JPRS ID: 8475 TRANSLATIONS ON USSR SCIENCE AND TECHNOLOGY BIOMEDICAL AND BEHAVIORAL SCIENCES AN ELECTRO - PHYSIOLOGICAL INVESTIGATION OF THE DOLPHIN BRAIN

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PHYSIOLOGICAL INVESTIGATION OF THE DOLPNIN 61~AIN
22 MAY i979 CFOUO i6r79~ - i OF 2

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JpI~.S L/8475
22 May 1979
~
TRANSLATIONS OP~ USS R SCIENCE AND TECHNOLOrY
BIOMEDICAL AND BEHAVIORAL SCIENCES
(FOUO l6/79)
' AN ELECTROPHYSIO~.OGICAL INVESTIGATION
OF THE DOLPHIN BRAIN
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JPRS L/a475
22 May' ~9 79
TRANSlATIONS ON USSR SCIE~;CE AN~J TECHfdOLOGY
BIOMEDICAL ArdD BEHAVIORAL SCIENCES
(FOUO 16/'l9 )
,
AN ELECTROPHYSIOLOGICAL INVESTIGATION
OF THE DOLPHIN BRAIN ~
rioscow ELEKTROFIZSOLOGICHE5KOYE ISSLEDOVANIYE MOZGA DEL'FINOV
in Russian 197g signed to press 27 Jun 78 pp 2-6, 86-160,
201-208, ~11-213
.
CONTENTS PAGE
MAItINE MA1~fALS
An Electrophysiologica~ Investigation of the Dolphin Bra~n 1
Chapter 3. betermining the Char~cteristics of Hearing in
Dolphins From the Electri.cal Reactions of the Cerebrnl
Cortex 7
Bibliography........... ~
85
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MARINE M~AMMA,IS '
AN ELECTROPHYSIOLOGIOAL INVESTIGATION OF TH~ DOLPHIN BRAIN
Moscow EL~KTROFTZIOLOGICHESKOYE ISSLEDOVANIYE MOZGA DEL'FINOV in Ituasian
1978 signed to preas 27 Jun 78 pp 2-6, 86-160, 201-2b8, 211-213
. (Annotation, introduction, table of contente, Chapter 3 and bibliography
from book by A. Ya. Supin, L. M. Mukhametov, et al., Izdatel'stvo Nauka, '
1200 copies, 215 pages]
Annotation
(Text] This book su~arizes experimental�data in an area not previously
investigated--the physiology of the brain of Cetaceans~-which are of
interest for both comparative physiology of the nervous system and general
neurophysiology. These data include information on the unusual nature of
the arrangement of the sensory areas of the cerebral cortex in dolphins and
the unusual nature of the generation of electrical reactiona (evoked poten-
tials) in the cortex; the unique nature of the regulation of wakefulness
- and sleep (the alternating development of sleep in the two cerebral hemi-
spheres); features of the suditory system that can be revealed according to
the electrical reactions of the cerebral cortex. Methods of electro- ,
physiological inuestigation of the dolphin brain are discussed.
The book is intended for biologists specializing in neuromorphology and
neurophysiology, zoology and ecology.
It has 102 illustrations and a bibliography of 160 titles.
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IN'1'lt()UUC'CxON
The intere~t which dolphin~ have aroused among re~earchere working in the
moet varied ~re~s of gcience is caused by many factors and ia, eo a con-
sidcrable exCenC, ro be expecCed, Hardly ~ny orher animal which ha~ became
~~vxtlable fnr reaearch has immediACely posed such a number o� dtverqe prob-
lems Eor speCialiaCs in varinue fi~lds. The chgracteristics of these
~nimals' behavior, which is organized in an extremely complex manner; their
sCriking ability to learn quickly and to enter inCo conegct with man; Cheir
developed system of communication by sound; eheir highly-develop~d tiearing,
including the ability to percetve ultrason~.c eignals; eh~ir cnpacity �ur
active echolocation; their brain, which is enormous (for animals); ttidir
ability Co move about in warer at high spe~ds wiCh a relatively small
_ energy ~xpenditure and Cheir capacity for prolonged diving--a11 ~f this i.s
far from being a complete lisC of Che anatomical and physiologica~. charac-
teristics of'dolphins, which deserve r.he tnost careful seudy.
Do dolphins (nr CeCaceans in general) actuaily constitute a group of
animals Chat are exceptional in their morphophysiological ~:haracterisCic9
and deserve very special artention from researchers? H~rdly. Rather,
dolphins provide an instructive example of how expanding the range of
animals which are the ob~ects of detailed morphophysiological atudy can
be unexpectedly imporCant and useful.
Unfortunately, a significant gap exists between the tremendous �number of
. animal species which are wiChin the sphere of attention of. zoologists wiCh
a broad range of specialties and the number of species which hnve become
model ob~ects for detailed study of their morphological or physiological
organization. This limitation of the number of objects for detailed study
is, in fact, essential. The constant intensification of the research
requires a thorough knowledge of the ob~ect, which would scarcely be pos-
sible with a large number of objects. On the other hand, for a compar:tson -
of the various wor.ks, it is advisable for studies to be performed of the
same standard ob3ects. Consequently, the selection of a relatively smail
numher of animal species as standard laboratory sub~ects is dictated by the
logic of biological research.
This situation also has a reverse side, however: one cannot always be con-
- fident that the data discovered in the course of the investigations reflect
universal phenomena and do not merely describe the features of the sp~.cies
being stud~ed or of the group to which this species belongs. For this
reason, detailed morphophysiological study of new species may some~imes ~
lead to unexpected results.
This is precisely what happened when dolphins in captivity became available
fQr detailed study as a result of the opening of oceanaria-dolphinaria in a
number of countrf.es. In this case, the morphologists and physiologists did
not acquire simply one more object for study; but an animal differing radi-
~cally from the well-known laboratory mammals. In the course of their
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Cvolution, Ch~ Cetnr.enn~ quite enrly se~~arr~r~d f.rnm the main trunk n� the
cvalueton~ry tre~ of th~ mammuls and followed nn extended d~veYopment~l
paCh, independpnt of the land m~mmals. ~or ehis re~~on iC is quite naturnl
Chnt many of. thp problemg of environment~l adraptxtion w~r~ solved in s
uniqu~ wny in this dirertion of Che evnlutinn af. mammdls nnd rhgt mnny
fciiturtq of rhe Cetucenns' morphc?physiolu};ical orgnnizatiun proved tn be
~ unusual Eor researchers ~orking with lnnd ldborarory ~n3m~ls.
An understanding of the fnct Chat the interest in dolphins is due to their
difference from the w~11-studied labor~tory animals rather thAn to gny
uniquenes~ of their position in the anim~l world rev~als the fullesC sig-
nif.icance of Che invesCigations being made of dolphins. They ~re import~nt
primar~ly because they can rid us of a one-sided concept of ~ number of
~specCs of mammalian orgttniz~tion. It should be particularly emphasized
th~t ehe comprehensiv~ study of an uriusual order such as the Cetacenns is
def.inirely of exceptional interest for an understanding of many questions
of m~mmalian evolution, including the evolution of ehe brain.
One of. the most important and interesting questiona in the study of.
Cernceans concerns the orgattizabion of their nervous sysCems. The Cetacenn
brnin is one of the largest in the animal world. Altltougli it is difficult
to compare the dimensions of the brain in animals with diEfering body size
(neither absolute nor relative brain weighL, nor the product of absol.ute
weight multiplied by the relative weight i~ considered to be ~ universal
criterion for such a comparison), the enormou,s and complex brain of the
dolphins, neverCheless, could not fail to dr~w extremely close atCenCion l
f.rom neuromorphologists and neurophysiolo~;ists.
Tlie study of the brain can be effective, however, ~nly if there is a har-
monious combination of morphological and physiological investigations. '
But while the study of the morphology of the Cetacean brain (which is more
accessible) has a relatively short history, until recently there were prac- ~
ticall.y no neurophysiological studies at all.
There are quite definite reasons for the inadeqt;ete development of neuro-
physiological, and particularly electrophysiological investigatiuns of Che
Cetacean brain. Despite the fact tha[ dolphins have been held in captiviCy
successfully for many years, until recently they remained expensive and
unavailable animals, the experimental use of which was extremely limited.
Furthermore, many diffi~ulties directly related to experimental methods are
encountered in the performance of neurophysi~logical investigations of
dolphins. The characteristics of the dolphins' structure and physiology
make it difficult o r impossible Co apply to them many well-developed methods
that are us~d in working with the usual laboratory animals. In addition to
the fact that the methodology for working with dolphins had to be developed
anew to a significant extent, there were, at the same time, ~erious diffi-
culties to be overcome, stemmfng from the problems of access to the brain
of. these animals, the need to take special measures to preserve their lives
during operations and expe.riments, etc. At the same time, the methodological
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pr.ab lems ~re int~nsif~ted b,y l~um~ni~aricin renuirem~nt~: t~~ wnrking wi.th
hi.f;hly nrgnnizcd anim~ls, r~ cate~ory whir.h de~initely i~icludes dalph:ins,
it is cg~enCial to exercise sCrice conrrol ~~nd to limit tt~~ trc~umn~ic effect
of Che research merhods ehar are used, avoiding gross vivisec~ion.
Tl~e inCeresC in Che neurophysinro~y of Che dolphin br~in, however, h~s
forced us to look for ways ~n overcome Chl?q@ difficulties. ACtempts have
repearedly been made ro perform physiolo~tcal investigations of dol~hin
brains (Lan~worthy, 1932; Yanagisawn er al,, 19G6; Lende, Adikman, 1968,
and others), alrhough the ma~oriCy of these ~tCempts hav,: yielded few
resules or have been unsuccessful due td rhe above-menCioned methodolo~ic.71
difEicul.rie~. Until recently the literature contained niily igolnted reports
of successful work on the electrophysiological study of the dolphin brain.
Among ehe pioneers of such invesCigations, Bullock and his coaurha,:n
(Bullock et al., 1968) should particularly be mentioned.
The Laboratory of the Evolution o� the Sensory Systems of the Tnstitue of
the ~volutionary Morphology and ~cology of Animals of the USSR Academy of
Sciences, which ~.s represented by the authorial collective of Chis book,
is one of the few laboratories ttiat conducts electrophysiological invesCi-
gations of Che dolphin brain. Various aspects of the morphological and
phy~iologic~l organizaCion of the brain of the dolphin have been sCudied by
this laboratory for a number of years. Furthermore, studies of a comprehen-
sive ~nd systematic nature are considered to be one oE the main goals of
Chis laboratory.
It goes with~ut sayinR rhat certain limitations on the work volume were
nevertheless ine~itable. The cerebral cortex, the highest analyzing center,
was c}~osen frr more detailed study, although other brain centers were
studied f~lon~ with it. A number of issues requiring pri~rity investigation
were outlined. They included, par.ticularly, the problem of the mafn fea~tures
of th~~ general morphophysiological organizarion of the dolphin brain, and
~specicilly of the cerebral cortex. The dolphins' analyzer activity was
thou~ht to be the second important problem, with the dolphins' key, acoustic
an~~lyzer attracting priority attention. Finally, it appeared that the
regulation of sleepin~; and waking states--in cannection with features of t}~e
dolphins' biology and behavi.or--deserved special artention.
In practice, during the process of the investigations all these problems
proved to he interrelated to a significant degree, and were worked on as a
sin~le unit. The division of all the re~ulrs obtained in thes~ areas, how-
ever, proved Co be convenient for purposes of their systematization, and
formed the basis for the contents of this book.
The major part of the investigations, the results of which form the basis of
this collective monograph, was carried out during expeditions to the Black
Sea, in the area of Anapa and Novorossiysk, rather than at stationary bases
or in oceanaria. The authors note with ~ratitude that it would have been
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~o~ o~ricT,~ us~ nNLY
' im~~assible tn c:7rr.y out the rey~~rnh pro~;rnm of Che expedi~inn~ withn~r ~he
help nf a large number oF ~19qiSCgnCs wi~n p~~rticipnred in thE l~bor-cnnsuming
~ ~uxil,inry work. '1'lic ~7uthors would lilce ~o express sincere graeiCude ro
everyone who assisted wieh the ~.nveseigttrions. The authors will also be
grateful Co their colleagues for ~ny criCical comments,
TAI3L~ bt' CONTENTS
InrrnducCion. . . . . . . . . . .
Ch~tpter 1. A Brief Morphulogic~l Descr.~ptinn of.th~�Ce~acean Brnin
1. Cephalizatinn nnd General Properties of. Che Bruin
Su1ci and Convolutions of the CerebrAl Cortex of Uolphins
3. Microscopic Structure of Che Dolphin Cerebral CorCex
Summ~ry . . . . . . . . . . . . . . . . . . . . . . . . . . . .
_ Chaprer 2. Neurophysiological F'egtures of the Dolphin~Cereb ral Cortex
1. Conditions for Experiment and Stimulus . . . . . . . . . . . . . .
2. Localizarion FeaCures of Evoked PoCenCials in the CorCex
3. I~ocaliz~Cion of the Auditory Pro~ecCion Area of the CorCex
in the Dolphin T. truncatus . . . , , , , , , , , , , , , , , , ,
4. Types of Responses and Functional Subdivisions of the Auditory
Area of Che CorCex . . . . . . . . . .
5. LocalizaCion of the Visual Pro~ection�Are~ oP~Che~Cortex in~the~ .
Dolphin T. truncaCus . . . . . . . . . . . . . .
6. Types of Responses and Functional Subdivisions of the Visual
Area of the Cortex . . . . . . . . . . . . . . . .
7. Localizarion of the Somatosensory Pro~ecCion Area.of the �
Cortex in the Dolphin T. truncatus
8. r(ap of the Sensory ProjecCion Areas of the Cortex.in the�
Dolphin T. truncatus . . . . . . . . .
9. Pro~ection Sensory Areas in the Cerebral Cortex�of the
P~rpoise Ph. phoc~ena . . . . . . . . . .
1G. Tl~c Question of the ~xistence of Pdlysensory�Zones.in the
Cerehral Cortex . . . . .
11. reatures of the Generation.oE Evoked.PoCentials in�the.
Cerebral Correx . . , . . . .
12. Features of the Arrangement o�.the.Pro~ection Sensory Zones
of the Dolphin Cerebral Cortex and Problems of the Evolution
of Che Cerebral Cortex of Mammals
. . . . . . . . . . .
Summary . . . . . . . . . . . . . . . . . . . . . . .
Chapte r 3. Dete rmining the Characteristics of Hearing in.Dolphins
From the Electrical Reactions of the Cerebral Corre x..
1. Microelectrode InvesCigation of the Pulse Activity of
Individual Neurons of the Auditory Cortex
2. Conditions f~r the Investigation of Evoked Potentials of�the�
Auditory System . . . . . . . . . . . . . . . . .
3. Evoked Potentials at the 5tart and Stop of the Sound. ~Force�
Curves and Absolute Thresholds of Excitation for
Evoked Potentials . . . .
. . . . . . . . . . . . . . . . . . . . . ~
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4. Reactions ot Che Cortex ro Chan~es in the ]:nt~nsity ot the
Sound. Diffcrential Threqholds for Lntensiey , , , . . , . , . .
5. 'C-ime SummuCion . . . . . . . . . . . . . . . . . . . ~ . ~ . . . .
6. Depression of the Gvoked Potenti~l~ WiCh ~1n Incre~h~~ in ~he
Duration of Che Stimuli . . . . . . . . . . . . . . . . . . . . . .
7. Dependence of the Evoked PnCenCials o� the Auditory Cortex on the
Characteristic Slope of tlie Front of the Auditnry 5timulus
- 8. ReacCions o� Che Dolphin's Cerebral Correx to a Cr~ange in
5ound Frequency. Thresholds of ~'requency Discrimin~ition
9. ReacCion CdpaciCy of the AudiCory Cnrtex Eor Reprod~ ~Cion With
Pa-i_r.e~ and Rhythmic Stimuli . . . . . . . . . . . . . . . . . . . .
� 1Ci. P.eactiuns of ttie Auditory CorCex ~0 5ound Stimuli nel.ivcred
Agninst a Background o~ Inrerference . . . . . . . . . . . . .
11. Functional Differentiation of Various Sections of the AudiCory
CorCex and Various Componenrs of Evoked PoCentials .
12. The Presence of Specialized (DetecCor) Elements in Che
Auditory System . . . . . . . . . . . . . . . . . . . . . . . . .
13. Clectrical Reactions of the Inferior Colliculus and the
Corpus Genicul~tum Mediale to Sound Stimuli . . . . . . . . . . .
Summhry . . . . . . . . . . . . . . , . . . . . . . . . . . . . .
Ghaprer 4. I:lecCroencephalographic InvesCig~Cion of the Cuncrional
SCates of the Dolphin Brain . . . . . . . . . . . . . . .
1. Conditions for an Experiment in ~EG-Investigation of Sleep
'l. Electroencephalogrnphic Characteristics of Waking and
Sleeping States . . . . . . . . . . . . . . . . . . . . . . . . .
3. MainCen~nce of Respiration in Natural Sleep . . . . . . . . . . .
4. Change in the Electrocorticogram Under the Influence of
Narcotics . . . . . . . . . . . . . . . . . . . . . . . . . . . .
5. In terhemispheric Asymmetry of Functianal Stares of Che Brain
6. The Daily Rhythm of Wakin~ and Sleeping in Dolphins
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Conclusion . . . . . . . . . . . . . � ~ . . . . . . . . . . . . . . . . .
Bibliography . . . . . . . . . . . . � . . . . . . . . . . . . . . . . .
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
COPYRIGHT: ]:zdat~l.'stvo "Nauka", 1978
12151
CSO: 81~+~+~1107
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~ ~ MARINE MAMMEIL5
_
CIlAP7'f:R 3. D~T~RMINTNG 'CFIE CHARACTERIS'CICS OC EiCARINC IN I)OLPHYNS ~ROM TH~
CLECTRTCAL ItCACTIONS OF TNT: CCItC4RA1~ COItTCX
By recording the elecCricnl reacrions of Che cerebral cortex to var3ous
ttfferene seimuli, importttnC information may be obtained on tl';~e mechanisms
of analysis of these stimuli and on the properties of Che co~rresponding
an~~lyzer. If one eakes into consideration the exceptional developmenC of
dolphins' acoustic annlyzer, as well as the fact thAe they h~ve the capacity
for ~coustic locatiott, iC is clear thaC the acoustic analyzer attracts the
;
greatese attention when studtes of this type are made of the dolphin's brain.
The characteristics of hEaring in dolphins have drawn the attentinn of a
number of researchers. tlfter ie was discovered that dolphins can perceive
ultrusonic frequencies (Kellog, Kohler, 1952; Schevill, Lawrence, 1953),
, aCtempts were made to measure more precisely the characteristics of Che
9olphins' auditory system, particularly its absolute sensirivity and
frequency-threshold charACteristics.
~ '!'he first sufficiently detaited quantitative studies of hearing in dolphins
~~er~~ made on specially rrained animr~ls under free behaviora2 conditions.
'rh~ hi~fm~il was taugh t to perform a certain movement in response to a sound
si~n7t. By gradually reducing the intensity of the signal, it was possible
to esrablish th e minimum intensity aC which the signAl cnuld still be per-
ceived by the animal~ w:~ich was indicated by the performance of the control
movement. This value was taken as the threshold of perception of the signal.
By using pure tones of vr~rying frequencies as the signals, it was possible
to determine how the sound perception threshold depends on its f requency,
i.e., to draw up an audiogram.
rteasurements made in this way (Johnson, 1967) showed thut the mittimum thresh-
olds of auditory perception in dolphins T. Cru�catus are ohserved at frequen-
cies of about SO ktlz, where they are (if the original data from the system
of units used by th e a~ithor ~7re converted into the Inte rn ational System) less
than 10-3 n/m2, corresponding to a power of nbour 10'13 w/m2. Quite low
rh resholds are characCeristic of the perception of sounds with freq~en cies
ranging from approximately 20 to 100 kNz, and with a stronger difference
in the frequency o� the soun~+ from the aptimum threshola, they rise:
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rowf~rd the lower frequencies--more sln~vly, ~nd tiow~rd ~hc higher frequencies
--qutt~ ~h~rply, since aC a frequency of L50 IcElz in tl~e work menttoned the
Chreshnlds exceeded the rhr~shnld~ for ehe op~:tmum ~rc:quencies by al~nroxt.-
m~~t~~ ly G(1 db.
'Clie dlsadvant~ge nf this mcrhadology, howev~r, is i~s extremc laUor-
tntensiveness and the very long time taken for the Experiments, as coe11
as Che need for good Craitting u~ Che F~nim~l. This makes it difficult to
repent the experimenCs And reproduce Che dat~ obCained when many t~nimal~
~nre used. A11 Johnson's d1CA were obrained by using the ,.,~me dnlph~in.
5ubsequenC ,~nnlogous s~udies were alsn macte using n single anim~~.l: on a
botelenone dolphin, T. CruncaCus (Ayrupet'yanes et a1., 1^69; Morozov et a.l.,
1971), cn Che porpoise Ph. phocoena (Andersen, 1970), the whnle Orc~.nus oica
i.inna~~s (Ha11, Johnson, 1972) and on a fresh-water dolphin (,lacobs, Ha11,
197?). At Che snme ~ime, various work performed using dolphins of a single
species yielded noticeably varyin~ results. The limited naCure of the
experiment~l marerial made it impossible to decide wheCher these dlfferences
wer~ cau;;ed by individual characteristics of the animals being seudied o r by
nrher reasons.
rinre efficient is the methodology in which the thresholds of n~adiCary per-
ception in dolphins were also determined by the conditioned ref].ex merhod, .
not ~inder conditions oE free behavinr, but rather witli the develo~~menr. in
the secured animal af a defensive conditioned reflex to a combinal�fo;t of a
~ound si~;nal and electrodermal reinforcement. The vegetative components of
t`ie defensive canditioned reflex evoked were recorded (electrodermal reac-
ti~~n, ct~ange in respiration, pulse rate, etc.) and according to their
manifesCaCion iC was possible to ~udge wtiether the soun~ signal was p~rceived
h~~ the animal. (Supin, Sukhor:ichenku, 1970, 1974; 5ukhoruchenko, 19~1). The
r:,i:idity of the development o~ the defensive conditioned re�lex makes it
pos~ihle to obtain representative material, dupl.icatinR the measurements
when many animals ~,re used.
Me~7surements made in this way using rhe porpoises Ph. phoccena yiLla~:d
results ~imilar to the data obesined using bottlenose do.lphins. The mini-
mum thresholds of auditory perception were de~ected at frequencies of 64-
90-128 kNz and were l.ess ehan lOr3 n/mz (Fig. 36). The minimum threshold
v~~lues obtained i? individual measuremen~s using both dolphins T. truncatus
(.lohnson, 1967) and porpoises Ph. phocoena (Supin Sukhoruchenko, 1974)
reached 10-4 n/mZ, which corresponds to 10-14 w/m~.
Measurements were also made of. other characteristics af dolphi~~s' auditory
system, including the differential thresholds �.~ith respect ro frequency and
intensity. Ti~e measurements of the differential threshol~is for Lrequency
were made under the conditions of a behavio ral e:cperiment using trained
animals, T. truncatus (Jacobs, 1972; Thompson, Herman, 1975) and also under
the conditions of developing a deEensive conditioned reflex in porpoises
Ph. phocoena (Suictioruchenko, 1973; Supin, Sukhoruchenko, 1974).
8
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. ~ori orrtc;r.n~ i~sl; ocv~aY
~ Thc lowesr differential threshol.ds Eor frequency, obea~tned in the lutiter
work, were 0.1-0.2%, although in thc ent~.r~, quite broad ranqe ot frequ~n-
cies studied (frc,m 4 ro 180 kHz) they did nnt exceed 0.5% (F�lg, 37).
. . . ~
0,!
~ . ` Oi0
J20 ~
/60 0,01
-?0 - � s0
. �
40 1
10 O,QOJ
-40
/0
J
4 d /0 J2 Q4 /10 .~1 t1 4J !0 ~/p O,OD01
F~gure 36. Audiogram of a Porpoise Ph. phocoena, ObCained by the Method of
Recording Conditioned Reflex VegeCaCive Indicators (Supin,
Sukhoruchenko, 1974) '
Along axis of the abscissa--frequency of the tonal signal, in kHz; dlong
axis of ordinates--threshold intensity of the acoustic stimulus relative
to a level of 0.1 n/m2 db
Fi~ure 37. Differential Th resholds According to Frequency, Measured in the
Porpoise Ph. phocoena Using the Method of Reco rding Conditioned
Vegetative Indi~ar~Ls (Supin, Sukhoruchenko, 197G)
Along axis of abscissa--frequency of the tone; 1--threshold deviation
(scale on the left, in Hz); 2--relation of the threshold deviation to the
initial frequency (scale on Che right)
The measurements of the differential thresholds with respect to intensity,
made using freely muving dolphins T, truncatus (Burdin et al., 1971b) and
porpoises Ph. phacoena (our data) showed that the animals can perceive
changes i.n acoustic pressure of less than 10%. A sCudy was also made of
the ability of dolphins' auditory system to differentiate temporal intervals
(Thompson, Herman, 1975).
The necessary data on the nature of the functioning of the auditory system
of dolphins are not, however, exhausted by information of this type. A ,
study must be made of the mechanisms of ope.ration of the auditory system,
particularly the neural mechanisms of analyzing acoustic signals. The mai~
method thar may give the necessary information on this problem is the
electrophysiological method of study, which makes it possible to record
the activity of the sensory neural centers directly during the process of
their analyzing afferent signals.
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i~'nlt dH'l~'xCIAL, US~ 0?~LY
Tf~c fir~t ~~~~empts ~e ~n ~l~ctrophy~ioln~ir.~1 Heudy uf. the dnlphi?t~'
rauditvey gy~e~m de~lt wi~l~ the interior Col.lir.ul.u~ ~t che hr.~in (13ullnr.k
rt ~1., L9b8). 'The resultg d~ t:hig work ~re nnrticulprly dig~u~red in
mnre ~et~i1 1~~1~w. '~h~re ~rn ~~1go ~~t~m~~tH e~ ntudy Che eleCCrienl.
~ceiviey oi othr_r qer.Cion~ of the nuditory ~y~tc~m, includin~ th~ inn~Y
~~ir (McCormi.e.k, ~t ~1., 1~7U; Ridgw~y, ~t .yl., 1974).
Ii~ our expc~rim~nt~1 rese~r~~h, p~rtfc~l~r ~ttentic~n wri~ p;~id Cc~ �~tudying Che
,~?~~lyr~iq nf Ch~ acoustic si.~zn~1s in rhe nuditnry re~inn~ of th~ ddlphin~' `
c~tehral cnr[ex. ~xami.nhd uldng with rhib txrc c~rt~in re~ il~s obt~ined in
an ~~lectrdphysioldgi~~1 ~tudy of other center4 of the ~~:dienry syst~m--the _
corpus genieul~C~m medi~le ~tnd the inferior colliculu~.
1. MicrnelenCr.nde Invegti~nti~n of the Pulg~ Activity of Individual NeurdnK
of the Auditnry Cnrr~x
The meChod df micrdelectrode recordinR of the ~ceivity of individu~l neurnng
shnuld b~ regard~d ~s a mnre ndvr~n~ed meChod nf srudying in~ormaCinn ~,rnCes~-
ing tn th~e n~ur~l cenCerg a~. Ch~ prescnC time. Tt~is meChod makeg it possible
to p~rform ~ more detailed unnlysig nf the processeg takinK plac~e in th~ ~
neuron~ :,~d affords an oppozCunity of dire~tly obt~ining infarmaCi~n t}~aC is
inttccessible or accessibl.e with difficuley when Che niethod of recording tlih
summ~eed el~ctrical re~Ctinns is used. Most modern concepts of the
mechanlsms of sensory analysis couid be formulated only in consitl~r~tion of
the informatio?~ ob tairted with a study of the activity of individual neucons.
'Cherefore~ nCtempts were ~niei~iiy made .~C rnicroelectroi~r:. ~tudy of Ch~ work
of individual neurons in the dolphins' Cerebral c:ortex wuen an~lyzin~
:~coustic signals (Capich, Supin, 1974). These studies were performed under
chnditions of an acute experiment using porpoises Ph. phoc:~ena. In the
animt~lG operated on with a local anesthet~c, the actu;~ting part of a distant-
reading hydraulic micromaninul~tor was placed on the cranial bone, and by
me~ns of it tungsten microelectrodes witt~ ttie diameter af the tip in the
ordcr of 1 mic:rometer were inCroduced into the anim~l's brain. The main
d?fficulty in recn rding the acr.i~ity of Lndividual neurons in unr~nesthettxed
and nonir.~mobiLized dolphins is that the sharp movement3 ~cr.urring mainly in
bre.~thing may lead to d~mage or lo~s of [he neuron being reco~ded. With thc
use of remote-contrc?lled micromanipulator and with reliable attachment nn
the skull of its a~tuaCing part, however, this difficulty could be overr.ome
and stahle extracellular assigning of the pul.se activity of Che neurons
could be obtained.
~ One of Che main requirements fmposed on the work of studying the activity of
~ individual neurons is that a sufficien tly large sampling of neurons abso-
lutely must be presented, so that further statistical analysi.s can reveal
what forns of reactions of the neurons may be regarded as typical, what
variants of the reactions, types of n~urans, etc. exist. When the brain
of dolphins is studied. this requirement comes into conflict with the
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N'nK CJA'I~'LCI:AL Uy1~: dN1~Y
d~.~.tic~ulry UCC~HJ t~ th~ mat~rixl. 'Cliere~~rc, th~ ee~u1C~ of Che ~~ri~g
a~ exptrimen~g in whi.ch wc ~tudied the ~ctivity n~' individu~l n~uron~ nf ~h~
- dnlphln~' uuditnry ~dre~x mny b~ cc~n~idex~d only pr~limin~ry, ~inC~ efr~
limited nqCu~e of th~ experimenr~l m~terinl dn~H not p~rmit ~i~niFicnni.
C~nc~luHinna tn lie dr~wn cdnc~rnin~ thc~ sp~ci~.l. f~r~ture~ c~f ennly~l~ nf
~cc~ur~tir. gi~n~~.,~ in Ch~~ dolphin'~ uuditnry syqtem.
In ehi~ serle~ of ~xt~~riments (Cnpich, Supin, 1y74)~ 45 neurnng of th~ '
gudieory region of Ctie c~r~ebral cortex ~f Che pnrpoi~~ Ph. phoco~nn w~re
~tudied with pxtrnc~11u1~r r~ggigning di thelr pul~~ a~Civity. ~n addiCinn,
fnr comp~ri~nn, 31 neurong frnm the section df ehe corr.ex le~C~ezd outgid~
Che ~udieory regi~n (nneeridr eo it) w~re gtudied. Mogt of Ch~ neurong hnd
. ~ b~ck~r~und activity r~nging from 1 Co 10 pulgeg/sec. A Ce rCnin dif�er~nc~
w~s r.cvec~led in the medi.um frequenCy nf the b~ckground activiCy in the
neurnns of th~ nuditvey ~nd non~uditary r~ginn~. In the ~.uditory region,
Che frequen~y of the b~~kground aeeivity of the neurong wa.s mose ofC~n
1-b pulseg/sec., is ~hawn nn the hi~to~ram of di~tribuCion nf neuron~
nccbrding to the f requencies nf the beckground ~crivity in ~'igure 38, A
(median nf e;~e hi~togr~~n, 4.4 pulses/gec). tn rhe nonauditdry region of
the corCex, locgted eoward the ~nterior, ,eher~ proved tn b~ numerous neurong
with a hi.~her-frequency background r~ctivity (F'i~. 38, B); th~ mod~ of the
histogram is 6-9 pulseg/sec, ~nd of the medi~n--ln.9 pulse~/~ec.
SJith res~ect to the nature df eh~ distribution of the pulses in time, the
back~round activity could be irregular or grouped, with n pulae group
repetition periad of GO-].20 msec. The nnture of the background pulse
~ activity is shown in the fnrm of histograms of thc intervnls between Che
puls~es for different neurons. 2rregular activity gives g monomodal dis-
tribuCion of tntervals between ttie pul~cs, the peak of which corresponde
to the lengtt~ of the most frequently encountered intervals (Pig. 39, A).
The grouped activity gives a bimodal distribution of the inCervals beCween
the pulses (Fig. 39, B). The first peak of the histogram (0-20 msec in
Fi.g, 39~ B) corresponds to the length of the intervals within the groups
of pulses and the second peak (60-80 msec in Fig. 39, B) co rresponds to the
most probable length of thE intervals between tlie groups of pulses.
A cnnsiderabl.e number of the neurons of the ~7udiCory re~ion of the cortex
rear.t~d to acousti.c stimuli--clicks or turnfng on tonal ~ignalg. Out of
32 neurons of the auditory region, the reactions of whicli were studied with
the effect of Clicks, ]4 neurons reacted to this stimulus by activation,
3--by inhibiti.on, and fn 15 neuroas no reliable res~ction wes detected.
but of 2b neurons of the auditory region in which the reactions to tonal
si~nals were studied, reactions in the fnrm of activation were detected in
7 neurons, in 1--only an inhibiting reaction, and in 12 neurons no reliable
reactions could be detected.
- In [heir capacity to react to acoustic signals, the neurnns of the auditory
region of Che cortex differ substat~tially f rom the neurons of the adjacent
regions. in whict~ such :eactions were practically nonexi~tent. Out of
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}~n[t tl~~rCtAL US~ ONLY
73 nrurnn~ in Che nonguditnry regidn of th~ Cnreex~ eh~ ~ceivity of whteh
wgg ~eudi~d wiCh Ch~ pffect nf ~~au~tie seimuli, only in 1 n~uron wa~ a
r~~GCiun tio ~11 ACnl19C~C rli.~k rev~~l.ed.
y. ~ A J 6
y.
J9 40 .
,
10 J JB /0
t0 ~
!0 f :
/0 ~
,
, ,
,
,
0 J1 1~ J6 O /t ,t6 J/
Figur~ 38. ~'requency Distriburion of Beckground Pu1ge Aceivity of Neurons
in the Cerebr~l Cortex of the Porpoise Ph. phocoenn
A--sampling of neurons from the ~uditory region of tne cnrtex (gupragylvian
gyrus), 45 elemente; B--sampling of neurons from the nonauditory region of
the rortex, 31 elements. Along the a~tis of the abscissa--the frequency of
the background pulse aGtivity, in pulse/sec; along the axis of ordinates--
number of elements in each claes of histogram, absdlute ~nd in ~ of total -
number of elements in sample. Vertical dotted lines--medians of ehe
histogr~ms
In the auditory region af the dolphin's cerebral cortex a considerable
variety nf reactions to acoustic stimuli is detected. As an examplp,
Figure 40 shows, in the form of poststimulus histograms, the reactions of
several neurone to uniform acoustic stimuli--clicka. Neurons cgn be detected
with a short regction of the phase type. lasting 30-SO msec (A). Neurons
wrth ~n internu~diate phase-tonic fvrm of reacCion rpspond with an intensive
initial activation, which gradually falls in the course of 100-300 msec (B).
Neurons with a typically tonic type of reaGtion respond with a prolonged
- (for hundreds of msec), slowly falling excitation to even a short click (C).
~in~lly, neurons were encountered which responded to stimulus with a pause
of silence lasting 200-300 msec, after which a burst of activit�? followed--
postinhibitory activation (D). Therefore, in the audita ry cortex of the
dolphin'~ brr~in there is apparently a considerable v~riety of functional
types of neurons t+hich, as may be assumed, fulfill different functions in
the processes of sensory analysis.
A study of the reactions of the cortical neurons to tonal transmissions
showed that the reactions of the neurons depend on the frequency of the
tonal signals used~ and one may single out an optimum (characteri~tic)
frequency for the given neuron, at which the neuron's response is maximal.
~
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~n~ n~~ricrat, us~ ntvvY
At~ ~~n r.xnmpl~ w~ m.~y ~xnminc ehp re~ctinn~ nf a n~uron, pr~~~nt~d ih th~
f~jr~n riC phgC~timulr~rinn i~tgtdgr~ma in l~i~urc 41. 'Che reac:tiion~ ~ride with
t~~r ctf~ct of Cdnnl si~n~~~ wieh frcquencihp of. ~0, 40 and 60 kHz. tt rnn
be ~~~n tih~e in the given n~ur~n rh~ r~~cCi~n to a ton~ o� 40 kHx ~ubetnnei-
, 811y ~x~~ed~ th~ r~nceinn eo tdn~g af rhp ~d3e~~nr frequ~ucies. Un�orCun-
~eely, th~ limit~d natt~re df Che experimentai material did not p~rmie u~
to m~k~ ~ dernil~d ~Cudy ~nd compariedn of rh~ ch~ract~ri~tic Curvee
(re~~tion of the r~ncrion ed the fr~quency of ehe tone) of diff~rent neurong.
JO ~ 6
40 '
~
J9 %
ra ;j
%
~
~u ~ ~ ~ ~
~
f~
ii � , ii~
:
f
0 J00 100 D !00 t~'!
F`igure 39. Histogrnms of Interval~ Between Pulses for Various Type$ of
Background Pul~e Activity of Neurons of the Auditory Cortex
of the 9rain of a Po rpoig~ Ph. phocoena
~,---irregular artivi[y; B--Grouped activity.
Along the axis of ehe abscissa-~length of the inte rvals between pulses,
in msec; along the axis of ordinates--number of intervals
The d~t~ pregented, obtained in a microelectrode study of the activity of
individual neurons of the dolphin's auditory cartex, as has already be~n
emphasixed~ we regard ~~s preliminary. Unquestionahly the study of the
activity of individual neurons raay give quite important and interesting
information on the charncteristics of sensory cmalysig ia the dolphin's
auditory system ns well. For this, however, such regearch must be
developed on a considerably broader scal~, which may be regarded as a
ta~k for future works.
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~f~Et O~~ICIAL U5~ ONLY i
_ s...,_. _ ___~.~r .~,_s-
J00 � A 6~g~ S~C~ ~Cn~
t9
,
.f0
/0 , - .
' ~~~r~~ ' /~y,~j,
j
i /:~''~�'g'
~ t00 600 200 , ~.?Ol~ lOD ~ t00 i00
Figure 40. Typ~g of R~action~ af N~urnns of the Auditory Cortex of the
Br~in o� th~ porpoiea Ph. phocd~na co Acou~ric Clicks
Pogtseimulgtion hi~tngrams of varinue neurons. ~ach higrogram is obCained
by gveraging 20 r~a~tion~ ~f ~he neuron. A--phas~ r~oction; B--phase-
tonic reaCtion; C--tonic r~gcrion; D--inhibiCing r~actinn with postinhibit- ;
ing output. Along th~ axig of th~ abecisg~~-tim~, in mgec; along the gxig
of ordinat~~--frequency of pulge acrivity of n~uran~, in puls~/sec. The
arrowg indicate the momenCs of stimuli
~
6 (B) e (C)
~
29
! % ~
~;;i! ~f~ ,
,
% i~i'i~ i ~ /~~1
0 t00 ~00 0 t00 ~00 0 100 4~
Figure 41. Renctions of Neuron of the Auditory Cortex of the Brain of thc
Parpoige Ph. phocoena to Ton~l Acoustic Signals
PostaCimulation histograms obtained by averaging 20 renctions. Ainng the
axis rf the ebecisg~--the time, in msec; along the axis of orc'inetes--
f requency of ~ulse activity, in pulse/sec. iiorizc,ntal liaes under the
histogram~ denote the tiaie of effect o� the acoustic stimuli. Norizontal
dotted lines--average level of background pulse activity of the neuron. ;
Prequency of the tone: A--30 kHz, B--40 kHz, C--60 kHz.
~
~
i
~ '
14 j
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~
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~
~dit dt~'~t~IAI, USl: nNLY
i
~ `L. Conditi~n~ f~r th~ Invcseigbtidn oE Lvnk~d por~ntial~ nE Ch~ AudiCOry
Sy~t~m
~ ~
; Sub~tc~ntial gnd important fnformation on th~ m~~hen~gmg nt ~~n~ary ~n~1y~i~
may also be obeain~d by m~ac~~ of nn~ther ~1FIJ~~C in@thdf~~ wh~eh 3~ mor~
npplirabl~ wh~n ~h~ numb~r nf gnimnl~ u~~d fdr th~ ~eudy i~ limited, `
; occurg in workin$ wirh dolphin~. 'Chi~ i~ the m~Chod of r~Cdrdin~ ehe ~um-
mat~d bioel~rtric r~g~riong tn gffer~ne ~tiimuli--evok~d p~tenri~ls. Ae Ch~
~ ~~rly ge~ge~ of invegtig~~ion the u~~ nf ehig m~thod prnv~d unquestionably
~ mn r~ us~ful and gc~ve a lnrg~ amoune nE inform~tidn worrhy nE attention on
the phygiological m~chani~mg of pror~eging gensory ~ign~lg in the ~uditory
~ rorCe x of thp dolphing' brain (Pop~v, Supin, 1975, 1976 b)~
~
Th~ main p~rt of the ~xp~rim~nt~ w~~ performed u~in~ be~ttlenoge dolphing
trunCaCus.
Le~dg were mnde fr~m v~riou~ ~ection~ of th~ audie~ry cnrtex. By pre-
liminery probing w~ found th~ paint of the cortex ~t which evoked pot~ntial~
with considerable amplitud~ w~re recarded using acougtic stimulu~. ~rom
this point we then m~de g chrnnic lead of the biapotentigls for ~ long
tim~, recording the rcnctinns to ?:iie acoustic srimuli with sysCemnticAlly
varied par~meters. A considerable amount of informgtidn was obtain~d f rom
eech nf :~he point~ of contact by this mechod.
~ Upon complnrion nf the qeries of experimenta, a cnagul~tiom m~rker was put
in plgce with Ch~ ~id of g diathermic Co~gul~Cor. Mor~hological moniroring
: nf. the lorelization of the points of cont~ct was subsequently made accord-
~ ing to tt~~ge markers. .
At the time of the experiment, which lgsted for several hours, the animal
was placed in a tank measuring 3 X 0.5 X 0.4 meters, filled With sea water.
Afte c tl~e experiment w~s completed the animal was released into a pen,
where it wns kepr constantly. ?'hese rncording sessions w~re repeaCed mnny
times (usually daily) over ~ long period of time (up to several weeks or
months) and made it possible to nbtain a considerable amount of information
from each of the impl~nted C LECCLOdC3.
The aroustic stimuli were fed th rough pie~oceramic transmitters f mm 1 to
~ 6 cm in dinmete~, immersed in the w~ter with which the experimental Cank
hAd been filled. The gignals actueting th~ transmitters (tones, noises,
clicks, etc.) were monitared directly s~t their in~?ut.
The animal's mabflity in the tank was essentially limited. This made it
possible, with tt?e aid of a hydrophone, to measure the actual character-
istics of th~ acoustir signr~l (fnrm, intensity, etc.) inanediately near the
animal'~ hend, Which contributed to increasing the accuracy of the
experiments.
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~Ott U~~LCIAL US~ C~NLY
~ t
~
V A ' ~It -
1.. . ~..1~
~ /ODNRB ~f~
tO~ctK ~~stc)
Figure 42. Re~ponse~ of the Auditory Cortex of th~ Brain of a Bottlenoae
Dolphin t~ Aedustic Stimuli (Clicke)
1--example~ of individugl regponaeg; 2--r~suYt of averaging SO responaee.
The arrows (AL and A:) ~how the mea~uring of the amplitude of the firaC
pha~e and the overal~ amplitude of the responee
The evok~d poeentials recorded were averaged (from 50-100 reactions) using
the method of synchronous accumulation with the aid of an automatic elec-
cronic analyzer. The measurementa of the ampl.itude ~nd temporal chorac-
teristics of rhe evoked pntentials were made according to these averaged
evoked potentials (~ig. 42), and the numericgl data thus obtained were used
to plot graphs reflecting the relation of Che p~rameters of the ~voked
potentinl to the churacteristics of the stimulug. In a number of cases
an additional averaging proc~dure wag curried o~t: th~ graph, pasging
through a series of points, was approximated by a straight line. In these
cas~s the position of the approximating stYaight line closest to the
experimental p~ints ~aas found from equations of the lines of regression,
i.e., if the approximating straight line is described by the equation
then
ysQ~Fb2,
� (s - rl (u Gl
b ~
and a~y~6=~
Where x, y sr~ the ~bscissa and ordinate of each point, and x and ~
the averages of all the values of x and ,y respecri~?ely. ~
~
1
;
~
~
.
~
;
1.6 ~
9
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t
,
~
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r0it nF~ICIAL US~: OivLY
~vnktd pneenei~lg ~t tt~e St~re ~nd Seop ~f ehc~ Sdund~ ~orce Curve~
~~nd Abyalute ~hr~shnldg nf 1:xcitaeian of ~voked i'ot~ntinls
(?n~ nE the m~in ~h~?r~cterisCics of ~ny nnalyzQr caasises r~.E ir~ nbsolue~
rhrr~hn.ldq nf perc~ption and forc:e ch~r~r.Ccri~Ctc~--thc relncion af the
r~nc~Cion ~a the int~ngiCy nP the.~ qtlmul.ii~, 1'h~reL�ore, thE~ fir~t. arage in
elic tl~crrophyginingicul seudy of the i~uditnry compl~:c nf a dotphin' d
brain w~~ ~ls~ Cl~~ d~eermin~~ion nf th~ :~bsnlute ~hreshnlda and forc:~
characterigties of th~ ~vok~d potenei~ls wirh different condieion~ of
srimulug. For th~s purpose ehe r~gponses aE Ch~ auditory reginn of the
dolphin'g cerebr~l cortex w~re recorded ~~t rh~e sr~~rt or stop of var3oue
gounds (noises, pure eones), As w~l.l ~s eo acoustic clicky, by varying
the inrensiCy of th~ stimuli (Popov, Supin, 1976a),
~ ~igure 4~ ehow~ rhe ren~ridns di the uuditn ry r~gion nf eh~ c~rebral cnrtex
� to ncnuaric clicks. As cttn be seen from ehe dingr~m, the t~mpornl chnrac-
CerisCic:a nnd form nf the r~spnnse~ ch~7nge liCtle with a v~ri~tion in ehe
intensiey of th~ stimulus; the aunplitude of rh~ respnnses is most obvinusly
changed. Therefore, to nbtain the force cu.~ves, ~h~ ~mplieude of the
re~pons~s was me~sur~d aC vnrious intensil.ies of the ~timulus.
The ~mplitudes of vnrious components of t1,t~s response were meg~ured. The
basic une~ were~the measurement of thp ;,,dgnieude of rhe firsC phase of the
response (from the isopc:tential line to the pcak) and measurement of the
overall ~mplitude of the ~~sponse (from Che penk of the Eirst eo the peak
of the s~cond phese), ns shown in Figure 43. The results of rhe experiments
showed, tiowever, Chat both these vdlue~ change in practtCally identical
mnnner with a change in the chardcC~risticg nf the stimulus; in p~rticular,
~ the force curves have an essentially identic~al form for both the amplitude
of the first ph~se of the response, and for the overall amplitude of the
response (Fig. 44). Therefore, in the future graphs were made that were
plotted only according to tlie changes in the amplirude o-f :he first phase
of the respons~s.
Figure 44 shows the relation of the amplitude of the response to the inten-
sity of the sound stimulus--a brief. bro~h~b00 1
B i
~
iQ
/O ~
so 's
/O~Z
i
i ,
~ � S f0 d6 !0 Z /O~~ /0~ ~
' Figure 46. Characteristics of Respanses of thG Auditory Cortex of a
Bottlenose Dolphin's Brain to a Change in Intensity of a~+
Tonal Signal
A--relation of the amplitude of the responses to ~he amount of increase in
the .intensity of the sound. Along [he axis of the abscissa--increase in
acoustic pressure, in db; along rhe axis of ordinates--amplitude of the
responses, in microvolts; 1--initial intensity of sound, 1 n/m2; 2--0.3 n/m2,
3--0.1 n/m2, 4--0.03 n/m2.; P--relation ot the difF~rential threshold
to initial intensity of the si~nal, Al.ang the a~is of the abscissa--the
initial intensity oE th e saund, in n/m2; along the vcis of ordinates--the
dif�erential threshold, in db; C--relation of the diff.erential threshold
to Che initial intensity of the saund. ;11ong the axis of the a'~scissa--
. the initial intensity o� the sound, in n/m2; 11ong the axis of ordinates--
. the threshold increase in a~nustic p ressure, in n/m2
The conformances to principle described are well illustrated by graphs that
show how the thresholds of responses to a change in intensity of the soun d
depend on the background intensity. Figure 46, B shows the threshold
values ~I/I at various background intensities_of sound. The thresholds were
23
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~,nx n~~icr~, us~: o~v~.Y
~ou~jd by c~xlr,~pdt.~~ti.un ni' th~ gr:~pha q11~wn in I~'i~urc 4C~, A. ~'igurc 46, li
Hlic~w;~ th~ r~dme dat~~ iu c~ dif.[erent form--rag th~+ r~l~~ti~~r~ ~t' thc thre~hdl.cf
r.}~anges in inten~ity ~I ec~ the b~~kgr~und inCnn~iCy nf r.he ~ound. IC enn
b~ e~~ily geen th~t with lnw ba~kgrnund inten~itir~~ (1!1-1.-~�10'2 n/m2), Ch~
ab~hlute thr~~hdid vr~lue~ of th~ rhung~ in inten~ity (Qr) remnin cnn~Cant--
eh~y r~r~cl~ v~lues cln~~ rn eh~ thr~shnld nf nrigin~rinn nf r~~pnn~~~ ro
eh~ ~t�~rt ~f ehe gound and dh nuti drop Lower. CJiCh hi~her int~neitie~
~d~~-3 n/m2), ehe relr~tiv~ e}~re~hold~ ( ~~/I) b~come c~n~eant, and nr~
~ytnbli~hed ~t ~ 1eve1 nf nbdut 0.6 cib.
T'~~ ~bnve-d~~~ribed experiments wer~ p~rform~d using purn tnne~ ng Ch~
- ~eimuli. An~lm~aus mer~gurc~ment~ w~re nbtcained fnr brn~d~,~nd noige~ The
chr~ng~~ in the int~nsity ehe noise result in ehe appearanc~ in the
rortex of r~nalogous evnked patenei~ls, rhus ch~nging, dep~nding nn th~
m~gnitude df eh~ iner~~s~ or d~cregg~ in the intiengity of the gound. 7'h~
threyhc~ld changes in ine~ngity fnr noig~, how~v~r~ proved tn be noticedbly
tiigt~er ~h~n ehe threghdld ~h~nges in t}ie inCen~iCy fnr the tone--30-50X
(2-3.5 db).
It is pn~sible, however, thet the higher Chre~hnlde o� ch~nge in inten~iey
�nr noise are c~used noC by nny specific reductinn in sen:~itivity tn thi~
type of ;~timuli, buC sim~ly Uy che fact ther in mtcrointhrvnls di tim~ Che
amplitude of th~ noise is not st~tion~ry. 'Thi~ lnck of st~bility in the
~mplitude may mask the moments of the chang~s~ in the averr~Ke intpn~ity of
the noi~e, if the~e ch~nges nre not gre~t. Th~refore, th~ thresh7lds~ of
rea~tions to a rh~ngc in the intengity nf the naise may prove ro be
raised.
5. Time Summatinn
A tiR,~~ summation is one o� the important characteristics of an auditory
syscem tl~et determf.nes the speci~l features of perception of sounds of vary-
:ng duration. Th is characteristic may also be studied using electricol
re~c[ions (evoked potenti~~ls) of *_he cerebral cortex.
The experiments were conducted as follows. The evnked potentials occurring
in response to the sound transmissions (noises, tnnes) of varying duration
were recorded. The rel~tion of the amplitude of the rexponses to the dura-
tior. of the transmission and intensity of the sound was studied.
The transmissions with a regul~ted transconductance of the fronr were formed
~ by an electronic key c~~ntrolled from a master oscillator. Th~ lengch of tt~e
sign~ls formed hy :ne electronic key could be regulated within a broad
range. Due to Che reverberatory properties vf thr tank in which the sCudiee
were made. however, the length of the sound transmissions actually oCCUrrine
in it could not be made shorter [han 1 msec. Therefore, the capacfty of
the dolphin's auditory system for time summation was studied in our experi-
ments for periods of. time of at l~ast 1 msec.
24
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~ ~tll~ ~~k'ti;tAL t1~l; r)Nf~Y
~ ~ _w_ _ e
~ rso _ . _ ~..w._.~..
,rv�~
~ ~ ~a~'t
10d . ~'~d 'r I -
~~in't _
n n J. ~
.~�rD ? 1d �
JD n r
�
~ 1 ,r
/ s~~~,~~'~ ~..Jp
~ --ZO
r r r
.1.... ' ' 1 . ~
~
~ ~ J /0 ID Id ~ JO "1 >d > ~ rv
~i~ur~ 47. ~!:fects of n Time Summ~tit~n in ~iie At.itlik~ry Gortex of tt~e
Biain o~' ~ 13~eel.cno~e Uolptif.n
A--reloCidn di the amplitude of Cl~e evoked n~c~~~~ iel;; to the dur~Cinn of
eh~ noige er~nsmission with varyin~ ir~tensit:ies uf Htimulus. Alnng th~
gxi~ nf the abscis~~~-=Che durx~Li.c~n of tt~~~ ~3r.imulu,~, in m~;~c; al~ng the oxi~
nf ort;inACet~--emp~itude n~ the re5pc~ttsrs, i.n mirt�~~volts. Shnwn for ench
grnph i.o the intensity of the nni_~e--tht~ ~ff~~etlve valu~~ of the tiCOUBC~C
pr~~gure, in n/m~; $--the s~~me data ~re presc~nt~c3 as the relaCion of the
~mplitud~ df Che ~cespt~nse (alon~ the axis of ordi~~ates, irt mfcrovdlts) to
Che inten~ity of the noise transmis~ton (f~lc~n~; rh~ axl~ of the abscigsa,
in n/m2) with varying dur~tinns of the t r~rismi.s~ion (in~ii~~t~ed for edch
greph in mg~c); C--relation of thc rhre~hc~lcir; of exclt;itian nf the regponseg
to the dur~tion of tlie noise tr.~n~mi~~~ic~~r ;~rcc~rdiny=, to the data of the seme
experiment. Along the nxis of the ab;~r_i:~~a--rttc~ dur~tion, in :nsec; elong
the axis of ordinatc~--the threshold i.n~ensicy of thcr noiye, in n/m2
~igure ~i7 shows the results of an ex;~r~r�r,~nt witli a study of the reactions
of the cnrtex to transmission5 of a t~~ne ot: varyiny, dur.~tidn. In the graphe
of the t'elation of the amplitude c~i' the c~vakcci potentials to the duration
o� the sound tran~mission (~ig. 47~ A}, {-1+~~~r incr~s~~:c~ c~rn be seen in the
tunplitude of [he respon,~es wfth ~~~1 tncr~~.jsc= iri th~~ dur.~tion of the trnns- _
mission from 1 to 20 m.~ec~ wliic}~ :,~.~y b~~ rc~f;ardr.d as a phenc~mc~non af [imp
summation. It mur~t be nnted, howevcr, r.hac C?:~ ohserved tncrease itt the
amplitude of the respt?nses i:~ rc3lativF~iy sr:::ll. i.e., in ehe ran$e of
durations Atudied (1-20 mse.~), t3~e t ir~c~ su~::~;at i~an 3.5 expreqsed relatively
weakly.
The m.7rked nature of tt~~ timc~ ~~ur.u~at Ic~n d~~p~nds on the intensity of the
sound. If one cortq~ares ct~r. relation ~~f ti~c ~~m,~litude c~f th~ respon~eg to
the di~ration of the ~signal wf tii varyfny; in;c~;i~it;c~s c~f �iih sound, it is
disr.~vered th:~t an i.ncreasr in thr~ > >
1971b, pp 1I2-115.
~
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7. V~si1'yev, A. G., "L'voked poCentigls and Che ~~acCions of Yndividu~l
Ncurons of the Inferior Colliculus of B~Cs eo U~.til't3SOlt~.C Stimu~.i With
~ DifferenC Duration of Their ~ronts," VCSTN. LCU. S~R. BId1,., Vnl 1,
No 3, 1968, pp 167-170.
~
8. Veyn, A. M., "Narusheniyh snA i bodrstvovaniya" [Sleep llisorder~ and
tJakefulness], Moscow, MediCsina, 1974.
9. Vel'min, V. A., dnd Titov, A. A., "Audirory Uiscrimin~tion o� Tntervals
IieCcoeen PuYses in BotClenose bolphins, " Morskiye m1..,copitayushchiye.
rtaterinly VI Vsesoyuz. soveshch." [Marine Mammals. Materials of the ~
+ Seveneh All-Union Conference], Vo1 1, Kiev, 1975, pp 77-78.
10. Gapich, L. I., and Supin, A. Ya., "Ac~iviry of Individual Neurons of
the AcousCic Area of the Dolphin's Cerebral CorCex," zHUR. EVOLYUTS.
BIOKHIM. I FIZIOL., No 10, 1974, pp 202-203.
11. Gapich, L. I.; Ladygtna, T. F.; Supin, A. Ya.; and Sukhoruchenko, M. N.,
"The Technique of Llectrophysiological Research on the Brain of the .
Dolphin," in the collection, "Morfologiya i ekolo~iya motiskikh
mlekopitayushchikh," rtoscow, Nauka, 1971, pp 95-105.
12. Gusel'nikov, V. I., and Supin, A. Ya., "Ritmicheskaya akti~rnost'
golovnogo mozga" [Rhythmic Activity of Che Cerebral Cortex], Moscow,
Izd-vo MGU, 1968.
13. Dubrovskiy, N. A., and ICrasnev; P. 5., "Discrimination by the Bottle-
nose Dolphin ~~tween Spheres With Respect to Material and Dimensions,"
TRUDY AKUSTICHESKOGO IN-TA [Works of the Institute o� Acoustics], ;
Vol ~17, 1971, pp 9-18.
14. D~ibrovskiy, N. A.,; Ti~:nv, A. A.; and Krasnov, P. S., "A Study of the
Resolving Power of the Ecliolocation Apparatus of a Black Sea Bottle-
nose Dalphin," TRUDY AKUSTICHESKOGO IN-TA, Vol a~J, 1970, pp 163-181.
15. Zaboyeva, N. V., "Time Summation in the Auditory Cortex With Exposure
to Sounds of Varying Frequencies and White Noise," in the collection,
"Mekhanizmy sl.ukha" [Hearing ;fechanisms], Leningrad, Nauka, 1967,
pp 90-101.
16. 'Lvorykin, V. P., "Morphological Bases of Ultrasonic and Locational
Pro~erties in the Dolphin," ARKH. ANAT., GISTOL. I EMBRIOL., Vol 45,
No 7, 1963, pp 3-17.
17. Kesarev, V. S., "Structural Organization of the Limbic Cortex of the
Dolphin," ARKH. ~~iAT., GI~TOL. I EMBRIOL., Vol 56, No 6, 1969,
pp 28-35.
86
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18. Kes~rev, V. S., "bnea on Che Neuron OrganizaCion'of Che NeocorCex nx
the tiolphin's Iir~in," AItKH. ANA'I'. , GZS'fOL, Y CMBRIOL. , Vnl 59, No 8,
1y70, ri> >i-7~.
19. Kes~rev, V. S., "'1'he Problem nf Homologiz~Cion of Che Nencortex of
C.he Brain nf Cetaceans," AItKH. ANAT. , G'~STOL. I CMBRIOL. , VoJ. 68,
No 3, 1975, pp 5-13.
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htotor Are~ of the Cerebral Cor�rex of the Dolphin," ARICH. ANAT. ,
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21. Ladygina, T. F., "Characteristics of ~voked Pntentials ot Che Cerebr~l .
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rIZioL., voi 10, 1974, pp 66-71.
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23. Ladyoina, T. F., and Supin, A. Ya., "Evolution of the CorCical Areas
of the Brain of Land and Aquatic Mammals," in the col.lection,
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[Morphology, Physiology and Acoustics of Marine Mammals], Moscow,
Nauka, 1974, pp 6-15.
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Cerebral Cortex in Bottlenose Dolphins," "Morskiye mlekopitayushchiye.
Materialy VI Vsesoyuz. soveshch.," Vol 1, Kiev, 1975, pp 172-173.
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~ collection, "Morskiye mlekopitayushchiye. Rezul'taty i metody
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' Moscow, Nauka, 1978.
26. Maruseva, A. M., "The Dynamics of Inhibiting Influences in the Auditory
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27. hiorozov, V. P.; Akopian, A. I.; Burdin, V. I.; Donskov, A. A.;
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Tursiops truncatus," FIZIOL. ZHURN. SSSR, Vol 57, 1971, pp 843-849.
28. Mukhamet~v, L. 'r(., and Supin, A. Ya., "Physiological Characteristics
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mlekopitayushchiye. Materialy VI Vsesoyuzn, soveshch.," Vol 2, Kiev,
1975~ pp 21-23.
87
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'L9. Plulchrime~~v~ L. M.. ~~nd Sup~tt~, Y,y., "L~C-SCudy of
V~riouq BChcivi.car,h]
St~?r~~ c~f I~'r~ely Movl~i~ Uolphin~," 'J.IiVNp, Vol 'l5, 1~)75, pp 3$6-40L.
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Itezul'taCy i meCody issledov~niy," Mo~cnw, N~ukfl, 1.978.
31,. Mukh~metov, L. hi.; Supin, A. Ya.; and 5trnkov~, I. G., "Inrerhemispheric
Asymme~ry of Che Functional StaCes oE the Iirnin During S1eep in Che
- DolE~hin," bOKL. AN 55SR" (Works of ehe USSR Academy of Sciences~,
Vo1 229, 1976, pp 767-770.
, ~
32. PcCukhov, V. V., and KoCelev, V., "CommuCaCional Device for Record-
ing the Electrical Activity of the I3rain o� an Animal Under. Conditians
of Free Movemenr," ZhVND, Vol 25, 1975, pp 1324-1326.
33. Ptlleri, G., and Gir, M., "The Nervous System," in the book:
A. V. Y ablokov; V. M. Be1'kovich; and V. I. I3orisov, "Kiry i del'Einy"
~ [~dhales and Dolphins], Moscow, Nauka, 2~i2, pp 200-232.
34. Popov, V. V., and Supin, A. Ya., "Determinins the Char~cteristics of
Nearing in the Dolphin Using the htethod of Evoked Potentials,"
"Morskiye mlekopitayushchiye. P-:aterialy VI Vsesoyuz. soveshch.,"
Vol 2, Kiev, 1975, pp 64-65.
35. Popov, V. V., and Supin, A. Ya., "De~ermining the Characreristics of
Hearing in the Dolphin Usir.~g the Method of Evoked Potentials,"
FIZIOL. ZHURN. SSSR, Vol (~2, 1976a, pp.550-558.
3G. Yopov, V. V., and Supin, A. Ya., "Reactions of the Auditory Cortex of
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Gol 62, 19766, pp 1780-1785.
37. Popov, V. V., and Supin, A. Ya., "Electrophysiologf.c~l Investigation
of the Auditory System of the Dolphin Tursiops truncatus," in the
collection, "Nlorskiye mlekopitayushchiye. Rezul'taL-y i metody
issledovaniy," rioscow, Nauka, 1978.
38. Rizhinashvili, R. S., and Mosidze, V. M., "The Significance of the
Commissural System of the Brain in the LateralizaCion of Sleep and
Wakef.ulness," SOOBSHCH. AN GSSR [Communications of the Georgian SSR
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39. Roytbak, A. I., "Bioelektricheskiye yavleniya v kore bol'shikh
polushariy" [Bioelectric Phenomena in thP Cerebral Cortex], Tbilisi,
Metsniyere6a, 1955.
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~ 40. RoyCb~k, A. I., "Cvolted PoCeneinls of ehe Cerebr~l CorCex," in the
. coll~ceion, "Sovremennyye problemy clekrrotiziologir.heykikh
~ :L~sledov~niy nervnoy s~.stemy" CCurxent Problems nf C1~cCrophysio-
logic~l Studies of Che Nervous Sysrem], Moscow, Medirsin~, 196G,
~ n~~ i~4-219.
41. Solcolov, V'. Ye.; Ladygina, T. F.; and Supin, A. Ya., "Localizarion c~f
Che Sensory Areas in Che Cerebral CorCe x of a Uolphin," DOKL. AN SSSit,
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' Mechanisms of Vieual Analysis], Moscow, Nauka, 19 74.
, 43. Supin, A. Ya.; Bogdanova, L. N.; Slezin, V. B.; Fedenko, M.. S.; and
Matisheva, S. K., "A MeChod of Impl~nting Chronic Electrodes in ehe
I3rain and Postoperative Treatment of Dolphins," "riorskiye _
mlekopitayushchiye. Materialy VI Vsesoyuz. soveshch.," Vol 2, Kiev,
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44. 5upin, A. Ya.; Bogdanova, L. N.; and redenko, M. S., "An Experiment in
~ Using Certain Narcotic and Immobilizing Subseances on Black Sen
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, Thresholds of Dolphins Using the Method of the Conditioned Cutaneo- '
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~ Moscow, Nauka, 1974, pp 127-135.
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the Seas of the USSR], Pioscoca, Izd-vo AN SSSR, 1962.
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8g �
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51. Shlcol' nik-Yarros, Y~. G. ,"N~yrony i mezhneyronnyye svyazi. '
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V~.sua1 Analyzer], Moscow, Meditsin~, 19b5.
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Cchidna, T~lchiglossus aculcatus. III. Sensory and tnterhemispheric
~voked Responses," AItCH. ITAL. BIOL., Vol 110, 1972, pp 195-216.
54. Andersen, S., "Auditory Sens3tiviCv of the Harbor Porpoise Phocoenn
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I ;

~

i ~ ~

~ . ~
PHYSIOLO~3ICAL INVESTIaATION OF THE DOLPNIN 6RAIN
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CO~YltIGNT: Izdatel'stvo "Nauka", 1978
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