three
to
accuracy.
other
procedure
S3 and the were
two alternations
and test
procedures
by S5 and the
were
each
and a .5
presented.
between with
would
first
Sl was presented
distractor
to
two keys
In the
S2 was followed
(i.e.,
the
before
procedures
on tracking
Sl was omitted
distractor).
three
different
administered
of
SL*_.
a deleterious
both
in several
were
intervals.
Omission
have
participated procedures
In
baseline phase
lasting
sessions. Repetition
determine
if
facilitate were
of
sequence.
repetition
tracking
of
repeated also
of
During three
of
sessions
on the
Sl,
to
illuminations
preceding
blocks
the
third
was implemented
key
As in the
three-session
test
times;
procedure
a sequence
accuracy.
two alternations
conditions.
This
S2.
repetition
of
would
procedures baseline
and test
SS sequence the
there
was
distractor
was
presented. duration
Kev
determine
if
accuracy.
the
varied.
This
duration
of
In each
equally
probable.
as usual.
of
eight
sessions
On one,
During
the
procedure
was implemented
key illuminations three
types
Sl and S2 were
other
two,
these
to
affected of
trials
presented
stimuli
tracking
for
lasted
were 500 ms,
250 and 2000
ms. Interval implemented affected
between to
tracking
equal
probability
trial
(short-short)
S2 and SS.
key
within
each
another
the
s then
2 s and 2 s then
of
of
varied.
simultaneous
if
a retention
.5 s,
Delays onset interval
varied. between
types
trials
28 sessions.
the
procedure
was
illuminations occurred During both
delays
and long-short)
with
one type
of
Sl and S2 and were
the
2 8.
delays
were
respectively.
were of
of
This
key
.5 s separated
(long-long)
two (short-long
and the such
delays Four
a delay
During other
if
accuracy.
During
Delav
illuminations
determine
imposed
between
S3 and the affected
the
offset
distractor
accurate
choice.
of
S2
to determine The delays
.5
292
occurred
with
were
2,
.5,
equal
probability
and 4 s.
There
Horizontal sequences
vs vertical
trials
sessions.
Horizontal 1.
trials
that
equal
sequences
Vertical
except
with
were
trials
S1 was selected
12 sessions.
between
and vertical in each
identical
were
from
the
of
either
The delays
S1 and S2.
probability
trials
sequence
of
Horizontal
sequences.
occurred
Experiment
in each
was no delay
to
identical top
of
33
those
to
in
horizontal
or bottom
row of
keys.
RESULTS Omission key
in the
of
S1ps2_.
sequence
had a deleterious
sequence
length
stimulus
had a larger
suggesting important
that to
omission
is
important
contiguity
effective
to
effect,
tracking.
than
effect, of
key
Renetition
incorrect
permitting keys,
of
first
or
second
demonstrating Omission
omitting
the
illuminations
of
first
that
the
second
(Figure
in space
2)
and time
tracking.
Percent of trials on which Fig. 2. key when Sl or Sp was omitted.
before
the
the
subjects
sequence.
Repeating
the
the
to
between
pigeon
facilitated
choose
tracking
(Table
chose
sequence
1).
the
the
correct
more than correct
and
once
is
293
TABLE 1 Results
Subject
of
some of
the manipulations
Number of CA JA KI S,
Subject
JA SA KI
Subject
BA CP FO
to
in Experiment
3 98 85 72
1 a7 70 65
Repetitions
S, Delay
2.
.5 8
2 s
4 s
72 77 79
74 79 a5
71 a2 85
Horizontal
Seouence
Vertical
64 65 69
Percent of trials on which the subjects Fig. 3. key when the duration of Sl and S2 was varied.
Seouence 60 58 71
chose
the
correct
294
Kev duration tracking
accuracy
illuminated was best
for
delays
factorially
short
than
the Delav
delays tracking
of
kev
a long
same delay varied. up to accuracy
Tracking
lit
the
S
When there 4 s between
was poorest
longest
(2
both
4).
when keys
were
tracking
s). Tracking
varied.
1 (Figure
affected
(HE and JA)
and S2 and between Tracking
S
accuracy
and S
2 3 was best when a
Sl and S2 and S2 and S3.
delays
between
imposed
(Table
key illumination
For two subjects
when these delay
of
illuminations
between
separated
was poorest
Furthermore,
were
manipulated
C.5 6) delay
Tracking
3).
250 ms.
keys
between
as the
Duration
(Figure only
when the
Interval varied were
varied.
S
between
1
were
long
(2 8).
and S2 was more disruptive S2 and S3.
were no delays
between
S2 and S3 had little
effect
ST and S2, on
1).
Fig. 4. Percent of trials on which the subjects chose the correct short (S - .5 s) key when the delay between Sl and S2 was either or long CL - 2 8) or a combination of short and long (S then L, L then S).
295
Horizontal horizontal (Table
vs vertical
sequences.
and vertical
sequence
Tracking
trials
accuracy
on
was approximately
equal
1).
DISCUSSION The major develop
purpose
a paradigm
tracking
of
procedure
visual
that
goal:
the
by manipulation
of
non-tracking
strategies
Pisecrete,
in this
(1979).
questions
such
sometimes
with
In such
as the
etc.
used
It
version
is
depending
pursuer
avoided,
or merely for
strategies
cited
in Ingle
Fleck,
1975)
overtake” pursuer
that
observed,
may also et
moving
al.,
to
al.,
1979).
may be used
strike
orient This
used
other
procedures
the
the
strategy the
prey.
a target
strategy “fixate target
the
moving
might
while
be celled
and overtake” moves
faster
being
species. and
1977,
(Lancaster e “fixate
strategy
be rate
pursued,
found
(Ingle,
upon the
6 and
ability
of
the
is
ineffective
end toads
at 3O’/s.
when pursuing target
might end the
been
fish
permit
moving
in different
has found--frogs
by gerbils of
is
have
This
also
object
be called
relies
used
sequences,
strategies
jacket
This
type
paradigm
track
and toads
and leather
as Ingle
the
the
will
object
strategies Frogs
pecking
be studied.
tracked
whether
be even
of
of
how animals
use what might
than
will
of
paradigm
tracking
of
exist.
1979)
ahead
the
could
and be different
behind
when the
ineffective--when
lit
effected
use
vs non-linear
prey
targets
The strategy is
is
stimuli.
to
matrix
version
present
different
to move faster
seed
the
procedure
linear
different
velocity
three
pursuing
consistently
of
arrangement
about
can move,
strategy.
fast
tracking
a much larger
accuracy the
questions
likely
the
Some evidence
of
of
upon the
et which
other
effects
at answering
objects.
for
successively
do with
an elaborated
on tracking
An elaborated attempts
potential
track of
discrimination
when implemeted
size,
discrimination
paradigm
or on e CRT-photocell-on-the-beak
target
to
es duration
unable
as they
have
Accuracy
such
are
conditional
by Blough
for
key.
to able
to
accurate
1982).
The delayed
keys
are
attempt
affecting
conditional
appears
lit
was to
factors
Pigeons
variables
subjects
research
The delayed
lest
Furthermore,
more useful
present
stimuli.
this
end choose
(cf.
the
investigating
was implemented
accomplishing keys
of
for
a moving
moving “predictive strategy than
the
sunflower
forward
(Ingle
tracking”
et end
is pursuer
or when
296
the
distance
between
Human infants catching
rapidly
move their to
evidence moving
it.
the
that
we are
light.
vision, that
the
the
pigeon of
keys
Hofsten.
with
the
visual
feedback
unsure
of
rewarded cues
influencing
great. of
object (1985)
is
strategy
1983).
when
They appear
and toward
it
to
so as
have described
important
when aiming
at
by which
animals
are have
accuracy are
able
pigeon
only
track
the
has a panoramic
a few degrees
proximity
to
the
of
location
field
visual
display
of arc
and
made
difficult.
a delayed choices
how our pigeons
the
occupied
was in close
seems to the
that
any strategy
In conclusion,
visual
type
(van
and Fleury
Hay,
The fact
that
observation
which
a different
objects
is
targets.
At present of
use
and pursuer
simultaneously Bard,
showing
target
seem to moving
hands
intercept
the
conditional contingent
potential of to
tracking perform
discrimination upon tracking
for
the
and for such
study
procedure a sequence
of
in
of
factors
elucidating
the means
behavior.
ACKNOWLEDGMENTS This research was supported by a grant from the Natural Sciences and D. Kennedy, and Engineering Research Council of Canada. R. Takai. K. Arbour also provided K. Arbour assisted with testing the birds. Send reprint requests to Donald helpful comments on the manuscript. I4. Wilkie, Psychology Department, The University of British Columbia, Vancouver, B.C. Canada V6T lY7.
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Ingle, D.. Cheal, M. D., 6 Dizio, P. (1979). Cine analysis of visual orientation and pursuit by the Mongolian gerbil. Journal of Comparative and Physiolonical Psychology, 93, 919-928. The nature and physiological basis of Kennedy, J. L. (1936). Psycholoxical Review, 43, movement discrimination in animals. 494-521. London: Routledge and KShler, W. (1925). The mentalitv of apes. Kegan Paul. Pursuit and prediction in Lancaster, B. S.. & Mark. R. F. (1975). the tracking of moving food by a teleost fish (Acanthaluteres spilomelanurus). Journal of Experimental BioloKy, 63, 627-645. (1983). Comparing frontal Bartinoya, C., Rivaud, S., & Bloch, S. and lateral viewing in the pigeon. II. Velocity thresholds for movement discrimination. Behavioural Brain Research, 8, 365-385. (1968). Interhemispheric transfer of a discrimination Hello, N. K. of moving patterns in pigeons. Brain Research, 1, 390-398. Borgan. C. L. (1930). The animal mind. New York: Longmans. Vision Bulvanny, P. (1978). Velocity discrimination by pigeons. Research, l8, 531-536. (1980). Movement detection by cats: Pasternak. T., & Merigan, W. H. Journal of Invariance with direction and target configuration. Comparative and PhvsioloKical Psycholoxv, 94, 943-952. Stimulus control of the pigeon’s ability to Pisacreta, R. (1982). peck a moving target. Journal of the Experimental Analvsis of Behavior, 37, 301-309. Siegel, R. K. (1970). Apparent movement detection in the pigeon. Journal of the Experimental Analvsis of Behavior, l4. 93-97. Siegel, R. K. (1971). Apparent movement and real movement detection in the pigeon: Stimulus generalization. Journal of the Experimental Analvsis of Behavior, l6, 189-192. von Hofsten, C. (1983). Catching skills in infancy. Journal of Experimental Psvcholoxy: Human Perception and Performance, 9, 75-85.
