72. Lilly, John C., and Alice M. Miller. 1962. "Operant Conditioning of the
Bottlenose Dolphin with Electrical Stimulation of the Brain." J. Comp. & Physiol.
Psychol.
Vol. 55 P. 73-79
Joul·,lalO/ComparaliL·e
and PnusiolugicolPsycnologU
196?, Vol. 55, No. 1, 73-79
OPERANTCONDITIONING
OF THE BOTTI,ENOSE
DOLPHIN
WITH ELECTRICIZL
STIMUL~'I'IONOF THE BRAIN'
J. C. LILLY AND i\. M. MILLER
Comm~clticaliolt
ResearcltInstil2tle,Mianti, norida
The stimulationof motivationalsystems
within the brain has been investigatedfor
with sea water sprays. The rostral restraint allowed the
animal to move the rostrum up anti down a distance of
smail-brained
mammals(Lilly,1958b).The and
2i".flukes
"ndlaterally
a distance
ofabout14in.Theflippers
were hanging free in the tank of water.
present report is on the investigation of the
Previousexperimentshad shownthat generalanes-
large brain of TzLrsiopstruncallls, the bottlenose dolphin of the Eastern United States.
thesia(Nembutal
or paraldehyde)
stopsrespiration
in
this species(Lilly,1958c,1961).Because
of the sus-
The bottlendse dolphin was chosen for these
ceptibilityof this animal to respiratorpfailureunder
experimentsto extend the comparative series anesthesia,a methodwas devisedto placeelectrodes
the brainof this animalusingonlya localanesof animals from those with brain weights within
thetic(Lilly,1958a).Iiigure2 showsthe principles
of
smallerthan thoseof man to brain weights this methodin diagrammatic
form.In brief,a small
equal to and greaterthan thoseof man. The pieceof hypodermic-needle
tubing(sleeve
guide)is
basic question is: Are these electricalstimula- h"mmered
throughthe skin,the blubber,the muscle,
tionsof thesubcortical
motivalionally
active and
islodged
inthesiinll.
Amandrel
which
carries
the
sleeveguideduringthe hammeringprocesscan be a~ithsystemsas demanding
in the large-br,zineddrawn
oncetheguide
penetrates
theinnertableofthe
mammalsas theyare in the smaller-brainedskull,
asinPhase
3,Figure
2.UptoPhase
3 thetipof
mammals (Lilly, 1958c, 1961)? A series of
the m""drelis heltiby the boneand oneis unableto
1959). The results of these measurements are
jectionof 1~ procainebeforethe hammering
process
carefulbrain-weight
measurements
weremade withdraw
it. Theprotruding
tipofthemandrel
isfreed
thecranialcavityat theinstantthatthesleeveguide
in relationto the sizeof the animals(Kruger, in
penetratesthe inner table. The site is preparedby in-
plotted in Figure i. The weightsare in the t"kesplace.Witha Tew
blows
ofa common
carpenter's
ranges 1,100 to 1,700 gm. The adult human
h"mmerontheupperendofthemandrel,
theoperation
is completed.
On the n~ithdrawal
of the mandrel,the
brainweights,as givenby Vierordt(1890),are skin
and blubberclose,leavingthe proximalend of the
in the lowerportion of this range. Earlier inves-
sleeveguidecovered.In the caseof the dolphin,a small
tigations
onbrainstimulation
in thesesystems markremains
intheskin.Later,using
thisasa guide,
were on the monkey, Ma·cocamulatta (80 to 105
gm.) (Lilly, 1958b).
The
present
account
is
concerned
with
O"eCa" find the track throughthe skin, blubber,anti
muscleandprobeforthe tip of thesleeveguideat any
timethat one~Yishes
to insertan electrodepair.
Specialelectrodes
are constructed
whichfit closely
showing
someresults
ofstimulation,
emphasiz-intothesleeve
guide.
i\nelectrode
driver
grips
theouter
ing those which differ from those for the
e"dofthesleeveguideuhichprotrudes
outoftheskull.
monkey.
An electrode pair can be lowered 1 mm. at a lime
M~TIron
throughtheguideintothebrainsubstance
bythistech-
nique.The trackis mappedmillimeterby millimeter.Al
The dolphinsin thisstudy wererestrainedin a small tl'e end of a day, electrodeand driverare removedand
laboratorytankand suspended
ill a slingin waterin the animalreturnedto the residence
tankfreeof any
intraspeciesisolation.Therostrumwasrestrainedby its
insertioninto a hole in a pieceof plywoodlinedwith
\vi'eSor connections.The experimentis continuedthe
f0llowi"6day, returningto the sameplacein the brain
polyurethane
foam.Theneckwashelddownward
bya Withthe electrodes.
The sleeveguidesplacedin~the
foam-covered
restraint:
bar. The dorsalaspectof the dolphin'sskullare No.15stainless-steel
(W316)
hyposkin was kept moistwith continuously
flowingsea de'mic-needle
tubing;the shieldinggroundingthe
water.Theexposedportionsof the headwerekept moist Oute'conductorof the electrodepairis No. 18stainless'This work was supported by funds from the follow- steel tubing.The innerelectrodeof the electrodepair is
ing sources:National Institute of NeurologicalDiseases
and Blindness,National Institutes of Health, Grant
B-3097;United States Air Force Omceof Scientific Re
search, Grant No. 61-62; Oflice of Naval Research Contract No. NR-2935(00). Iiacilities were constructed in
part under National Science Foundation Grant No.
11417. A portion of the work was done while the authors
were associated with the National Institute of ~Iental
insulatedfromtheouterelectrotie
bya tubeof Teflon,
and consistsof a pieceof No. 30 hypodermic-needle
tubingwhichextendsbeyondthe tip of the No. 18tut,ing 1 mm.ThisexposetltipaIldnearbyreturn electrode
give a localization of the stimulus at thresholtl over a
volumewhichhas a radiusof approximately1 to 2 mm.
n'ithin the brain. The diameterof the outer tubing is
1 mm.Suchan electrodeis stillenoughto givea straight
track through this large brain. A dorsal-ventral track
Health,National
Institutes
of Health,Bethesda,canbeupto120mm.inlength,
antia side-to-side
track
Mnry]and.
canbeupto200mm.
long.
With
theoseoftheproper
7·f
J. C. I,ILI,Y ANI) ~. M. ~IILI,EK
BODY LENGTH IN FEET a
INCHES
DIREC
TURSIOPS
ii
TRUNCATUS
I
1800
-
~ 1600
·-··
1 t II
t-
I
C~
SLE~VE`
i
I
,;~?~-
sn/N
I
~~,US
GUIDE
Y~
I
~
BoNE
DURd
5mm
W
2
1400
Frc. 2. Stages of penetration of sleeve-guide into
bone. (The sleeveis hammeredinto the bone by hitting
z
"m 1200~
the uI'perend of the mandrel.The directorkeepsthe
sleeve and mandrel lined up in the desired direction. At
the instant of penetration of the sleeve through the
1000 '
iOO
---~--~---~
150
200
inner tal,le--fhase
250
300
350
j---the mandrel is freed up from the
bone, and is easily withdrawn.)
BODY WEIGHT IN POUNDS
FIC.1~Brainweight,
bodylength,
andbodyweight tions are extremelycomplexand cover the
for the bottlenosedolphin(TzcrsiopsIrMncnlzcs).
(The whole known range of whistles, clicks, squawks,
solidcurveis brainweightversusbodylength;the and other complexnoises(Lilly,1958c,1961;
broken line curve is brain weight versus body weight.
Recently,a babydolphin5 ft. 4 in. long,weighing100
Ib.,\~asfoundto havea brainweightof 1,100gm.)
Lilly esMiller, 1961a,1961b)in a fewminutes.
In the case of the punishment or the self-
sl:opsystem, the first experiencewith a naive
electrical
wave
form(Lilly,
1960;
Lilly,
Hughes,
Al animalgivestheemission
ofdistress
whistles
vord, & Galkin,1955)observationscan be continued
formany
hours
perdayforseveral
weeks.
Reproducible
!Lilly,1958c,1901)
results are obtained testing a track
during
withdrawal
Self-Sfart
orRe~wardinR
Syslems
of the electrodes,on reinsertions to the same depth at
laterdates,andduringcontinuous
observations
at the
Illustrationsof the kindsof resultsobtained
samesite.
are given in I;igures 3, 4, and 5. With the
A minimum-effort
switch(Lilly,1942)is placed electrodesin a zone near the head of the cau-
above
thesaltwater
andoutofthespray.
Amovabledatenucleus,
a dolphin
canbequickly
induced
rod is placednear the rostrum of the animal,which
usuallypushesat onceand exploresits degreeof free- to turn on the switch to start a train of, stimuli
dom.Therodis usuallyadjustedto movein a vertical of 0.5- to l.0-sec. duration repeatedly.I;igure
direction when touched and return to the initial po-
3 shows the first such contacts
of a naive
sition
onrelease.
TheS quickly
determines
thatthisis dolphin.
the properway to push the rod. As soonas an active
spot(I,illp,1958~3)
is found,theS is permittedto push
When the S pushes upward on the rod and
therodin ordereitherto (a)turnthestimulus
onor makescontactwiththe radio-frequency
switch,
(b) turn thestimulusoff,depending
on thesiteof the th, t,,,,,,
electrotlein its brain,
RESULTS
With a naive, untrained S of a previously
this recordgoesdownward.When
the animal releases the rod, the trace then re-
turns to the baseline in an upward direction.
If S maintains contact with the rod, the trace
uninvestigat-edSpecies, the first: e?tperiences willstay in the downwardposition.The release
stimulal:ing either the "rewarding" or the of the rod triggers the stimulus. As I;igure 3
"punishing"systemstax the ingenuityof the shows,during the first 12 sec. of the record the
pushedoncein a very incisivefashion,in a
investigator.In the caseof monkeyswe noted S
hesitant fashion the second time, and the third
that stimulation of a rewarding or self-start-
ing system and a punishingor self-stopsystem time released it quickly. Immediately he
touched it again, releasedit again, and during
at threshold for the animal to turn on or to
shut off the stimulus apparently can be a very
mild experience (Lilly, 1958b). In General,
the next 4 sec, touched and released once per
second. The S started a train about once or
littleor novocalizstions
areemittedat these twiceevery4sec.fortherestoftherecord,
exthresholds
in the monkey.
StimulatioI1 of either of these systems in a
cept for briefburstsof aboutone per sec.,
three-quartersof the way through the record.
This kind of immediately successful bedolphinleadsto vocalization;
in the caseof the havior
of a nai\;e animal we did not i~ndin the
rewardingor self-startingsystemsthevocaliza-
OPERANTCONDITIONING
OF TI-IEDOLPHIN
'ij
TO&16H
RELEASE
4sec.
time
)------4
record
of
a
naive
bottlenose
dolphin.
(Everp
time
the
animal
touches
and pushesupFIc. 3. "Start-capture"
ward on a lever, the trace moves do~~nwardand stays downuntil the animalreleasesit. Each touchof the lever
is symbolized by a small peak in the downward direction. The electrode pair is situated at a depth of 95 mm.
below the surface of the rostral dorsal cortex in subcortical white matter near the head of the caudate nucleus.
The pulsepair repetitionfrequencywas60 per secondat a positivepeakcurrentof 6.0ma. with a train duration
df 1.0 sec. Every
10 sec.,
the animalwasgivena freetrainby theapparatus.Thisis theS's firstsetof touches
andreleasesin orderto stimulateits ownsubcortex.Dolphin7, 1957.)
~OUCH
~f~Ji~tf~BW~t~t~
RELEASE
tim~
FrC.4~A later"start-capture"
record.
(Theanimal
hasdeveloped
a regular
rhythmofpressing
thelever:for
thetrainduration
of1.0sec,theanimal
pressed
at a rateofabout1every2 sec.;andfora shortertrainduration
3,parameters
andlocithesame.Dolphin
7.)
--0.1 sec.---S pushed at a peakrateof3 persec.Cf.Figure
case of the monkey. i~Iany random contacts
with the radio-frequency
switch were necessary
before the monkey made the association be-
tween touching the switch and the onset of the
under water, the monkey in air. This animal
achieved rates of pressing (not shown on these
records) up to three pushes per second.
If during this long sequenceof self-starting
stimulus. In the case of the dolphins, the cause
and effect relatiyn between pushing on the rod
of the brain stimulus the current is interrupted,
made after the first two to three experiences.
immediately starts a series of vocalizations.
either by accidentor by design,and not al-
andtheelicitation
oftheeffectinitsbrainwas lowedto enterthebrainof theanimal,theS
"
L
From this point on this animal continued to
These sounds resemble those produced by the
push on the rod for manyminutes,
naive animal when first stimulated
is shown in I;igure ii. This record is quite com-
ing the switchto receivethe electricalstimu-
and l.O-sec.trains (Lilly, 1958b).
zation.
Asegmentofa laterportionoftheS'srecord
in these
During the previous sessions of presssystems.
parablewiththat ofsomemonkeysusing0.5- lus, the S worked silently without any vocaliA monkey can average three limes per sec-
I;igure 5 showssuch a vocal effect in a
ondwiththeminimum-effort
switchwith0.1- graphic
form.Attheleftofthefigurethecursec, trains. The dolphinwith O.l-sec.trains can rent has just been interrupted. This record is
reach these rates also. The dolphin is working the rectifiedintegrated output of an amplifier
~. (3. LTLLY AND ii. R·I. MILLER
jt~_7-i--
I
II
·~:
-\·
QMg---eOFF (sn dume9s)r8
·---
··~.
oisuse
4 sec.
Iirc.5. Vocalization
afteraninterval
of'Ltart-capture"
activity.
(An
upx·ard
rieilccrion
in~his
recot~d~lts
vocalactivity
ofsomesort.Thepenis deflected
by the
of therectifieti
electrical
acoustic
output.The
loutierand the moreprolonged
the noise,the farthertheintegral
recordstartsimmediately
afterthecurrentis removed pen is denectedin a g~venshort periodof time.This
fromthe stimulating
electrodes
by turningo~ theswitch
to the animal--"ondummy''--sothat.S no
canstimulate
itself.
Within
a fewseconds
ofthattime,theS
starts to vocalize,in this casecontinuingits longer
vocalization
of40sec.Parameters
areasinFigures
3
and4. Previousto the timeof thisfigure,the
S had foraninterval
beennonvocal
while
stimulating
itsownbrainbypressing
fed by an air microphonesuspendednear showedthat one can increasei-helolalamounl
the lever at a rate of about one per sec. Dolphin 7.)
the blowhole.The seriesof air-bornesounds
daily vocalization, but one cannot induce
emittedhereconsistedofwhistles,bronx--cheer-of
the
monkeyto usevocalization
thewayit uses
like noises,squawks,blats, very rapid and its hands
in order to start a desiredstimulus.
loudclickings
and otherextremely
comp]er: dolphinapparently
usesvocaloutputsand
vocalizations
(Lilly& R/Iiller,
1961a,1961b), The
may even prefer vocczlization
to pushinga lever
theresultof thewithdrawal
ofa "rewarding"withits beak.As in the case
of the human,
~p~~ely
reinforcing)
experience
inatrained
t)lere
isless"central
effort"
forvocalization
Anotheranimal(Dolphin8, 1958)stimulated than there is for the monkeyand for the action
in a similar region was
of thelargermuscles
of thebody(Lilly,1958h;
induced
tovocalize
ina Liily&Miller,
1961a,
19Blb).
particular uray in order to obtaiII the brain
stimulation.
Fromitsvocalizations
a whistle,in SlopSyslenzs
air, of a particularfrequency,
duration,and As was stated above in the case of the naive
amplitude was chosen as the sufficientcondition for giving a braiI1 stimulus.
animal,we have found that dolphinstend to
emit a more stereotyped series of vocalizations
It was found that the S cluicklylearned whenwestimulatea negative,
or "punishing,"
some of the rules of this game. The S emitted
whistles of varying characteristics from one
whistle to the next. The proper whistle was
quickly shaped up. Every time the S whistled
a particular pitch, it was "rewarded"; within a
or "stop"systemzone.Themonkeycaneasily
be trainedto shut off a stimulusat a critical
threshold,and the thresholdfor such phe-
nomenais determinedby means of a crescendo
train of stimuli (Lilly, 1958b). The train of
periodof 10min.S waswhistlingthis particu- stimulistartsnearzero,buildsuplinearlywith
]ar
duration,
pitch,
andamplitude.
Asinthe respect
to time,i.e.,thepulses
increase
in
case of the monkey,it attempted to reduce the
effort put out in order to obtain the reward: it
intensity over a period of time, are auto-
maticallyshut offby the apparatus,and after
triedto lowerthe amplitude,keepingthe pitch a pause are started once again. At any time the
and duration constant; by thewithholding
of. S callshut.offthecurrent.A typicalresultis
reward, it could be
again.
induced
to whistleloudly shownin Figure6.
The S is given a switch which shuts off the
A similar series of experiments with rhesus stimulus the instant that:S releasesits contact.
monkeys in similar systems within the brain
This task a monkeylearnsrelativelyrapidly
i
OPERANT
CONDITIONING
after rather few trials. If the peak of the
crescendo is carried· high enough to cause vio-
OI: THE DOLPHIN
77
6 illustrates the threshold level at which the S
pushes the lever to shut off the current and
lent L'escape"behavior, a trained monkey will the immediate cessation of pushing when the
shut off the stimulus at a threshold
most of
current is turned off. It is to be noted that in
the time well below that where one can detect
this figure there is no particular time relation-
any clinicalsignsof increasedautonomicac- ship betweenthe respirationsof this S and
tivity or "escape" behavior (Lilly, 1958b).As the pushing of the switch. The head movewas reported previously,the threshold for de- ments of respirationdid not interfere with the
tection by the monkeyof stimulationin such a head movements necessary for moving the
system is about one-third to one-fifth the rod.
threshold for its imperative shutting-off of the
stimulus. At currents only a few percentage
As in the case of the rewarding systems, the
S did not vocalize during this period while
pointsabovethe imperativevalue,the monkey pushingthe switchin order to shut off the
is no longer able to shut off the stimulus. A
current. In contrast with the stimulation of
physiological
state of violentescapeactivity "rewarding"systemsthere was no vocalizatakes over all the paths to motility.
tion when the current was cut off from the S.
Similarresultshavebeenfoundwiththe Oneco;lldobtainvocalization.,
thatis thedisf
dolphin.The only dramatic differencesare the
tress call, at a threshold somewhatabove that
task and its developed control of the escape
to shut off the current.
speedwithwhichthenaivedolphin
learnsthe at whichtheS waspushing
theswitch
inorder
pattern. Within the order of 3 to 20 trials, the
Figure 7 shows a quantitative plot of the
dolphinis pushing regularlyat the value it will analysisof 588 successivecrescendoexposures.
be pushing several hours later. If one Shuts off The total number of times that the S shut off
the current, the dolphin immediately stops
the current was 564 in 588 trials, i.e., its over-
pushing the lever. The monkey may push it
two or three times beyond the instant at which
80
the current is shut off by the operator. Figure
P
60
RELEASE
w
~
3.0
i
~
I
o:
bD:
TOUCH
~L040
RESPIRATION
1.5n
~B
~z
,,
lime
ONc-tOFF
ton dumny)
3Osec
FIc. 6. "Stop-capture" record after a short period of
2
0'20
z
~I
~
~S
-I
learning.(Respirationis shownby a slight downward
deflection of the pen. Increasing sfimulus intensity is
shownby an upwarddeflectionof the pen occurring
regularlyevery13 sec.Afterthe fourteenthupward
D
0'
4
8'
0.3
TIME
1NCRESCENDO.
SEC.
FIc.7. Analysis
ofthedistribution
of thresholds
for
de8ection, the current was shut off, i.e., the S was nqt
"self-stop" activity. (The dotted curve is the course of
allowedto receiveany further stimulationin this por-
the crescendoof current with respect to time if the
tion of its brain. The left half of the record shows the
animal dots not shut off the current. The solid curve is
accuracywithwhichthe S waschoosing
the valueof the distributionof frequencies
of occurrence
of the
currentat whichit wouldpush the shutoffswitch.No
shutoffsat each valueof currentfor the exposureto a
errors
onthepartoftheanimal
areshown
onthisrecord. total
of 588 crescendos. Of these total exposures, the
The righthalf showsthe peakcurrentto whichthe animalshut off 564.The baselinegivesthe timein
apparatuswouldcarrythe currentif the S had not shut
it off. There is no regular relationship between the
seconds after the beginning of the crescendo. The ordinate
for the solid curve
is the number
of shuto~s
at
respiration
andthepushingof thelever.Theelectrodes eachvalueofcurrent;forthedottedcurve,thecurrent
are locatedin the thalamusat a depthof 60 mm.below at the instantof shutoffin milliamperes.
Thepeakof the
the dorsal corticalsurface.The electricalstimulusis solidcurve occursat approrcimately
4.1 sec, after the
occurring
at a rateof60pulsepairspersec.increscendos beginning
of thecrescendo,
i.e.,at a currentof 1.7ma.
startingat a currentof apprordmately
0.3 ma, and Beyondthis peak,as in the caseof the macaque,
the
risingto a peakof2.9ma.unlessshutoffearlierbythe animalbecomes
tooemotionally
upsettopushthelever.
animal.The crescendorepetitionrate is one every 13 The parametersof the electricalstimulusare 60 pulse
sec.controlledby the apparatus.Dolphin6-1957. De- pairsper sec.,from0.3 to 2.9 ma.,lasting8 sec.and re
tailed analysis of such records is shown in Figure 7.)
peated once every 13 sec. Dolphin 6.)
78
J. C. LTLLY.4ND ~. M. R·IILLER
1
:1
FIc. 8. Locationof the electrodesin the Chalamus.
(Thiscrosssectionillustratesthe sizeof the dolphin'sbrain
and showsthe locusof the stimulationsfor Figures6 and 7. The trackdownto this point wasexplored,and is seen
filledwith the No. 18hypodermic-needle
tubingof the outer,groundedelectrode.Somemotorresponsesincluding
respiratory ones were found above this negative zone. Dolphin 6.)
all scoreof 95.9% was comparableto that of a
appearance of clinicalsigns. In the case of the
well-trained
monkey.A detailedanalysisof the dolphin,the first:clinicalsign is vocalization,
currents for each of these shutoffs gives the
the distress call. This call is elicited only by
statisticaldistributionshownin this figure.As trains not under the control of the dolphin
in the case of the monkey, there is a small which are of a current larger than 1.6 ma.
"detection"peak at the beginningof the step
(0.3ma.)of the crescendo.
Mostof the imperative shuto~soccurat a thresholdof slightly
over 1.7ma, with theseparticularparameters.
Asin thecaseof themonkey,thisimperative
peak precedestin termsof risingcurrent)the
Afterthe firstfewexposures,violentescapebehavioris not elicitedat any of the currentsto
whichwe exposedtheseSs: the most activity
elicitableis brieflyshutting both eyes tightly
las theseSs do in painfulneedlepricks),sharp
withdrawals,jumps of the head and body, or
OPERANT
their "warning
i,
CONDITIONING
DOI,PHIN
79
or hori-
wanted, or positive, or rewarding stimulus or to
zontal rapid head-shaking. This apparent
cut off an unwanted, or negative, or punishing
control is contrasted with the violent pushing,
pulling, running, and biting of the macaque
similarly stimulated. However, after a long
period of coniinement with humans, a dolphin
begins to act more violently. A naive dolphin
in restraint tends to inhibit the violence for
unknown reasons. Over months it tends to become less inhibited toward humans and its
restraints. If we remember that a 3CO- to 400Ib. dolphin has great muscular power and
could destroy, or at least badly damage, itself,
the restraint systems, and possibly even the
investigator, it is very surprising to find that
stimulation started by the apparatus. In contrast with the monkey, the dolphin uses its
vocal output when this is effective in order to
modify the responses obtained from the environment tin terms of brain stimulation). The
dolphin can inhibit violent escape behavior
caused by "punishing" brain stimulation; the
macayue does not. The large brain of the dolphin thus affords: (a) faster learning, (b)
greater control over reactions tostimulation of
subcortical systems which are motivationally
active, and (c) control and use of vocalizations
to obtain "rewards" and to stop L'punish-
dolphins
ments."
are
negation"--vertical
OZ; THE
not
more
violent.
under
such
"punishing"
stimulations
as these.Theirbe-
havior resembles that of a very strong-minded
human being who, during bouts of intense pain,
can sometimes attenuate his desperate behavior more than can the "lower" animals.
The particular
REI;ERENCES
RRUGER, L. The thalamus of the dolphin (Tursiops
Iruncelus) and comparison
with other mammals.
J.,,,p. Neltrol.,
1959,3, 133-194.
LILLY,
J. C. Electricalcapacitancediaphragmmanom-
spot in the brain stimulated
eter. Rev.scienl. Inslrutn., 1942,13, 34-37.
for the previoustwo figuresis shownin Figure LILLY,
J. C.Electrode
andcannulae
implantation
in the
8. It is about
8 mm. lateral
to the mid-plane
in
brain by a simple percutaneous
1958, 127, 1181-82. (a)
method.
Science,
the S's thalamus. Presumably this is a portion
L,,,,~ J. C. Learningmotivatedby subcorticalstimu-
of the system which in the human causes the
lation: The "start" and "stop" patterns of be-
thalamic pain syndrome when stimulated
havior.In H. H. Jasper,LorneD. Proctor,Robert
(iLlonnier,
1955).Othersuchspots have been
S·
K"ighto",
William
C. Noshay,&of Russell
T.
Costello (Eds.), Relicular formalion
l~e b~ain.
studied
also.
SUM~AKY
It is shown
that
Boston: Little, Brown, 1958. Pp. 705-721. (b)
LILLY, J. C. Some considerations
regarding basic mechanisms of positive and negative types of motivations.
AND CONCLUSIO~S
it is feasible
to place
elec-
trodes into the large brain of the unanesthe-
An2er. ~. Psycl~iat. 1958, 115, 498-504. (c)
LILLY,J. C. I,earning motivated by subcortical stimu-
tized bottlenose dolphin (I'ursiops 1IZIIZGUIUS). ]ation:The"start"andthe"stop"patternsof beUsing roving electrodes, one can demonstrate
separate zones of the brain which are positively
and negatively reinforcing, i.e., in which "self-
start" and "self-stop" activity can be elicited.
It is shown that the dolphin learns these tasks
more rapidly than does the monkey. It is also
shown
that
in a rewarding,
or positively
rein-
forcing zone, the animal tends to vocalize in
complex and startling ways. It vocalizes when
first
stimulated
in this
area
or when
stimula-
havior.
Injury
and excitation
of brain by electrical
currents.In ~leclricalsl2cdies
on Iheunaneslketized
b,,;,,~ N,, y,,k: p,,l B~ Hoeber, 1960. Pp. 78105.
L'LLY,J. C. il~IaW
and dolpltin.Garden City, Long
Island: Doubleday, 1961.
LILLY, J. C., HUGHES, J. R., ALVORD, E. C., JR., &
GALKIN,
T. W. A briefnoninjuriouswaveformfor
stimulation of the brain. Science, 1955, 121, 468-
469.
LILLY, J. C., & ICIILLER, A. M. Some audible sounds
emitted by the bottlenose dolphin. Sciellce, 1961,
133, 1-5. (a)
tion is withdrawn after a period of "self-start"
LILLY,
5. C., &MILLER,
A. M. Vocalexchanges
between
activity.
Thus me conclude that the behavior of the
dolPhins.Sca`elrce,
1961,134, 1873-1876.(b)
MONN'"R~M. La stimulation electrique du thalamus
bottlenose
dolphin
differs
from
that
of the
chez
I'homme
r~sultats
nettrol., 1955, 93, 267-277.
somatotopiques.
Rep.
monkey in the following respects: Like the
V,,,,,,~ H. DasMassenwachsthum
derKorl'erogane
monkey, the dolphin uses any available external somatic motor output in order to push
switches to stimulate its own brain for a
des Menschen. Arch. nltal. Physiol. nnal., s62,
1890.
(Earlp publication receivedJune 19, 1961)