Morgane 1980

The Anatomy of the Brain of the Bottlenose
Dolphin (Tursiops trunca . Surface tus)
Configurations of the Telencephalon of the

Bottlenose Dolphin with Comparative
Anatomical Observations in Four Other
Cetacean Species’
PETER J. MORGANE, MYRON S. JACOBS AND WILLARD L. MCFARLAND
Worcester Foundation for Experimental Biology, Shrewsbury, MA,

New York University, New York, NY
and
National Institutes of Health, Bethesda, MD
Received 30 November 1979
MGRGANE, P. J., M. S. JACOBS AND W. L. MCFARLAND. The annfomy of the bruin of the boftknose dolphin
(Tursiops truncatus). Surface configurations of the telencephalon of the bottlenose dolphin with comparafive anatomical
observations in four other cetacean species. BRAIN RES. BULL. 5: S~ppl. 3, l-108, 1980.-By analyzing the surface
configurations of the telencephalon of the bottlenose dolphin and four other cetacean species, we have been able to
delineate the major sulcal and gyral patterns, define the basic lobular organization of the highly gyrencephalic cetacean
brain and describe the general organization of the brain surface into major cortical territories. Although the cetacean
endbrain presents many general mammalian features, the surface configurations of the brains of these aquatic mammals
differ in many significant details from the patterns of other mammals. These departures are subtle in the rhinencephalic
(median) zone of the telencephalon, are more prominent in the limbic (paramedian) zone and become outspoken in the
dorso-lateral and posterior zones of the fundus of the hemispheres. The latter zones of the fundus display the widest
interspecies variability as well as the most marked individual intraspecies variability among Cetacea. The more salient
general features we have observed are: (1) pronounced antero-posterior foreshortening of the forebrain with enhanced
dorsal proplasia of the parietal lobes, blunting of the occipital lobes, and ventro-lateral proplasia of the temporal lobes; (2)
great transverse width of the dorso-lateral and posterior (epihippocampal) parts of the hemispheres and reduced width of
the antero-ventral (epistriatal) parts; (3) conspicuously small hippocampus contrasting with the great mass of the epihip-
pocampal parts of the hemispheres; (4) large olfactory lobes and massive septal area contrasting with small orbital lobes and
striatum; (5) relatively large insula, buried under a massive operculum, the latter disposed as a discrete lobule demarcated
from the dorso-lateral wall of the hemisphere by a deep arcuate fissure; (6) presence of a paralimbic lobe, unique in
Cetacea, intercalated on the medial wall of the hemisphere between limbic and supralimbic lobes and covered by a special
heterotypical cortex of the sensorimotor type-agranular motor anteriorly, granular somesthetic dorsally, and modified
koniogranulose visual posteriorly; (7) some degree of cytoarchitectonic homogeneity though areal differences can be
resolved and some apparent homogeneity may be due to the gradual transition of one area into the other which is
characteristic of the cetacean cortical areas; and (8) unusual organization of the homotypical cortex of “associational” type.
However, a resolution of this latter must wait thorough architectonic evaluation of the entire supralimbic lobe. The surface
demarcation of the rhinencephalon and the cleavage of each hemisphere into concentric tiers (limbic, paralimbic and
supralimbic) are sharper and clearer in the cetacean brain than in any other mammalian brain. The segregation of the special
heterotypical cortices in the paralimbic lobe, cleaved from the medial hemispheric wall by arcuate longitudinal clefts, is, as
far as we have been able to determine, unique for Cetacea. In contrast to primates, in which the somatic sensorimotor and
special sensory cortices are widely separated by broad expanses of homotypical cortex (association cortex), these inti-
mately thalamus-bound special heterotypical cortical areas in Cetacea remain largely on the dorsomedial wall of the
hemisphere bordering on the epihippocampal limbus, and do not appear to extend onto the supralimbic fundus of the
hemisphere. This uniquely cetacean feature characterizes one of the most interesting aspects of the evolution of the brain in
these totally aquatic mammals.
Dolphin brain Whale brain Cetacea Tursiops truncatus Iniu geoffrensis Balaenoptera physalus
Cetacean brain Delphinapterus leucas Phocaena phocaena Supralimbic lobe Paralimbic lobe
Limbic lobe Rhinencephalon Comparative neuroanatomy Olfactory lobe Cytoarchitectonics
Cortex cytoarchitecture Isocortex Neocortex Paleocortex Archicortex Aquatic mammals
Telencephalon Association cortex Homotypical isocortex Heterotypical isocortex
Sulci and gyri of whale brain Insula Limbic cleft Paralimbic cleft
‘Supported by NSF grant BNS 77-08660.

3
MORGANE. JACOBS AND MCFARLAND
THE Delphinidae are the family of Cetacea, including dol- tions of each hemisphere were identitied and photographed.
phins and porpoises, that are most accessible to observation The serial sections, in addition to defining architectonic
and to experimental inquiry into the mode of their compara- tields, provided a very important guide for description of the
tive anatomical and behavioral organization. In the past few surface morphology of the forebrain since areal differentia-
decades there has occurred a notable increase of interest in tions could be determined and architectonic boundaries
the study of behavior and the nervous system of these a- more clearly defined at high magnification. Our observations
quatic mammals. In this country the studies of Cetacea have of the surface configurations of the brains of Tur.siop.\ and of
been led mainly be ecologists and ethologists rather than by the four other cetacean species were compared with those in
anatomists and physiologists. The present study is one of a the earlier descriptions of Beauregard (1883). Guldberg
series we have been carrying out on the brain of the bottle- ( 1885), Kiikenthal and Ziehen (1889), Bianchi (1905). Pettit
nose dolphin (Tursiops truncatus) and other cetacean species (1905), Rawitz (1910), and also in the more recent studies ot
which, in addition to its comparative neuroanatomical inter- Kojima (1951). Friant (1958. 1974). Pilleri (1964. 1966). and
est, may also serve as an anatomical reference for physio- Hammelbo (1972).
logical studies on the brain of this species.
The present paper emphasizes the surface morphology of
the telencephalon of the bottlenose dolphin and of four other I. GENERAL SURFACE CONFIGURATIONS UF THE FOREBRAIN
cetacean species used for comparison. Our chief concern is AND ORIENTATION OF THE CEREBRAL HEMISPHERES WITH
REFERENCE TO THE “BEAK-FLUKE” AXIS
to describe the common basic pattern of the surface config-
uration of the hemispheres of Tursiops trunccrtus which The nomenclature of sulci and gyri of the brain of Cetacea
would then serve as the topographic reference for later cyto- in general usage has been that used mainly for the brains of
architectonic studies. ungulates and carnivores [2,4, 25, 32, 33, 34,48,55]. Never-
theless, the comparative anatomical, physiological, behav-
MATERIALS AND METHODS
ioral, and ecological studies of the bottlenose dolphin indi-
cate that this highly specialized, exclusively aquatic mam-
The material for these investigations consists of six mal. exhibits cerebral morphological peculiarities of its own
cerebra of adult bottlenose dolphin (Tursiops truncatus~ and which not only set it apart from land-living and from other
four cerebra of other Cetacea (beluga whale, Delphinapterus para- and semi-aquatic mammals, but also, in some features.
ieuws; common or harbor porpoise, Phocaena phocaena; distinguish it even from other Cetacea [ 19,24,40,44.45,49].
Amazon River dolphin, Inia geoffrensis; and the finback We have been mainly interested in the differences which the
whale, Balaenoprera phpsalus). For the gross comparative brain of the dolphin exhibits rather than its many obvious
anatomical aspects of this study, the cerebra of five Tursiops stimilarities with the brains of other mammals. This orienta-
and one brain each of the four other cetacean species were tion has made it desirable to use, as far as possible, a de-
dissected and compared to the brains of ungulates, cami- scriptive nomenclature which would not force, by purely
vores, primates, and man. Three other cerebra of Tursiops semantic implication, unwarranted homologies upon certain
were prepared in whole brain serial section and used for special formations of the brain of Cetacea in general and of
histo-anatomical analysis. These latter cerebra were gravity the bottlenose dolphin in particular.
perfused in situ with Windle’s fluid, removed on a vertical Among the characteristic features of the brain of the dol-
milling machine, photographed, weighed, and measured. phin are those pertaining to the position of the forebrain irl
The three brains weighed, respectively, 1408, 1632 and 1569 situ with reference to the longitudinal body axis (beak-fluke
g and had volumes of 1315, 1600 and 1475 cc as determined axis), and those pertaining to the surface configurations of
by the water displacement method. They were then dehy- the cerebral hemispheres. The first of these features makes
drated in graded alcohols, embedded whole in celloidin, and the conventional reference to planes of section of the fore-
cut in a gapless series of sections at 35 micra on the Goddard brain in general usage for the brains of other mammals often
[22] brain microtome (Mica Instrument Co., Cambridge, ambiguous when applied to the dolphin. while the second
MA). Each of these brains was cut in one of three cardinal feature raises problems of the identity of parts of the hemi-
planes: the sag&al; the frontal, which was at approximately spheres in cetacean brains with parts in apparently homolo-
a right angle to the beak-fluke axis; and in a horizontal plane. gous positions in the brains of other mammals. Here. in the
which was parallel to the beak-fluke axis and at a right angle surface configurations of the neopallial formations. the
to the frontal plane. Every fifth section was stained by the homologies based on mere positional similarities existing be-
Loyez modification of Weigert’s hematoxylin method for tween the dolphin and other mammalian brains become often
myelin sheaths 131, and each adjoining section throughout tenuous and uncertain.
the series was stained for nerve cells and glial nuclei by the As compared with the brains of land-living mammals, the
Bielschowsky-Plien cresyl violet method 131. The serial sec- peculiarities of the dolphin brain are most obvious when vis-
tions were studied on the Galileo contour measuring projec- ualized in its natural orientation in the cranial cavity (Fig. I).
tor and Nikon optical comparator and with higher power In the dolphin the forebrain is foreshortened (“telescoped.”
microscopy in close conjunction with regional dissections of Kojima. 1951) in the rostro-caudal (beak-fluke) or body axis
the formalin fixed whole brain specimens. of the animal. It exhibits pronounced mesencephalic, pon-
The gross dissestion of brains of all five cetacean species tine, and cervical flexures (Fig. 2A). These flexures in the
used in this study were carried out as follows: After each brain of man (Fig. 2B) and of most land-living mammals are
intact brain was photographed, it was cut in midsagittal plane prominent only during the embryonic state (Fig. 2C). In a
through the interhemispheric cleft, septum, and raphe of the lateral view of the cerebral hemispheres the antero-posteriol
brainstem to expose the medial walls of the hemispheres and foreshortening of the forebrain of the bottlenose dolphin
third and fourth ventricles. Then. the opercula were re- (Fig. 3) and of other cetacean species (Figs. 4. 5, 6. 7) is
moved in stages to expose the insula. At consecutive steps of evident when compared to the generally elongated brains ol
the dissection the cardinal furrows and major lobar forma- primates (Fig. 8A. C, D) and carnivores (Fig. 8B). In con-
SURFACE ANATOMY OF WHALE BRAIN
splenium of corpus callosum
P
‘CERVICAL
FLEXURE
PONTINE
FLEXURE
rostrum of corpus callosum Chop

MtiSENCEPfjALIC FLEXURE post Hyp
I
Ant Hyp
FIG.?A. Photograph of a mid-sagittal brain section of the adult dolphin brain indicatmg brainstem flexure pattein. The brainstem exhibits
right angle turns at the pontine and mesencephaiic ftexures. The spinal cord twrns directly at an approximately 90 adgle from the cervical
flexure in the body (beak-fhike) axis (See Fig. 1). Note the extreme anterior rotation of the hemisphere on the upper brainstem so that the
corpus callosum has an almost vertical otientation. Abbreviations: Ch @+-optic chiasm: 3 V-third bentricle; Ant Hyp-anterior hypothal-
amus: Post Hy-posterior hypothalamus. (approx. II/:x).
SURFACE ANATOMY OF WHALE BRAIN 5
splenium
roe1 of -of corpus
corpus orum’ collosum
pontins
- flexure
BOD; AXIS / ah& I m&encephalic flsxure

3V INFUNDIBULUM
FIG. 2B. Photograph of a mid-sagittal section of the adult human brain indicating brainstem flexure patterns. The stem below the mesence-
phalic flexure is approximately at right angles to the long axis of the corpus caliosum. In this histological section there is a slight artifact with
the lower brainstem being twisted to the right rather than continuing at a right angle to the corpus callosum. (approx. ‘/4x).
trast, when viewed from the anterior aspects, the foreshort- and “dorsal.” All these space reference terms, explicit and
ened brains of Tursiops (Fig. 9) and of other Cetacea (Figs. understandable when describing the forebrain of man and
10, II, 12, 13) are greatly widened in the bitemporal (trans- land-living animals, become ambiguous when applied to the
verse) diameter. forebrain of the dolphin. Formations such as frontal lobes,
A peculiarity, of considerable import from the descriptive which are “frontal” and “anterior” in the hemisphere of
standpoint, is the orientation of the orbito-occipital axis of land-living mammals (Figs. 8A-D), are ventral in the hemi-
the hemispheres with reference to the body axis (Figs. 1, sphere of the dolphin, and formations which are “occipital”
8E). Owing to foresho~e~ng of the forebrain, the cerebral or “posterior” in terrestrial forms are dorsal in the dolphin
hemispheres (including the striatum and thalamus) of the (Fig. 8E). To be consistent, positional references of parts of
dolphin, and of other Cetacea, appear as if they were “ro- the brain relative to three planes of space in the case of the
tated” on the transverse axis to a full 90 degree angle with dolphin must preferably be given with reference to the oro-
reference to the beak-fluke axis and brainstem (Figs. 1, 2A, caudal (beak-fluke) axis (Fig. 8E) of the animal, rather than
8E). This positioning of the cerebral he~spheres in Cetacea to the anthropomo~~c planes of the human brain. As will
results in certain ambiguities and a conflict of terms such as be described below, this is not just an inconsequential se-
“frontal,” “occipital,” “anterior,” “posterior,” “ventral,” mantic exercise, but has other important implications.
Text continued on page 15
MORGANE, JACOBS AND MCFARLAND
FIG. 2C. Brainstem fiexure pattern in fetal human brain at approximately 12 weeks of gestation age. In this draying the bmin is rotated
counterc~~kw~se from the normal in situ position to approximate the po&ian of the dolphin brain far campa&on, Note the remarkable
similarity of the brainstem flexure pattern in the human fetus with that of the adult dolphin. Modified fram His.
INFERIOR
.ATERAL SUL .cus
PARIETAL
OPERCULL
TEMPORAL
IPERCULUM
ORBITAl
OPERCULl JM
OF
CLEFT
LATERAL ORBITAL -SULCUS SYLVIAN CLEFT
FIG. 3. Left lateral view of brain of bottlenose dolphin (Tursiops truncatus) showing the pronounced anterior-posterior foreshortening and
the anterior rotation of the cerebral hemispheres. Note the position of the orbital lobe below the tip of the temporal lobe. Also note simile of
the hemisphere in lateral view to a pronated and cupped “boxing glove.” The Sylvian cleft cuts into the massive parietal lobe at a right angle,
i.e., “vertically,” with respect to the body axis. The temporal lobe exhibits marked foreshortening. (approx. VEX).
8 MORGANE, JACOBS AND MCFARLAND
FIG. 4. J+atera.l view of brain of beluga whale (Delphimgterus leucas) showing marked anterior-posterior- foreshortening and the anterior
rotation of the cerebral hemispheres. Note the position of the orbital lobe below the tip of the temporal lobe. Also note simile of the
hemisphere in laterat view to a pronated and cupped “boxing glove.” The Sylvian cleft cuts into the massive par&& lobe at a right angle,
i.e., “vertically,“ with respect to the body axis. The temporal lobe exhibits marked foreshortening. In this and several other subsequent
figures the gyri have been spread by akohol immersion to reveal with more clarity the various sulci. String has been inserted into the depths
of certain sulci to aid sulcal analysis. (approx. 1 x).
FIG. 5. Lateral view of brain of harbor (common) porpoise fPhocnena phocaenu) showing marked anterior-posterior
foresho~ening and the anterior rotation of the cerebral hemispheres. Note the position of the orbital lobe below the
tip of the temporal lobe. Also note simile of the hemisphere in lateral view to a pronated and cupped “boxing glove.”
The Sylvian cteft cuts into the massive parietai lobe at a right angle, i.e., “‘vertically,” with respect to the body axis.
The temporal lobe exhibits marked foreshortening. In this and several other subsequent figures the gyri have been
opened by alcohol immersion to reveal with more clarity the various sulci. String has been inserted into the depths of
certain sulci to aid analysis of sulcal patterns. (approx. 2x).
FIG. 6. Lateral view of brain of Amazon River dolphin (hia geoffrensis) showing marked anterior-posterior foreshortening and the anterior
hemisphere in lateral view to a pronated and cupped “boxing glove.” The Sylvian cleft cuts into the massive parietal lobe at a right angle,
i.e., “vertically,” with respect to the body axis. The temporal lobe exhibits marked foreshortening. In this and several other subsequent
figures the gyri have been opened by alcohol immersion to reveal with more clarity the various sulci. String has been inserted into the depths
of certain sulci to aid analysis. (approx. l’ldx).
FIG. 7. Left lateral view of brain of fin whale (Balaenoptera physalus) showing marked anterior-posterior foreshortening and the anterior
hemisphere in lateral view to a pronated and cupped “boxing glove.” The Sylvian cleft cuts into the massive parietal lobe at a right angle,
i.e., “vertically,” with respect to the body axis. The temporal lobe exhibits marked foreshortening. In this and several other subsequent
figures the gyri have been opened by alcohol immersion to reveal with more clarity the various sulci. String has been inserted into the depths
of certain sulci to aid analysis. (approx. 0.6x).
CALLITHRIX ARGENTATA CANIS FAMILIARIS

(Marmoset Monkey) (Dog)
PONGO SATYRUS
(Orang-utan) %
,r
HOMO SAPIENS
(Man)
5ody Axis
i
i
TURSIOPS TRUNCATUS
(Bottlenose Dolphin)
Beak-Ftuke 03ody)Axig
4
FIG. 8. Illustration showing in lateral view the brains of the (A) monkey, (B) dog, (C) orang-utan, (D) man, +d (E) .dolphin, ayd
indicating the position of each with respect to the long axis of the body. Note that in man and dolphin the fronto-occlpltal axis of the brain
is approximately at right angles to the body axis, whereas in the dog this axis is more nearly parallel to the body axis. In the monkey this
axis is in an intermediate position.
PARIETAL LOBE pAmt~htc CLEFT (entolakral fissure ; rulcur confinis)
*SUPERIOR LATERAL
FISSURE
(ectoloteral wlcus)
‘CRUCIATE SULCUS
TEMPORAL ILOBE
r,
0RBITAL”lOBES CORONAL SULCUS
FIG. 9. Frontal view of Tursiops trunca?uS brain showing the course of the cruciate sulcus and the origin and course of the parahmbic cleft.
Note the proximity of the orbital “lobes” and the great bitemporal width of the hemispheres in this view. (approx. V3x).
SUPERIOR LATERAL FISSURE (ectoht-1 SUlCUd
INTERMEDIATE LATERAL
FISSURE
(ruprasylvion rulcur)
PARALIMBIC CLEFT
(entolaterol fisrur* ;
rulcur confinis\
INFERIOR LATERAL
FISSURE (cleft!
(sctosylvian rulcur)
TEMPORAL LOBE
*CRUCIATE SULCUS
SYLVIAN CLEFT
ORBITAL LOBES coronal sulcus

FIG. 10. Frontal view of Delphinapterus leucas brain showing the course of the cruciate sulcus and the origin and course of the parahmbic
cleft. Note the proximity of the orbital “lobes” and the great bitemporal width of the hemispheres in this view. (approx. r/zx).
SUPERIOR LATERAL FISSURE
(ectolaterol SUlC”S)
PARALlMBlC CLEFT
(entoioteral fissure;
sukus confinis)
SYLVIAN CLEFT
TEMPORAL LOBE
-.
TRANSVERSE TEMPORAL
GYRI
SYLVIAN CLEF i CRUCIATE SULCUS ORBITAL LOBES
FIG. 11. Frontal view of Phocaena phocaenn brain showing the course of the cruciate sulcus and the origin and course of the paralimbic clc:ft. Note the
proximity of the orbital “lobes” and the great bitemporal width of the hemispheres in this view. (approx. 1 x).
superior loteral fissure (ectoloterol sulcur)
PARALIMBIC CLEFT
[entolateral fissure;
rulcur confinis)
superior lateral
ectolateral sulcus)
ORBITAL LOBES
FIG. 12. Frontal view of Inia geoffrensis brain showing the course of the cruciate sulcus and the origin and course of the paralimbic cleft.
Note the proximity of the orbital “lobes” and the great bitemporal width of the hemispheres in this view. (approx. ‘/4x).
It should be noted that in the dolphin and other Cetacea at a much wider angle open dorsad (Fig. 9) than in man and
the deep Sylvian cleft cuts into the massive hemisphere most land-living mammals. This feature of the cetacean fore-
at nearly a right angle to the body axis (Figs. 3-7, 8E). brain is particularly prominent in Phocaena phocaena . In the
Viewed from the side, the cerebral hemisphere of Cetacea Delphinidae the “frontal” lobes, oriented sharply ventrad,
presents a simile of a boxing glove pronated and cuffed over are small relative to the massive parietal and temporal lobes.
the thalamus and brainstem. The hemispheres arch dorsad Because of the ventral orientation and relative hypoplasia of
from the orbital (frontal) to the temporal poles in steep arcs the frontal lobes, we refer to them as orbital rather than
concave ventro-mediad. Taking the foramen of Monro as the frontal lobes. It is noteworthy that the orbital lobes in
center of the arc, its circumference from the orbital to the Cetacea, as in all mammals other than man, do not project
temporal pole may be taken as the index of rotation of the forward above the orbits as do the frontal lobes in man but
hemisphere [60] and has been taken by Pilleri [49] as an index rather are postero-mediad to the orbits (Figs. 1,14). They are
of “encephalization” of the mammalian central nervous sys- situated in the narrow anterior cranial fossa immediately
tem. In the rabbit the index is 260 degrees, in the dog 280 behind the “blowhole” and bony nares in Cetacea (Fig. 1).
degrees, in the primates (rhesus monkey and orangutan) 300 The orbital lobes in Tursiops truncatus (Fig. 9), Del-
degrees, and in man 320 degrees [60]. In Cetacea (Physeter phinapterus leucas (Fig. 10) and Phocaena phocaena (Fig.
catodon) it is 330 degrees [49]. The temporal and parietal 11) are in close apposition separated only by pia-arachnoid;
lobes expand far lateral from the median plane (Figs. 9,lO). there is no falx in the interhemispheric cleft between these
In consequence of the rostro-caudal foreshortening and lobes. There is, however, a slight diastasis between the rela-
transverse widening of the hemispheres, the shape of the tively larger and more prominent orbital lobes of Znia geof-
brain of the cetaceans we have examined (8E) is globoid and frensis (Fig. 12) and of Balaenoptera physalus (Fig. 13). In
presents no sharp polar angulations comparable to the fron- all the Cetacea we have examined, the interhemispheric cleft
tal and occipital poles of the primate and human hemispheres is shallow anteriorly and ventrad from the corpus callosum
(Figs. 8A-D). Since the transverse diameter of the temporal (Figs. 15-19). However, over the corpus callosum it deepens
lobes is extremely wide in Cetacea, the long axes of the two and widens posteriorly between the blunt occipital “poles”
hemispheres viewed frontally diverge from the orbital lobes of the hemispheres. A short, bony falx projects from the
SUPERIOR LATERAL FISSURE SUPERIOR LATERAL FISSURE
(ectolateraf sulcur) (ectolaterol sulcur)
INTERMEDIATE
LATERAL FISSURE
(suprasylvian
INFERIOR LATERAL ‘:
FISSURE [cl&)
(ectosylvian fissure)
PARALIMBIC CLEFT
SYLVEAN CLEFT-
(entolateral fissure;
Sulcus confinis)
(entolateral fissure ;
p::::n::ys==l
cruciate SUICUS d
CIRBiTAL LOWS’
FIG;. 13. Frontal view ofl3ultri,r?oprrrtr)pf[~r~~

ph~.scr/rr.\ brain showing the course of the cruciatc sulcus and the origin and course’ oi the p;~ralimhic
cleft, Note the proximity of the orbital “lobes” and the great bitemporal width of the hemisphere in this view. (approx. 1: ).
17
6 GORILLA
c DOLPHIN t
body axis
FIG. 14.Sketch showing the positions of the brain in the skull of man (A), gorilla (B), and dolphin (C). Note the
position of the frontal (or orbital) lobes in relation to the orbits (or) in man as compared to the gorilla and dolphin.
Drawings are not to scale.
18 MORGAN& JACOBS AND MCFARLAND
CLEFT
(suprospkniaf sulcur)
tectam.
massa intermedia
thaiomi Csciola cinsze&
/
corprts callorum
external intwcalate tasnSa Ierta

ltnibir sufcus 14,
fornix MESOCELE
‘ZI. +iSyfvkn aqueduct)
LiMBiC CLEFT
(suprosplenial s&as]
oblique sufcus
$‘p‘
internal intarcalate
Iimbic rulco$
RHINIC CLEFT
jrulcus of corpus
ftHiNlf CLEFT
collosum ) hippacompol rufcws)
ontsrior commissurs’
AVUM.SEPTI
LfMBIC CLEFT - LAMtNA ROSTRALJS

fsplenial sulcus)
\ \
anterior par&factory rulcus’ anterior rhinai SU~CVJ oilactary tuber& L
infondibutum
parolfactory tobute of Brcca4
posterior parolfactory solcvs
FIG. 1.5. Photograph of medial view of right hemisphere of T~siops lru~(‘~~us brain shawing the general profile of the he~~hore and
relations of the hemisphere to the upper brainstem. The relations of the cortical fo~ations to the vertically oriented ccq~t~s callosum are
also shown in this figure. Note that the orbital and temporal lobes point ventrad. The paucity of cortex anterior to the genti of the cOrpus
callosum points up the reduced frontal lobes in this species. The great expansion of the medial cortical formations (timbic and paraiimbic
lobes) is also indicated in this figure. Note also the great dorsal proplasia of the parietal lobe (cortex above the parajimbic cieftl. This figure
also shows the limbic cleft in almost its entire course. (approx. ‘ir xl.
INTERCALATE SULCUS OF THE OVAL LOBULE INFERIOR MARGINAL WLCUS
PARALIMBIC CLEFT
(ENTOLATERAL FISSURE)
PARALIMBIC CLEFT
(ENTOLATERAL Flssu RE)
\ &
SUPERIOR MARGINAL-_
LIMBIC CLEFT
(SUPRASPLENIAL SULCUS)
,FASCIOl.A CINEREA
,TAENIA TECTA
LFIMBRiA
-0BLlOUE SULCUS
(SULCUS OF COR
CAVUM SEPTI
- LIMBIC CLEFT
(PARAH;;Q$MPAL
PAROLFACTORY
Clll~,,C
PAROLFACTORY LOBULE OF BRtJCA OLFACTORY LOBE MASSA INTERMEDIA THALAMI
FIG. 16. Photograph of medial view of right hemisphere of Delphinapterus leucas brain showing the general profile of the hemisphere and
relations of the hemisphere to the upper brainstem. The relations of the cortical formations to the vertically oriented corpus callosum are
also shown in this figure. Note that the orbital and temporal lobes point ventrad. The paucity of cortex anterior to the genu of the corpus
callosum points up the reduced frontal lobes in this species. The great expansion of the medial cortical formations (limbic and parahmbic
lobes) is also indicated in this figure. Note also the great dorsal proplasia of the parietal lobe (cortex above the paralimbic cleft). This figure
also shows the limbic cleft in almost its entire course. (approx. l/2x).
20 MORGANE, JACORS AND MCFARLAND
cokarine *uikw (?i

7
PARALtwK CtEFT
(entoloterol firrur4
ia@%aiaf# 5UkU8 Qf
- -=tlT* 0”Cl lobuk
mrdiaf senicufote
body
inferior
merqinol rulcur
bliqu* 5UlCUE
LIMBIC CLER
(supraspknial
sukur)
intorcolak limbic
SUICUS
frplanial sukur)
sulcur of
corpus catlorum
foromrn of
MOflWJ
antodor
parolfactory sulcvr
%
porolfoctory lobule of firnca anterior rhino1 rukur
FIG. 17. Photograph of medial view of right hemisphere of Phocaena phocctena brain showing the general profile of the hemisphere and
relations of the hemisphere to the upper brainstem. The rektions of the cortical formations to the vertically oriented corpus caltosum are
also shown in this figure. Note that the orbital and temporal lobes point vent&. The paucity of cortex anterior to the genu of the corpus
callosum points up the reduced frontal lobes in this species. The great expansion of the medial cortical formations (limbicand paralimbic
lobes) is aIs0 indicated in this fgure. Note also the great dorsal proplasia of the parietal lobe (cortex above the paralimbic cieft). This figure
also shows the limbic cleft in almost its entire course. (approx. 1 x).
farciolo cinema PARALLMBIC CLEFT (sntokzterof fissure)
OLFACTORY LOBi hippccompol rutcur UBXUI
FIG. 18. Photograph of medial view of right hemisphere oflnia geoffrensis brain showing the general profile of the hemisphere and relations
of the hemisphere to the upper brainstem. The relations of the cortical formations to the verticalfy oriented corpus callosum are also shown
in this figure. Note that the orbital and temporal lobes point ventrad. The paucity of cortex anterior to the genu of the corpus cailosum points
up the reduced frontal lobes in this species. The great expansion of the mediaI cortical fo~ations (limbic and paralimbic lobes) is also
indicated in this figure. Note also the great dorsal proplasia of the parietal lobe (cortex above the paralimbic cleft). This figure also shows the
limbic cleft in almost its entire course. (approx. VEX).
M
PARALIMBIC CLEFT (entolateral fissure)
thalam
us\
LIMBIC CLEFT
7
er cternal intercal late
limbic sulcus \
ini ternal i ntercal ote

s&us
\
limbic
ante1 *ior
Pa rolfacto ry sulc :us\ \ oblique I sulcus
iporohi ppocampS al sulcus
OLFAC:TORY LO 4%
(Qlfoctol ry tuber< :le
- ‘lobu lie d&serf 3
LIMBlC CLpFT kterior rhino1 sulcus
FIG. 19. Photograph of medial view af right’ hemisphere ofBalqenopteru,physalus br+ showing the general profile of the hemisphere and
relations of the hemikphefe to thp up&r @rahtes@l The telakms of the cortical formations to the vertically oriented corpus ca&sum are
also shown in this Fiiure. NoWthat the orbit&&l tempoti to&s point ventrad. The paucity of cortex anterior to the genu of the corpus
callOsum points up the reduced firontal l&es ‘in .this ~pec&s. Tfie great expansion of the medial cortical formations (Iimbic and paralitibic
lobes) is also indicated in this figure. Note also the great dorsal propgsia of the parietal lobe (cortex above the paralimbic cleft). This figure
also shows the limbic creft in almost its entire course. (ap$rox. 21’3 x).
bony tentorium between the occipital lobes, but its anterior rotation of the hemisphere on the transverse axis with refer-
border does not extend above the corpus callosum beyond ence to the brainstem (Figs. 15-19). The corpus callosum in
the splenium. The medial walls of the parietal lobes remain in Tursiops, as compared to that of a human brain of compara-
close juxtaposition to the level of the splenium. However, ble size and weight, is smaller and shorter antero-posteriorly
posterior to the splenium in all species examined the in- (Figs. 2A, 2B), yet the ratio of the length of the corpus cal-
terhemispheric cleft opens widely into the transverse cleft of losum from the knee to the splenium, to the length of the
Bichat over the massive cerebellum (Figs. 20-24). The oc- foreshortened hemispheres from the orbital to the occipital
cipital poles are rounded and blunt, as if foreshortened. As pole is greater in the dolphin than in all other mammals we
indicated, the diastasis between the occipital lobes is widest have investigated [42]. Taking the length of the hemisphere
in Phocaena phocaena (Fig. 22). as a constant, the ratio of the length of the corpus callosum
Viewed from the basal surface, the forebrain of Tursiops to that of the hemisphere in the common dolphin, Delphinus
truncatus (Fig. 2.5), Delphinapterus feucas (Fig. 26) and delphis, is 51:lOO as compared to 44:lOO in man, 43:lOO in
Phocaena phocaena (Fig. 27) present some differences as chimpanzee, 41: 100 in haphale, 34: 100 in seal, 36: 100 in dog,
compared to Inia geoffrensis (Fig. 28) and Balaenoptera 34:lOO in anteater, and 28:lOO in rabbit (Filimonoff, 1949).
physalus (Fig. 29). In the latter two species the orbital lobes Furthermore, in Tursiops truncatus the main mass of the
are narrower and longer than in the former three species. corpus callosum appears shifted posteriorly towards the
The olfactory lobes are relatively smaller in Znia geoffiensis splenium as is the main mass of the hemispheres. The splen-
(Figs. 18,28) and, particularly, in Balaenopteru physalus ium extends caudad and is deflected ventrad at nearly a right
(Figs. 19,29). The optic nerves and tracts in the latter two angle to the body of the corpus calIosum (Figs. 2A and 15).
species are also smaller. In all the toothed whales or odonto- The thickness of the splenium in midsagittal section is con-
cetes (Tursiops truncatus, Delphinapterus leucas, Phocaena spicuously greater than the width of the genu (knee) in the
phocaena, and Znia geoffrensis) the massive eighth nerve dolphin (Fig. 2A), a relationship which does not hold in man
roots and the absence of olfactory bulbs and tracts (Figs. (Fig. 2B). The forceps major (Fig. 36) “flares out” from the
25-28) contrast with the relatively small roots of the eighth splenium at a wide angle open posteriorly. The rostrum of
nerve and the presence of long olfactory tracts in Balaenop- the corpus callosum in Tursiops truncatus is short and nar-
tera physalus (Fig. 29). row and contains few commissural callosal fibers. The for-
The medial wall of the hemisphere in Tursiops (Fig. 15), ceps minor (Fig. 36) is open ventrad from the knee in dolphin
and in other Cetacea (Figs. 16-19), as compared to the me- (Fig. 37) rather than rostrad as in primates and man (Fig. 38).
dial wall of the hemisphere of carnivores (Fig. 30), primates The cavum septi is wide in transverse diameter and nearly
(Fig. 31), and of man (Fig. 32), is greatly expanded dorsal round in shape (Figs. 15-18). However, the cavum septi,
and posterior to the corpus callosum. In contrast, anterior which in ungulates, carnivores, and monkeys communicates
and ventral to the corpus callosum, the orbito-medial wall of anteriorly with subarachnoid space, is closed in Tursiops
the hemisphere, corresponding to the medial wall of the fron- truncatus by the lamina rostralis (Fig. 15), as it is in an-
tal lobes of the carnivores and primates, is greatly reduced in thropoid apes and in man [54].
Tursiops and other Cetacea (Figs. 15-19). The temporal
lobes are also foreshortened (Figs. 3-7,8E) and the temporal
II. THE RHINENCEPHALON
pole does not project beyond the uncus (Figs. 15-19) as it
does in man (Fig. 33). In cetacean brains, the long axis of the The earliest and clearest statement of the distinction be-
temporal lobe is at a right angle to the beak-fluke axis (Figs. tween the rhinencephalon and the cerebral hemispheres is
3-7, 8E). Viewed from the postero-medial and basal aspects found in Turner’s (1891) writings. He observed that when the
(Figs. 15-19), the planes of the baso-medial wall of the tem- surface of a cerebral hemisphere is carefully examined, it is
poral and of the occipital lobes form an obtuse angle, and the seen to be capable of a natural division into two parts: a basal
foreshortened and blunt occipital pole blends with the region or rhinencephalon, and a superior portion, or pallium.
baso-medial wall of the temporal lobe in a gently curved He further noted that the separation of the rhinencephalon
plane. Because of the prominence of the parietal convexity from the pallium or cerebrum proper by a rhinal fissure is
of the massive and globoid hemispheres, the profile of the fundamental and is seen even in the lowest vertebrates. Not
medial wall of each hemisphere (Figs. 15-19) presents four until Filimonoff provided the tectogenetic (1938) and com-
rounded ’ ‘poles”-the orbital and temporal poles pointing parative cytoarchitectonic (1949, 1965) criteria for definition
ventrad, and the obtuse parietal and occipital poles pointing of the rhinencephalon and pallium has it become possible to
dorsad and posteriorly. remove the grounds for confusion and controversy. The pal-
The configuration of the lateral ventricles in Tursiops lium (neocortex, isocortex) is defined by the presence of the
truncatus conforms with and underscores the peculiarities of cortical plate, the superficial part of which (supragranular
the configuration of the hemispheres. When viewed in their layers II-IV) develops in all mammals in the marginal layer of
natural orientation with reference to the longitudinal body the embryonal hemispheric evagination. The cortical plate of
axis, the casts of the lateral ventricles show a striking differ- the hemisphere has a sharp clinoid border which superlami-
ence in the dolphin (Fig. 34) as compared to those present in nates but does not extend over the rhinencephalon through-
terrestrial mammals, such as the dog (Fig. 30), monkey (Fig. out the circumference of the hemispheric stalk. The rhinen-
3 l), and man (Fig. 32). The lateral ventricles in Tursiops cephalon itself has no cortical plate. The superficial cell con-
truncatus form arcs concave ventrad. The anterior horns are densation in the olfactory lobe (paleocortex) and the cell
rudimentary and there are no occipital horns. The long in- plate of the hippocampus and subiculum (archicortex) corre-
ferior horns are oriented ventrad at nearly a right angle to the spond at best to the deep layers of the pallium (infragranular
beak-fluke axis (Figs. 34,35). layers V-VII). The lamina zonalis or molecular layer of the
The orientation of the corpus callosum, as seen in mid- cerebral cortex corresponds to the marginal layer of the em-
sagittal section through the interhemispheric cleft and sep- bryonal hemisphere. The stratum zonale of the rhinencepha-
tum pellucidum, reflects in Tursiops and other Cetacea the lon (“white cortex” of Meynert) develops later and is not
PARALIMW. CLEFT
~(eitfoktwal
FRANSWERSE CLEFT OF IlKHAT
FI( 3. 20. Dorsal view of brain of Tursiops trmcatus showkg the great width of this brain in its bitemporal (transverse) diameter. Most of
doI-Sal surface of the cerebellum is not covered by the cerebral hemispheres in this species. Note that the irrterhemi>pheric cleft open s widely
intoD the transverse cleft of Bichat over the large cerebellum. (approx. ‘/4 XL
25
PARALIMBIC CLEFT (entolateral sulcus) superior lateral fissure (ectoloteral sulcu 4
TRANSVERSE CLEFT OF BfCHAT fNTERH~MfSPHERfC CLEFT
FIG. 21. Dorsal view of brain ofD&hinupterus leucas showing the great width of this brain in its bitemporal (transverse) diameter. Most of
dorsal surface of the cerebellum is not covered by the cerebrai hemispheres in this species. Note that the interhemisphe~c cleft opens widely
into the transverse cleft of Bichat over the large cerebellum. (approx. )/4x).
26 MORGAN& JACOBS AND MCFARLAND
PARALCMW CLEFT (ontofated fissure) superior lahrol firsufrr (etctolaf~al sukus]
\ n
FIG. 22. Dorsal view of brain of Phocaena phocaena show@ the great width of this brain in its bitemporai (transverse) diameter_Most of
dorsal surface of the cerebellum is not covered by the cerebral hemisphere> in this species. Note that the @terhemisphetic ctift opens wkkly
into the transverse cleft of Bichat over the large cerebellum. In this species thehemispheres are spread very wi&apart posteriorly.~@pprox.
Ix).
TRANSVERSE CLEFT OF illCHAT -INTERHEMISi’HERIC CLEFT

FIG. 23. Dorsal view of brain of Inia geoffrensis showing the great width of this brain in its bitemporal (transverse) diameter. Most of dorsal
surface of the cerebellum is not covered by the cerebral hemispheres in this species. Note that the interhemispheric cleft opens widely into
the transverse cleft of Bichat over the large cerebellum. (approx. 1%x).
interamdiate latqwal fissure (suprasylvian sulcus]
INFERIOR LATERAL
superior lateral
) FISSURE (cleft)
fissure (ectosylvian sulcus)
(ectolateral sukus]
SU?RALtMBtC LORE
-PARALIWIBiC CLEFT
(entolatafal fissure)
tNTERHEMtSPHLRtC
CiEtT
TRANSVERSE Ct.WT OF BtCNAT

I-IG. 24. Dorsal view of bram ofBtr/~c~*~roprfrcr ph.vva1rr.s showing the great width of this hrain in its bitemporal (transverse) dlametcr. Most ot
dorsal surface ofthe cerebellum is not covered by the cerebral hemispheres in this species. Note that the interhcmisphcric cleft opens wide&
Into the transverse cleft of Bichat over the large cerebellum. (approx. V~XI.
ORBITAL LOBES
OLFACTORY LOBE gyrur proreur medial orbital rulcus
lateral orbital
V SENSORI sulcus
tronrverre
orbital gyrur
OPT‘IC NERVE
V MOTOR
VII
VIII
FIG. 25. Ventral view of brain of Tursiops truncatus showing orbital and olfactory lobes and orienting surface landmarks including several
cranial nerves. (approx. l/2x).
ORBITAt LOBES medial
‘OLFACTORY
lral orbital,
,porterior rhi
rulcus
,v
VII
VIII
VI
FIG. 26. Ventral view of brain of Delphinapterus leucm showing orbital and olfactory lobes and orienting surface landmarks inI ludiog
several cranial nerves. (approx. ‘/2X).
ORBWAL LOBES
orbital opwculum Olfactory *yru* (*trio) tranrvbrre orbital gyru~ Jakrol orbital
OLFACTORY LOBE n/h

olfwtory tubercle
gyrur ambimnr L
LIMBIC CLEFT-+
FIG. 27. Ventral view of brain of Phocaenu phocarno showing orbital and olfactory lobes and orienting surface landmarks m&ding several
cranial nerves. (approx. ‘/4X).
31
anterior rhino1 IUICUS medial orbital SUICUS ORBITAL LOBE
gyrus proreus parolfactory lobule of Broco

cruciate s&us
I I I I / /
lateral orbital
OLFACTORY LOBE
sulcus Ifoctory tubercle-
posterior orcuate
olfactory gyrus
se~micircular sulcur
of the insula
diagonal band-
d
- sulcus
porte rior rhino1 SUICUS _remiannularir
‘VIII
rhino-hippocompol/ ,.
sulcus
‘VII
LIMBIC CLEFT
parohippocampal
‘UrICUS
‘IX, x
hippocampal sclcus
FIG. 28. Ventral view of brain of Inia geoffiensis showing orbital and olfactory lobes and orienting surface landmarks including several
cranial nerves. (approx. ‘/2X).
32
0
U
0
=
t
0
,’
.;
0.
0
Sylvion aqueduct
III VENTRICLE INFUNDIBULAR RECESS
FIG. 30. Medial profile of the right cerebral hemisphere in the dog indicating the orientation and configuration of the ventricular system with
respect to the hemisphere in this species. Note the very limited dorsal-posterior expansion of the medial wall of the hemisphere as compared to the
dolphin (Fig. 34). (approx. 1X).
foramen of Monro BODY OF THE LATERAL VENTRICLE
ICT
/
Ill VENTRICLE
INFERIOR HORN lateral rec*ss of IV ventricle

FIG. 3 1. Medial profile of the right cerebral hemisphere in the rhesus monkey indicating the orientation and configuration of the ventricular
system with respect to the hemisphere in this species. Note the very limited dorsal-posterior expansion of the medial wall of the hemisphere
as compared to the dolphin (Fig. 34). (approx. %x).
34 MORGANE, JACOBS AND ~~~ARLAN~
olfactory bulb olfactory tract temeoral Dole
ORBITA 1 LOB
OPTIC NERV
ANTERIOR PERFORATE SUBSTANCE (olfactory lobe) uncus
FIG. 33. Ventral view of human brain showing the extension of the temporal poles far anterior to the uncal area. Note the great Projection of
the frontal lobes anterior to the anterior perforated space in the human as compared to the dolphin (Fig. 25). (approx. 1X).
36 MORGANE. JACOBS AND MCFARLAND
FIG. 34. Me&i profile of the right cerebral hemisphere and of the ventricular system in the do&tin indicating hew the [email protected] of
components of the vent&&r system, especially the lateral ventricles, reflect the rotation and ~~~~0~. of the hemisphere. Note that
the lateral ventricles in Tursiops truncafus form arcs that are concave vetirad and tha& there are onl$ rudimentary anterior horrrs and
complete absence of occipital horns. The inferior horns are foreshortened and oriented vertically in the plane of the Sylvian cleft. (approx.
V3x).
RCtDY INFERIOR HORN
MESOCELE
lvian aquedu 1ctl
VENTRICLE
.TERAL RECES IS
IV VENTRIC ILE
FORAMEN
OF MONRC
MASSA
INTERMEDIA
THALAMI
ANTERIOR HORN
FIG. 35. Cast of the entire cerebroventricular system of the dolphin brain. Note that the long inferior horns, as well as the bodies of the lateral
ventricles, are oriented ventrad at nearly a right angle to the beak-fluke axis.
FORCEPS ANTERIOR (minor] caudots nucleus

\ ,
rlomus
forni
‘-’v
FoRcEPS 605~~~0~ (m&r]
FIG. 36 Photograph of dolphin brain section in horizontal plane showing the forceps major and forceps minor. Note that in the dolphin I bram
the j:iWS of the forceps major or “posterior” forceps are wider, longer, anddiverge from the spknium laterad rather than caudad as they do ‘in the
ml ,rain (Fig. 38). Loyez myelin stain. (approx. 1x).
OVAL LOBULE OCCIPITAL LOBE iNTERHEMISPHERIC CLEFT (supracaliorof part)
.EFT
bit
\
forceps porteriorl,
(major)
(ET
*s
I
forceps anterior
(minor)
optic nerve
ORBITAL LOBE INTERHEMISPHERIC CLEFT (rubcollorol part)
FIG. 37. Photograph of dolphin brain section in frontal plane showing that the jaws of the forceps minor (anterior) in the dolphin open less
widely than in the human (Fig. 38). This section also shows the “triple-tiering” of the cortical formations corresponding to the three main
lobes of the hemisphere, i.e. limbic, paralimbic and supralimbic. Loyez myelin stain. (approx. */4x).
40 MORGANE, JACORS .4NI> Mc,t-ARL,ANl>
FIG 38. Photograph of horizontal brain section (frontal poles downward) in human. Note that the forceps minor opens rostrad in
rath er than ventrad as in the dolphin (Fig. 37). Loyez myelin stain. (approx. l’/zx).
comparable to the molecular layer of the cerebral cortex. It is compared to 0.6% in man, 1.4% in macaque, 6.8% in dog,
comparable rather to the superficial white matter of the spi- and 14% in rabbit. We have described the olfactory lobes in
nal cord and brainstem. On the basis of these cardinal mor- our earlier study on the paleocortex of Cetacea [28].
phological premises by which the rhinencephalon is distin- Anteriorly and laterally, the olfactory lobes are clearly
guishable from the cerebral hemispheres in all mammals, we demarcated from, respectively, the orbital lobes and the in-
shall describe first the surface morphology and boundaries of sulae by the shallow but constant anterior rhinal sulci (Figs.
the rhinencephalon. 39-47) and posteriorly, by the telodiencephalic sulci (Fig. 47)
The rhinencephalon in Cetacea, as in all mammals, forms from the area of the septo-hypothalamic junction and cere-
the common median pediment of the paired cerebral hemi- bral peduncles. Each sulcus rhinalis anterior arises medially
spheres and consists of three divisions (Figs. 39,40): (1) the in the septal area under the rostrum of the corpus callosum
septal area (area septalis) dorsal to the lamina terminalis; (2) as a shallow groove from the posterior parolfactory sulcus
the so-called olfactory lobes; and (3) the hippocampus [28]. (Figs. 40, 41A, B) and extends ventrad to the ventromedial
The rhinencephalic formations of the basal forebrain are re- border of the orbital lobe and, thence, in an arc, laterad and
duced in the Cetacea to their elementary components and are posteriorly to the isthmus of the temporal lobe (Fig. 47). The
revealed with an almost diagrammatic clarity. Here, similar- olfactory gyrus (gyrus X of Breathnach, [Sl) of the olfactory
ities to the formations of the basal forebrain in all other lobe follows the sulcus rhinalis as a whitish band (olfactory
mammals are as if underscored and allow the least doubt stria, G 01, Fig. 47) from the septal area to the uncus where
with regard to existing homologies [5, 16, 17, 281. this band of superficial fibers blends with the gyrus semilun-
aris of the uncus. At the isthmus of the temporal lobe the
I. Septal Area sulcus rhinalis becomes shallow and turns at a sharp angle
laterad onto the medial wall of the temporal lobe (Figs.
The septal area (Figs. 39,40,41A, B) is well-developed in
47,48) and extends rostrad towards the uncus. Here it be-
Tursiops truncatus and the other four cetacean species
comes deeper and turns dorsad between the gyrus ambiens
examined. It extends from the lamina terminalis to the floor
of the uncus and the temporopolar end of the parahippocam-
of the cavum septi. It is traversed posteriorly by the slender
pal gyrus (Figs. 40,42,47). In the medial wall of the temporal
anterior commissure and is bounded bilaterally by the mas- lobe the sulcus rhinalis is designated as the sulcus rhinalis
sive precommissural fomix (Figs. 40, 41A). It forms the tel-
posterior (Figs. 40, 41B, 42, 47, 48, 49) demarcating the
encephalic wall of the median (rhinencephalic) telocoele
uncus from the parahippocampal gyrus (Figs. 50, 51, 52,
(cavum of Monro) anterior to the large massa intermedia
53A, B, 54A, B). In all cetacean brains examined the sulcus
thalami and third ventricle (diocoele). The septal area is a
rhinalis posterior was found to be continuous through a shal-
median formation folded upon itself in the geometric mid-
low connecting groove with the hippocampal sulcus (Figs.
sagittal plane in the floor of the interhemispheric cleft and
40,47). Because of the constancy of this connecting groove in
blends bilaterad with the diagonal band of Broca of the olfac-
Cetacea we designate it as the sulcus rhino-hippocampalis.
tory lobe and dorsad with the septum pellucidum and hip-
The sulcus rhinalis, throughout its course from the sulcus
pocampal rudiment (Figs. 40, 41A, B). A shallow posterior
parolfactorius posterior to the sulcus rhino-hippocampalis
parolfactory sulcus (Figs. 41A, B), barely perceptible to the
demarcates the limes (threshold) of the olfactory lobe
naked eye, extends from the rostrum of the corpus callosum
(paleocortex) of the rhinencephalon from the limbus (border)
towards the lamina terminalis and may be taken as the sur-
of the hemisphere (neocortex).
face demarcation between the diagonal band of the olfactory
The olfactory lobe in Tursiops truncatus and other
lobe and the septal area with taenia tecta in its median fold.
Cetacea is a large and prominent elevation bounded an-
The posterior parolfactory sulcus is a well-defined, distinct
teriorly by the olfactory gyrus (Figs. 42, 43, 44, 45, 46, 47).
groove (Fig. 41A, B) which is a continuation of the sulcus of
The elevation, designated by Broca [6,7] as the “lobule dbs-
the corpus callosum from the rostrum onto the medial wall of
ert,” corresponds in position to the olfactory tubercle of the
the orbital lobe and demarcates the olfactory stria or gyrus of
anterior perforated substance and to the projection of the
the olfactory lobe from the parolfactory lobule or area on the
head of the caudate nucleus on the surface of the olfactory
medial wall of the hemisphere (orbital lobe).
lobe 151. Anteriorly, it is demarcated from the olfactory
gyrus by the anterior arcuate sulcus of Guldberg (Figs. 41A,
2. Olfactory Lobe
B, 44, 47). Posteriorly, this elevation is demarcated by the
The olfactory lobes (region of olfactory turbercle and an- shallow posterior arcuate sulcus of Guldberg (Figs. 41B, 44,
terior perforated space) in the adult Tursiops truncatus, as in 45,46,47) from the diagonal band of Broca (Figs. 28,39,42,
all Cetacea, are prominent formations at the base of the or- 44, 45, 46) which is lighter in color than the “lobule d&sert”
bital lobes (Figs. 15-19, 42-46). Relative to the size of the and is pitted by blood vessels. The diagonal band extends
massive hemispheres, however, the olfactory lobes are from the septal area (Figs. 39, 40, 41A, B) ventrad and
small. Nevertheless, relative to the orbital lobes of the hemi- laterad, parallel to the optic tract, to the isthmus of the tem-
spheres (pallium) which overlie them, they are large in all poral lobe.
Cetacea (Figs. 15-19). The contrast between relatively large
olfactory lobes and reduced orbital lobes in Cetacea becomes
evident when compared with greatly reduced olfactory lobes 3. Hippocampus
(anterior perforated substance and olfactory tubercle) and We have described this formation and other parts of the
greatly expanded frontal lobes in man (Cf. Figs. 33 and 42- archicortex in a recent paper [29]. In contrast to the olfactory
46). The size of the olfactory lobes in Cetacea has no relation lobe, the hippocampus in Cetacea is an inconspicuous for-
to the Presence or absence of the olfactory bulbs and tract mation. According to Filimonoff [16], its surface in Del-
(Cf. Figs. 25-28 with Fig. 29). According to Filimonoff [16] phinus delphis represents 0.8% of the total surface of the
the surface of the olfactory lobe (paleocortex) in dolphins cerebral hemisphere as compared to 2.2% in man, 4.0% in
represents 0.9% of the total surface of the hemisphere as macaque, 6.3% in dog, and 23.8% in rabbit.
SULCUS MEDIANUS
TELENCEPHALI MEDll AND TAENIA TECTA
I- SUBCALLOSAL SECTOR TAENIAE TECTAE
(gyrus subca Ilosus, A
hlppocampal rudiment) ALLOSAL SEC TOI
lpal rudiment)
SULCUS /
PAROLFACTORI US POSTERIOR A
ANTERIOR COMMISSURE.\
OLFACTORY
LOBE
B-
SULCUS RHINALIS “2,
POSTERIOR SULCUS OBLIQUUS
SI II Cl IS
RHINO-WIPFOCAMPALIS
------
SULCUS w
PARAHIPPOCAMPALIS -
SECTOR
I”-TEMPdRAL SECTOR’
(hippocampus ma jar)
FIti. 39. Diagram of the generai con!iguration of the cetacean rhinencephalon (rhinic lobe or median tclencephalon) rllustratinp palcc)~Mt~c‘al rlnu
archicorttcal divisions of the rhinencephalon and their regions of junction in t$e two unci and in septal area. The hemispheres are abstracted exposing the
olfactory lobes (paleocortexl and the hippocampus (archicortex). ‘The septum pellucidum. t’ornix and corpus callosum arc omitfed for clarity The
rhinencephalic formations shown In this figure are demarcated hilaterad hy the rhinic cleft from the hemispheres proper.
WPPOCAMPAL COMMISSU~E
TAENIA
SUBCALLOSAL GYRI 310LA CINEREA

JS FASCIOLARIS)
SULCUS S!.RO‘FACTOR,“S
FoslEmoR -
-DENTATE GYRUS
MEDIAL BOROER
OF ORBITAL LOBE
FIG. 40. Diagram of the basomedial aspect of the right hemisphere (end brain) of Tursiops truncafus in the mid-sagittal plane of section. This
view illustrates the relations of the archicortical and paleocortical formations to the limbic lobe, paralimbic lobe, basal-temporal part of the
supralimbic lobe and the orbital lobe. This schema shows well the surface relations of the retrosplenial and temporal sectors of the limbic
lobe to surrounding formations (also see Fig. 55 which is photograph of dolphin brain in same position as illustrated in this drawing). It is
noteworthy that the temporal pole in Tursiops forms a hub of four cardinal grooves arising anteriorly in the region of the septal area and
extending posteriorly to the region of the uncus in the temporal lobe: (1) the posterior rhinal and the hippocampal sulci between, respec-
tively, the olfactory lobe and hippocampus and the limbus of the hemisphere; and (2) the circular and parahippocampal sulci which
demarcate the insula and parahippocampal gyrus, respectively, from the operculum and from the basolateral (temporal) wall of the
hemisphere. The converging polar ends of these cardinal and constant grooves in the temporal lobe form a characteristic S-shaped gyrus
(Fig. 32) which may be designated as the temporo-polar or sigmoid gyrus of the temporal lobe. Abbreviations: G-gyrus; S-sulcus;
Splen-splenium; For Ma&--forceps major; Lat Genie Body-Lateral geniculate body; Sep Pell-Septum Pellucidum.
44 MORCANE, JACOBS AND McFARtANU
2.
IRD VENTRICLE
(diocaslel
AvUM OF MONRO
rdion
ruior
afmiilory body
f:lobula dirert’j
FIG. 41A. Medial view of the right hemisphere in Tursiops~~~cmu showing the details of the orbital area, including the origin and course
of the sulcus rbinalis anterior. Note the continuity of the sulcus of the corpus callosum via the sulcus paroifactorius posterior with the sulcus
rhinalis anterior. Note the attenuation of the body of the corpus callosum and the refative thickening of the spienium. Also. observe that the
paucity of cortex anterior to the genu of the corpus callosum points up the reduced “frontal lobes” in this species. Great expansion of the
mediai cortical fOmdOnS (limbic and paralimbic lobes) in this species compared to primates and man (Fig. 32) is also emphasized in this
figure. (approx. t %xf.
\ CILim SPfA--+/
FIG 41B. Schematic interpretation of the medial surface of the hemisphere in Tursiops rruncatus illustrating various sulci, gyri and other
formations in this area. This scheme represents an interpretive view of the areas of the brain shown in Fig. 41A. Abbreviations: CC-corpus
callosum; S CC-suicus of corpus callosumr Cav Sp-cavum septi; Spt Pl-septum pellucidurn; Fo-fomix; For Mo--foramen of Monro;
Th (Ms It@--thalamus (massa intermedia); Ar Spt-septat area; AC-anterior commissure; G Sc-subcallosal gyms; D-area diagonalis;
ind Gr-induseum griseum; P-ports; Fos I Ped-inte~duncular fossa; SH-hippo~ampai sulcus; S Rh H-rhino-hippocamp~ sulcus;
GP~-parahip~~pal gyrus; S Rh P-posterior rhinal sulcus; Tub &-tuber cinereum; GTP-temporal polar gyrus; Ch Op.--optic
chiasm; lnf-i~undibulum; T-area tubercula~s: S Arc A-anterior arcuate sulcus; S Arc P-posterior arcuate sulcus;
S Rh A-anterior rhinal sulcus; A Pf B-parolfactory area of Broca; S Pf A-anterior parolfactory sulcus; S Pf P-posterior parolfactory
sulcus; R Sup M-superior mammillary recess; G Pr-proreus gyms; G T Or-transverse orbital gyrus; Got--lateral olfactory gyrus (stria);
Cl Lim-limbic cleft; S Pf Pm-sulcus parolfactorius postremus; G Am-gyrus ambiens; G Sk-gyrus semilunaris; U-uncus; R S Op....-
supraoptic recess; GLII-internal intercalate limbic gyrus; S SAn-suicus semiannularis.
46 MORGANE. JACOBS AND McFARLANI.)
IJATUS anterior rhinal sulcus oifoctory gyrus transverse orbtital 9y,,,,

\ h
diogonol band
\
h
lotorol orbital
rvlcur
rculor tulcur
rf the inrulo
posterior rhinoi,,
sulcus
wnsv*rsa inrulor
BY’US
iyrus ombionr
optic trot p+arior rhino1
rulcur
Wmionnutcrrir
rhino-hlppocompa
sukur
LIMWCCLWT
olfectory fuberct osohippoconpot
sulcurJ
hypophysis rplmiem of corpus culiorum gyrus samtlunoris
FIG. 42. Basal view of brain of dolphin, Tursiops truncaatus. This figure shows the prominence of the olfactory lobes and the gross relation*
of the olfactory and orbital Iobes, optic tracts and temporal polar area. Note that the orbital lobes are clearly demarcated from the olfactory
lobes by the olfactory stria (gyri). (approx. 1x 1.
FIG. 43. Basal view of brain of beluga whale (Delphinapterus irucas) indicating orbito-basal structures and the prominence of the olfactory
lobes. (approx. ‘/2x).
ant*riar arcuato rulcur anterior rhinal sulcur
If
(111
ria )
SYLVIAN CLEF
to
optic nerve
olfactory tubercle (‘lobule dbrert’) vi1 vill
FIG. 44. Basal view of brain of common harbor porpoise (Phocaena phocaena) indicating orbito-basal structures and the prominence of the
olfactory lobes. (approx. 2x).
transverse orbital _.
wrus
v (motor)
trcrnv*rre insular
optic nerve
9Yrus
diagonal bond
““CU. posterior arcuate sulcus olfactory tubercle rlobule dbrsrt’)

FIG. 45. Basal view of brain of Amazon River dolphin (Ink geoffreensis) indicating orbito-basal structures and the prominence of the
olfactory lobes. (approx. %x).
olfactory tubercle (‘lobule dbsert’) mediai olfactory gyrus olfactory tract
latera II
factory gY’
pastel vior
arcuate su 1
tranrver se
sulclr g vu
antsrio r
inal sul cus
diagonal ba
ncus
FIG. 46. Basal view of brain offinback whRletRnlnc~n,,~rc,rcr ~~h~sc~lrts)indicating orbito-basal structures and the prominence of the olfactory
lobes. (approx. “4 y ).
H
‘AL
Loiim(RSp)
TENTORlAL
SURFACE OF
TEMPORAL LOBE
FIG. 47. Interpretive schema of basal view of dolphin ~Tursiops truncutus~ brain showing su~ivisions of uncal or temporal portion of rhinic
lobe and their relationship to basal olfactory areas. Note, in particular, the course of the anterior and posterior rhinal sulci and the relations
of these to the hip~~ampal sulcus. Abbreviations: S Rh A-sulcus rhinalis anterior; S Arc A-anterior arcuate sulcus; G Or T-transverse
orbital gyrus; S Or T-transverse orbital sulcus; G 01-lateral olfactory gyrus (s&a); S Or L-lateral orbital sulcus; G IRS T-transverse
insular gyrus; S Cir I-circular sulcus of insula; G Sl-gyrus semilunaris; S SAn-sulcus semiannularis; G Am-gyrus ambiens; GPH-
parahippocampal gyrus; S Rh P-posterior rhinal sulcus; SPH-pamhippocampal sulcus; S Rh H-rhino-hippocampal sulcus; SH-
hippocampal sulcus; S Obl-oblique sulcus; Cl Lim-limbic cleft; Lo Lim (R Sp)-limbic lobe (retrosplenial sector); Sp CC-splenium of
corpus callosum; Fcp Mj-forceps major; F Tr B-transverse cleft of Bichat; P Amg-periamygdalar area; D-diagonal area; T-tubercular
area; S Arc P-posterior arcuate sulcus; PP-prepyriform area (lateral olfactory gyrus); S T Di-telo-diencephalic sulcus; T Op.-optic tract.
50 MORGANE. JACOBS AND McFARLAND
inferior lotoral fissure (Aft) intarmodiato lotoral fissure superior Iatorol firsuro
fwtosykian rulcur) \ (ruproqlviin rulcurf (sulcus ectolabtr~lir of’ Guldborg )
/central rulcus
of inrulo
tranrvorre ins vlw

Qyrus
’ (ontwior an, d
portwlor pat ‘W
circular sulcv
/ Of inculo
waporcll IO&
tronwem
&mpor0l gyri
cruciatr mhua
ontorior
- tranworw
tomporol sulci
olfactory gyrur
SUICUS ilwlo-
istrial 1 ’ parahippocam
\
(orbital rector] ant&or and pwtwior gyrvr r*nitunaris porohippoc~pak gyrur
rhino1 sulci of the uncus
FIG. 48. Photograph of the left cerebrdl hemisphere of the brain in the dolphin (Turrit~p.t~ruwtrtrrs)showing the topogrdphlc relations ot the
olfactory lobe to the insular and temporo-uncal borders of the hemisphere revealed by dissecting away of operculum of lateral convexity of
hemisphere. Note sharp definition of lateral olfactory stria and its blending with the gyrus semilunaris of the uncus. Clearly shown is the
transition from anterior rhinal sulcus to the posterior rhinal sulcus that occurs at sharp angulus of the genu of the olfactory gyrus. In this
antero-lateral view the insular formation has been exposed by removing the orbital, parietal and temporal opercula. The central sulc~~sof the
insula lies in the same plane as the Sylvian cleft. (approx. 0.8~).
outer banks of the

inferior later .al
fissure
(sctosylvian su Icus)
PARALIMBIC
CLEFT
Ienltalateral fissure)
transverse central rulcur

insular of the insula
circular rulcus
ante
rhino1
olfactory
cruciate sulcuS insular gyros
gyrus ambiens parahippocampal g;rus parahipbocampal dcus~parterior rhino1 sulcus

sulcus ins&o-parahippocampalis
FIG. 49. Photograph of left hemisphere of brain of beluga whale (Delphinapterus leucas) with opercular plug removed to expose the insula
and dorsal surface of the temporal lobe. Note the continuation of the anterior rhinal sulcus onto the medial aspect of the temporal lobe as the
posterior rhinal sulcus. The sulcus rhinalis, throughout its course from the sulcus parolfactorius posterior to the sulcus rhino-hippocampalis,
demarcates the limes or threshold of the olfactory lobe (paleocortex) of the rhinencephalon from the limbus or border of the hemisphere
(neocortex). (approx. 3/4x).
outer bank of the inferior lotoral fiisuro (cleft) (ectosylvian rulcur)
control
@Sulcus of inrula
circular sukur
of inrula
@&tol rdcw)
circular tukuo
I of indm
[temporal rockr)
tranwwro
Ansular gyms
anterior
~&not sukur
tranwwso
in$daf gyms
cirr~ular~ r&em of thkr inrula (orbitof roctor)
FIG. 50. Photograph of left hemisphere of brain of common harbor porpoise ~Phocarntr phoc~na) with opercular plug removed to expose
the insula and dorsal surface of the temporal l&e. (approx. 1’14~).
PARALIMBIC CLEFT
(entolateral fissure; central
sulcus confinis) ~sulcus of insula
transverse
insular gyrur circular sulcus
of insula
(temporal sector]
parahippocampal
sulcus
/ \
circular sulcus of insula (orbital sector] porahippocampal gyrus
FIG. 51. Photograph of left hemisphere of brain of Amazon River dolphin (Ink geuffrensis) with opercular plug removed to expose the insula
and dorsal surface of the temporal lobe. (approx. ‘LX).
-L
2
_ .E
0
2”
c 0
a
” H
E
The hip~campus of Tursiops ~r~nca~us and of other extension of the hippocampus and contains in its core the
Cetacea we have examined consists, as in all mammals, of amygdala (Fig. 39). The uncus thus represents the region in
two divisions 1291: (1) the hippo~amp~ rudiment covering which the olfactory lobes (paleocortex) and bifurcated hip-
the corpus callosum between the hemispheres and including pocampus (archi~o~ex) interlock bilaterad in the temporal
the septum pe~ucidum; and (2) the Ammon’s horn or “hip- olfa~to-hip~campal junction.
pocampus major” in the medial wall of each temporal lobe The Ammon’s horn formation in Tlmiops truncates is
(Fig. 39). The hippocampal rudiment (“paraterminal body” short and is buried deep in the media1 wall of the temporal
of Elliot Smith, 112,131) arises under the rostrum of the cor- lobe between the chorioid fissure and the parahippocampal
pus callosum (Figs. 39,41B) dorsal to the olfactory lobe from gyms. The two Ammon’s horns diverge from the unci in two
the septal area in the subcallosal gyri (Figs. 40,41A, B) and arcs laterad and dorsad towards the oblique sulci (Fig. 39)
extends over the corpus callosum as the induseum griseum and encompass in their concavities the thalamus as if be-
and taeniae tectae to the fasciolar gyri upon the splenium tween two brackets. The small dentate gyri with fimbriae and
(Figs. 15, 16, 18, 39, 40). The two sulci of the corpus cal- the posterior crura of the fornices converge from the Am-
losum (hemispheric sulci) demarcate the supracallosal hip- mon’s horns dorsad and sharply mediad into a massive hip-
pocampal rudiment (archicortex) bilaterad from the medial pocampal commissure (psalterium) under the splenium (Fig.
border (limbus) of each hemisphere (neocortex). These two 40). As seen in a median section of the forebrain, the bulk of
bilateral longitudinal sulci are continuous anteriorly below the fibers of the fomix pass anterior to the septal (anterior
the knee of the corpus callosum with the posterior parolfac- cerebral) commissure into the septal area and into the area of
tory and the anterior rhinal sulci of the olfactory lobes (Figs. the septo-hypothalamic junction (Figs. 40, 41A, B, 57). The
40, 41A, B) and, posteriorly, below the splenium with the post-commissural fomix to the hypothalamus is small.
hippocampal sulci of the Ammon’s horns (Fig. 39,40,48,49, On the base of the forebrain, between the optic tracts, the
50,51) 52). In Twsiops truncatus the taeniae tectae arise from lozenge-shaped hypothalamic area is greatly foreshortened
the septal area in a common stem between the subcallosal in the antero-posterior diameter from the optic chiasma to
gyri (Figs. 39, 40, 41A, B) from which they are demarcated the posterior perforate space (Fig. 57). The posterior wall of
bilaterally by the postreme parolfa~to~ sulci (Fig. 41B). the in~ndibulum and the floor of the third ventricle (Figs.
Posteriorly the taeniae tectae diverge from the median stem 2A, 34, 41A, B, 57) face the inte~eduncular space from
over the body of the corpus callosum (Fig. 39) as two sepa- which they are separated in the transverse plane by a deep
rate strands of Iongitudin~ fibers. The bifurcated taeniae vertical cleft of the mesencephaiic flexure (Fig. 2A). The
tectae form a slight Ion~tud~aI ridge in the floor of each mammillary bodies are barely recognizable in gross (Fig.
sulcus of the corpus callosum and extend to the splenium 41A) and histological (Fig. 57) preparations. The quotient of
(Fig. 39). Under the splenium they blend with the fimbriae of lengths of the hypothalamus and of the cerebral hemisphere
the dentate gyri. Around the wide spleuium of the corpus has been taken by Griinthal[23,24] and by Pilleri [491 as one
callosum the taeniae tectae thicken into fasciolar gyri of index of “encephalization” of the central nervous system of
Retzius or fasciolae cinereae (Figs. 15, 16, 18,39,40). Below mammals. The quotient falls off from rodents to primates
the splenium the rudimentary dentate gyri form the telen- whereas, in Cetacea, the quotient is in the range of that of
cephalic lips of the chorioidal fissures in the medial walls of primates.
the temporal horns of the lateral ventricles. At the junction The large anterior hypophysis is lodged in a shallow sella
of the sulcus of the corpus callosum with the hippocampal turcica (Fig, 1) and in all species of Cetacea we have exam-
sulcus and in the transition from the retrosplenial hippocam- ined is wide in the latero-lateral dimension and narrow in
pal rudiment (fasciola cinerea) to the hippocampus major antero-posterior diameter (Figs. 42,46). The posterior
(dentate gyrus plus hippocampus proper and subiculum) a hypophysis is situated dorsal to the anterior hypophysis (Fig.
constant torsionfold in the slender parahippocampal gyrus 57) and consists of a transverse bar of neuroglial tissue in a
cuts obliquely across this gyrus from the hippocampal to the meningeal sac completely separated from the anterior
parahippocampal sulcus. We designate the oblique groove hypophysis (Pilleri, [51,521). In a median section through the
formed by this torsionfold as the oblique sulcus (Figs. 15-19, ventricle the cavity of the anterior and dorsal part of the third
39, 40, 47). The hippocampus major (Ammon’s horn) ex- ventricle is filled by a large massa intermedia (Fig. 15-19,
tends in both banks and floor of the hippocampa1 sulcus from 4tA). Anterior to this mass the cavum of Monro (median or
the oblique sulcus to the uncus in which the hip~c~pus rhinencephaIic telocoele) between the columns of the fomix
major (~chi~o~ex) is joined to the olfactory lobe (paleo- communicates posteriorly with the third ventricle (diocoele)
cortex). and, bilaterad through the narrow slit-like foramina of
In Tursiops ~~~~e~~~~(Fig. 42) and in other Cetacea we Monro, with the lateral ventricles (lateral or hemispheric
have studied (Figs. 2629, 45, 46) the uncus is an elaborate telocoeles) as shown in Figs. 34, 35 (also see McFarland et
and conspicuous formation. It projects from the medial wall al., [42]).
of the temporal lobe into the transverse cerebral cleft of From the above description of the rhinencephalon as a
Bichat (Fig. 40) between the temporal pole and the optic median pediment of the paired cerebral hemispheres, it ap-
tract (Figs. 26-29, 40, 41B, 42, 45, 46, 53A, 53B, 54B). The pears reasonable to designate it as a median or rhinic lobe
uncus is grooved by the sulcus semiannularis which sepa- [591. It has clear and constant surface demarcations from the
rates it into two parallel, almond-shaped gyri (Figs. 39, 40, hemispheres proper. Beginning from the septal area as the
42, 47, 54B, 55, 56), the gyrus semilunaris facing the optic primordial region of the endbrain, these surface demarca-
tract mediad and the gyrus ambiens facing the parahip- tions are: (1) the anterior rhinal sulci, (2) the posterior parol-
pocampal gyrus laterad (Figs. 40, 41B, 42, 47, 54B, 55, 56). factory sulci, (3) the sulci of corpus callosum extending to
The latter gyrus is demarcated from the uncus by the the (4) hippocampal sulci and, thence, via the rhinohip-
posterior rhinal sulcus and by the sulcus rhino- pocampal sulci, the latter sulci are joined to the (5) posterior
hippocampalis. The gyrus semilunaris is a club-shaped ex- rhinal sulci which are once again continuous with the an-
tension of the olfactory gyrus (&a). The gyrus ambiens is an terior rhinal sulci back to the septal area, thus closing the
PARALIMBIC CLEFT (ENTOLATERAL Sut_CUS)

n LATERAL SUPERIOR
CRUCIATE FISSURE
SULCUS
i_ATERAL INTERMEOtATE
GYRUS PROREUS SULCUS
LATERAL tNFERiC+4 SULCUS

(ECtiSYLVtAN SULCUS)
GYRUS CC$RONAttS
TRANSVERSE SYLVIAN CLEFT

ORBITAL evdus
LOBULE
DESERT OF BROCA
TRANvERSE TEMPORAL
LA1 ‘ERAL OLFACTORY GYRUS
GYRI
(AREA PRAEPYRIFORMISI
SEMILUNARIS GYRUS
MEDIAL TEMPORAL POLAR
GYRUS
SULCUS SEMILUNAR6
LATERAL TEMPORAL
GYRUS AMBtENS POLAR GYRUS
PARAHIPPOCAMPAL GYRUS ’ \
LATERAL SUPERIOR SULCUS
I
PARAHIPPOCAMPAL SULCUS
FIG. 53A. Direct anterior view of brain of dolphin fTursiops rruncutus) indicating details of the gyri. suki and )&ular format&s of the
orbital and anterior temporal lobes. (approx. I/Zx).
GYRUS CRUCIATE
LIMBIC CLEFT PROREUS SULCUS 5R5lTAL LOBE
LOBULE DkSERT
~OLFACTORY
Lll WBIC LOBE TUBERCLE 1
JUNCTION OF
CRUCIATE SULCUS
GEN IU OF AND LATERAL
CORPUS CALLOSUM ORBITAL SULCUS
ISTHMUS OF TEMPOR AL
SULC :US OF LOBE (FLOOR OF
CORPUS CALLOSUM SYLVIAN CLEFT)
PAROLFAC:TORY AREA
OF SROCA
/ ,TEMPORAL POLAR
MASSA INTERMEOIA
GYRUS
3rd VENTRICLE
END OF
SPLEN IUM OF SULCUS CIRCULARIS
CORPUS CALLOSUM
SULCUS INSULO-
PARAH~F+~CAMPALIS
HIP POCAMPAL
cc >MMlSSURE . AREA DIAGONALIS
PARAHIPPOCAMPAL
OPT IC CHIASM
SULCUS
BYRUS SEMILUNARIS
AQUEDUCT
SULCUS RI-IINALLS
\
POSTERIOR
SU LCUS
V
TELO- DIENCEPHALICUS
/
/ OPTIC TRACT HIPPOCAMPAL YRUS AMBIENS
CEREBRAL PEDUNCLE SULCUS
TENTORIAL SURFACE SULCUS RHINO--WIPPOCAMPALIS
OF TEMPORAL LOBE
FIG. 53B. Antero-medial view of brain of dolphin (Tursiops truncatus) indicating certain gyri, sulci, and lobular formations of the antero-
medial, orbital, and anterior temporal regions of the left hemisphere. This view provides various perspectives not seen in Fig. 53A. (approx.
3/4x).
58 MORGANE. JACOHS ANTI Mt~tAKI.ANI)
PARALIMBIC CLEFT
(ENTOLATERAL SULCUS) SUPERlOw LATERAL SU‘CuS
INTEHMEDIATE LATERAL
SULCUS
CRUCIATE SULCuS
MEDIAL ORBITAL GY ;NFERIOR LATERAL 3UI.CU:,

!FCTOSYLVIAN SUI.CUS)
GYRUS
MEDlAL ORBITA
DISSFCTION
_ATERAL ORBITAL SULCUS

(HERE ORBITAL END
OF CRUCIATE SULCUS) MtDlAL 7EMpORO POLAR
G VRUS
AMBIENT GYRUS
SVLVIAN CLEFT
~TERAL OLFACTORY GYRU
(AREA PRAEPIRIFORMIS
LOBULE DESERT
(AREA TuBERCULARI
TRANMRSE TEMPORAL GYRl

SUL .CUS RHINALIS ANTERIO
CLEFT OF BICH END 0F CIRCULAR SULCUS
SEMILUNAR GVRU OF PARA~I~PP~CAMPAL

SULcUS
~lpp0CAh4PAL SULCU
END OF RHINAL SULCUS
E ‘ARAHIPPOCAMPAL GVRUS
LA1 FRAL TEMPORO POLAR

GYRUS
/
PARAHIPPOCAMPAL SULCUS SULCUS RHINALIS POSTERIOR
FIG. %A. Direct anterior view of brain of dc$hin (Tursiops fruncufus) indicating various gyri, sulci and lobular formations of the anterior.
orbital and temporal regions of the brain. (approx. ‘14x).
Ryrus remiiunarir
circular SUICUS
onteriof
rhino1 sutcur
anterior tronsvor*o
par0lfactory area
ontsrior tronswrse
of Broco
temporot rutcur
loterol olfactory
ircufar sulcur
gyros (are0
praepiriformis)
(posterior ssctor)
olfactory tubarcls
- sulcus insulo-
(lobule dbssrt)
parohippocampalis
diogonol bond
TEMPORO-POLAR
(orso diogonalis)-
optic tract
gyros ombiaor 4
~inrulor *yrur
hippocompa
sutcua (2)
rhino-
‘porohippocompal
hippocompol
rulcus (4)
sulcvs (l-2)
J \
posterior rhino1 sulcur (1) PARAHIPPOCAMPAL GYRUS
FIG. 54B. Antero-baso-medial view of the left cerebral hemisphere in the dolphin (Tursiops ~runcalus) showing the details of the uncal
region, temporo-polar region and orbital lobe. The temporal pole is revealed as the hub of the four cardinal grooves (rhinal, hippocampal,
circular, and parahipp~~pal) as discussed in text and caption for Fig. 40. (approx. 1x).
LIMSLC CLEFT icie4ar uulcur of the inruk, Tm R ‘.tnrulo-prrrahippoccmpol gfoore

hmporo-polar e*tr*mity) (3) OnVS (‘pli de porrag;’ of Esdurepordl
FIG. 55. Photograph of the right cerebral hemisphere of the dolphin(Tursiops fruncurus) in me&ctbasal view as schematized in Fig. 40. Thrs
photograph reveals the ~ste~rn~~ and basal aspect of the oIf&ctory lobe as well as the fo~~ons suKo~ing it in the basaJ region of the
hemisphere. The rhino-hippocampal sulcus, prominent in this specimen, is shown linking the posterior rhinai sulcus with the pit-tile. uncal
end of the hippocampal sulcus and constitutes a groove in the floor of the annectant gynts formed by the mergence of ambient and
~a~ip~~p~ gyri-termed by Broca (6,7j ‘“le pli de passage retr~l~bi~e.” (approx. */.sx).
rhino-hippocampal sulcus Sulcus semiannularis
posterior rhino1 UNCUS

sulcus
para
QYrus
4emilunaris
Iolfactory str ia
medial bank of
the limbic cleft
(parahippocampal Yhippocampus
sulcus)
\ hippocampa I
oblique sulcus
sulcus
dentate gyrus
FIG. 56. Enlarged basal view of dissected temporal extremity of right limbic lobe (parahippocampal gyrus)
and uncal formation. The details of the various sulci in these areas are revealed in this figure. The division of
the uncus into the almond shaped gyrus semilunar-is and gyrus ambiens is shown and the separation of the
gyrus ambiens from the parahippocampal gyrus by the posterior rhinal sulcus is indicated. (approx. 5x).
+ *
taenio tecta
sulcus telc-diencephaiicur \
durb optic chiarm anterior hypophyrir
FIG. 57. Photograph of sag&tally cut section of brain of dolphin (Tursiops truncafus) revealing some details ofthe septal area. hypothalamus.
and pituitary body. Note the large anterior (pre-) commissural fomix system. Note that the posterior hypophysis lies dorsal to and i\
completely separated from the anterior hypophysis. Loyez myelin stain. (approx. 5 X)
\osuAA
ISTHMUS
TEMPORAL
RHINO-HIPPOCAMPAL
SULC u s
FIG. 58. Schematic representation of the sulci which, in continuity
bilaterad, form the rhinic cleft in the cetacean main. The olfactory
lobe (paleocortex) of the rhinic formations is demarcated from the
hemisphere proper by the posterior parolfactory, anterior rhinal.
posterior rhinal and rhino-hippocampal sulci. Abbreviation.
RH S-rhinal sulcue.
PARALIMSK CLEFT LIMBIC CLEFT oval lobule
RHINK CLEFT
@Jlcus of corpur
collosum)
CA .UDATE NUCLEUS
rplcnium of the
corpus callorum
inhrior colliculus
dentote nucleus
lahral Iomnircur
and nucl~r
inferior cereb4ar
peduncl*
subrtantio nigra
trapezoid body
\ \
RIIINK CLEFT (anterior rhino1 rulcur~ sulcus telo-diencephalicur pen,
II
FIG. 59. Photograph of sagittal section of dolphin brain showing the massive, triple-tiered cortical formations corresponding to the three
main lobes of the hemisphere (limbic, paralimbic, and supralimbic). Note the relatively small corpus striatum (putamen and caudate)
compared to the bulky hemisphere. The main mass of the striatum is situated ventral to the thalamus. This figure also illustrates the great
length of the ventrally deflected splenium of the corpus callosum and that it extends caudad to the hippocampal commissure (psaltarium) at
nearly a right angle to the body of the corpus cailosum. Loyez myelin stain. (approx. 0.9~).
ring of cardinal limiting grooves around the circumference of gulates, carnivores and primates) are much less certain and
the hemispheric stalk (Figs. 40,58). Because of the con- present problems which can be resolved only by ontogenetic
stancy, rather than depth, of these uninterrupted cardinal and comparative architectonic studies of the cerebral cortex
grooves delimiting the rhinencephalon from the hemispheres (isocortex). Accordingly, in the present description of the
proper (pallium), we designate the entire complex of grooves surface morphology of the cerebral hemisphere of Tursiops
as the rhinic cleft (Figs. 15, 58). truncatus we shall use a nomenclative schema which is
purely descriptive-anatomical. We will first describe the
III. THE CEREBRAL HEMISPHERES
striatum, the insula, the temporal pole and the orbital lobe of
In Cetacea the morphological distinctiveness of the ex- Tursiops and will then describe the hemispheric mantle (pal-
panded cerebral hemispheres from the rhinencephalon in the lium) as a whole in detail and with its surface subdivisions
median position is explicit. In contrast to the rhinencepha- cleaved by deep longitudinal fissures into discrete lobar for-
lon, which exhibits essentially the same basic structural plan mations.
as in carnivores and primates, the cerebral hemispheres of
I. Striatum
these aquatic mammals exhibit a pattern of surface confg-
uration and morphology which in some respects is radically The corpora striata in Cetacea are relatively small as
different from that of the hemispheres of most land-living compared to the bulky hemispheres (Fig. 59). The main mass
mammals. In the limbus of the hemisphere, nearest the rhinic of the striatum is situated anterior and ventral to the
cleft and rhinencephalon, the departures in surface contig- thalamus. The striatum blends with the gray matter of the
urations from the familiar pattern of land-living mammals are olfactory lobe and is covered by the orbital lobe and, more
the least. They are greatest in the fundus or crown of the laterally, the insula. In the core of the olfactory lobe the
hemisphere, i.e., in the supralimbic parts, farther removed putamen and caudatum are joined into a massive fundus
from the median rhinencephalic pediment of the paired hem- striati (Fig. 60). The tails of the caudate nuclei are short and
ispheres. Here the homologies of the cortical areas of the slender and follow the steep arcs of the lateral ventricles into
hemispheres of Cetacea with the positionally homologous the temporal horns and diverge widely dorso-laterad to the
parts of the hemispheres of the more familiar mammals (un- well-developed amygdalae.
64 MORGANE. JACOBS AND McFARtAN
putaman fundus ttrioti iunction

olfactory tuber&
head of caudote nucleus lantsrior -rhino1 sufcus~
FIG. 60. Photograph of frontal section of dolphin brain (Tur.sio~s trmcxms~ showing the fundus striati in the care of the otfacrory Iohc
Loyez myelin stain. (approx. ? X).
2. Itr.srrln insula into 8-10 anterior and shorter g!yrl and 3--i posteriol
and longer gyri. These radial insular gyri and the s&i he-
The Sylvian cfeft (Figs. 6114-66) opens beneath the oper-
tween them dovetail with the transverse gyri and sulci of the
cufum into a large insular pocket (Figs. 48, 49-52, 6lB)
operculum facing the insula. In the temporal operculum the
which extends in the parasagittal plane from the isthmus of
two anterior of the 4-S transverse opercufar gyri facing the
the temporal lobe to the orbital pole of the hemisphere. The
iflsufa are larger and more prominent and are bounded by
insula in the dolphin (Figs. 48.61B) and in other Cetacea
deeper transverse sulci than the posterior transverse gyri
studied (Figs. 49-52) is large and is elaborately sulcated. It
(Figs. 48-52). The transverse opercutar sulci extend onto the
borders on the olfactory lobe (paleocortex) of the rhinen-
free lateral wail of the hemisphere and radiate from the Syl-
cephalon, buried deep under the massive operculum, and
vian cleft as the orbital, the parietal. and tempof;ll branches
forms the ventrolatera1 wall of the hemisphere. It is clearly of the cleft (Figs. 61A-66).
demarcated from the olfactory stria or gyrus by the anterior
Posteriorly the sulcus circularis duplicates the course of
and posterior rhinal sulci (Fig. 48). A prominent and constant
the rhinal sulcus and turns in a steep arc later+ventrad onto
transverse insular gyrus (Figs. 48, 4%52, 54B) (gyrus lim-
the antero-medial wall of the temporal lobe (Figs. 48. 618.
itans insulae or gyrus Y of Breathnach, 1953, the “limen” or 66). In most specimens it is continuous laterad from the
threshold of the insula) follows the hemispheric bank of the p~ahip~~mpal gyrus with the parahippocampal sulcus
rhinal sulcus and represents the insular limbus of the hemi- and, faterad from the uncus, the transverse insular gyrus
sphere. The insula is inscribed by the buried sulcus circularis blends with the parahippocampal gyrus (Figs. 48. 54B, 55)
(Fig. 48, 4%54B, 61B. 66) which demarcates it from the which represents the temporal limbus of the hemisphere.
opercufum. In Tur.siops truncatus and the other odontocete
Cetacea studied, 1I to 14 radial insular gyri, separated by .+. 7’~~tnportil I’olc trtiti 7‘c,ttll,ctrttCcrllrr (i,bWt.\
sulci, radiate fanwise from the transverse insular gyms
towards the circular sulcus (Figs. 48-548, 6lB). in The blunt and foreshortened temporal pole ot‘ the hemi-
Rulucnopferu phy.sal~~s some 20 radial insular gyri are pres- sphere in Tursiops rrun~urus and other Cetacea examined is
ent (Fig. 52). A deep central suleus of the insula, (Figs. the region of junction of four cardinal limiting sulci (Figs.
48-52) oriented in the plane of the Sylvian cleft, divides the 54B.55). These sulci are: (1) the posterior rhinai SU~CUS. (2)
1’2.~1continued on pas 70
INfERoR LATERAL FISSURE {deft) INTERMEDIATE LATERAL FISSURE INFERIOR LATERAL FISSURE {daft)
(ivprorytvion svlcur) [ectorylvion sulcur)

isetcrylvion sulcus) J
INTERMEDLATE
LATERAL FISSURE
{ruprorylvion rulcu
111161 OPERCULUMw.
r==-
TEMPORAL 01 iRCULUM
INFERIOR LATERAL
FISSURE icisftl
c SYLVIAN CLEFT
Iectervlvicm sulcud
optic n*rv*
cil VJICUI of
FIG. 61A. Left lateral view of brain of dolphin (Tursiops tr~~cut~s~ showing the general co~g~tion of the sulci and gyri on the
dorso-lateral aspect of the hemisphere. Note the opercular gyri forming the opercular lobule surrounding the Sylvian cleft. The course of the
inferior lateral and intermediate lateral fissures are indicated. The pronounced foreshortening and anterior rotation of the forebrain is clearly
evident in this photograph. Also, observe the position of the orbital lobe below the level of the tip of the temporat lobe. (approx. ‘/zX).
,NFER,OR LAiERAL FISSURE C
FIG. 618. Photographs of iaterai view of hemisphere of do~pbin ~Tf~~~~u~~ rr~~cufu~~. Left pbotog~p~ (1) shows intact hemisphere while
right photograph (2) shows a hemisphere with the opercular lobule removed revealing the insula. Note that the circular sulcus of the insula
parallels on the insular surface of the hemisphere the course of the inferior lateral sulcus on the lateral aspect of the hemisphere. (approx.
‘/2x).
66 MORGAN& JACOBS AND ~~~ARL.A~~
INTRUDE LATRRAL FtSSURE INFERfOR LATERAL FISSURE (cleft)

(iupraryfvion rulcu I)
lbJFEWR L&TERM
FiSSURE (cleft)
(+ctorylvion sulcun)
$vlW CLi&FT OLFACTORY L08E
FIG. 62. Left lateral vi,ew of br&n of beluga whale (Delphinapt~ms /mcmi showing the general configu~tio~ of tfre subi and gYri on the
dorso-lateral aspect of the hemisphere. (approx. V4X).
SUPERIOR LATERAL FISSURE (ectolateral sulcus)
INFER !IOR LATERAL

FM iURE (cleft)
(ectoq rlvian sulcus)
SYLVIAN CLEFT OLFACTORY LOBE INFERIOR LATERAL FISSURE (c

:left)
(ectosylvian rulcur)
FIG. 63. Left lateral view of brain of harbor porpoise (Phocaena phocaena) showing the general configuration of the sulci and gyri on the
dorso-lateral aspect of the hemisphere. (approx. 1x).
68 MORGANE, JACOBS AND McFARLAN D
ion
MTERMEDIATE IR 1 LAL
LATERAL FISSURE IRE 4
(suprasyluian sulcurl tion curl
INFERIOR LATettbd
WWRE (cleft1
(rctoorylvian ruicul
/ t
SYLVIAX awl CM&TORY LORE
FIG, 64. Left lateral view of brain of Amazon River dolphin (lnia gcoffrensis) showing the general configuration of the sulci and gyri on the
dorso-lateral aspect of the hemisphere. (approx. %x).
a
u a 4
3
33 a
(* (L .E
.E
4a “0
2
._
”
70 MORGANE, JACOBS AND McFARLAND
sphere covering the insula and parahippocampal gyruh. The

annectant gyrus in the floor of the “pli de passage” of Beau-
regard (insulo-parahippocampal sulcus) expands from this
groove on the temporal pole into a roughly S-shaped gyral
configuration designated here as the tcmporo-polar- pyrus
(Figs. 40, 53A. B. 54A. B. 5).
The sulcus circularis of the insuln I\ continuous ante1o-

ventrad with the lateral orbital sulcus (Fig. 47) which demar-
cates the orbital operculum in the lateral bank of the sulcus
from the lateral orbital gyrus in its medial bank (Figs. 42.47).
The transverse insular gyrus is continuous anteriorly with
the transverse orbital gyrus (Figs. 43. 47, 481. The latter
gyrus borders on the sulcus rhinalic anterior and olfactory
gyrus (stria) and forms the orbital limbus of the hemisphere.
The orbital lobes project antero-ventrad from the transverse
orbital gyri between the orbital opercula and the interhemi-
spheric cleft. Viewed from in front (anteriorly) the orbital
lobes of Cetacea are comparable to two pyramids juxtaposed
in the plane of the interhemispheric cleft (Figs. 9-13) with
their converging apices (orbital poles) oriented ven-
tromediad. The broad bases of the two pyramids blend dor-
sad with the expanded dorsal parts of the hemispheres
ANT PAROLFACi SULCUS (parietal lobes). The orbital lobes in Cetacea are notably
small relative to the prominent and wide olfactory lobes. The
FIG. 66. Schematic representation of the ring-like configuration of contrast with the inverse relation between the olfactory
the limbic cleft in its entirety beginning in the anterior parolfactory lobes (anterior perforate substance) and frontal lobes of pri-
SU~CUS and extending as the splenial and retrosplenial components of
mates and man is striking (Cf. Figs. 25,33).
the limbic cleft into the parahippocampal sulcus and, thence, via the
circular sulcus of the insula. onto the orbital surface of the hemi-
In the medial wall of the orbital lobe the transverse orbital
sphere as the cruciate sulcus. The different components of the limbic gyrus blends with the parolfactory lobule of Broca which
lobe are indicated. The epihippocampal and epistriatal components represents the parolfactory limbus of the hemisphere (Figs.
of the limbic cleft are illustrated. The orbital, parietal and temporal 15-19, 40, 41A, B). The parolfactory lobule of Broca i> de-
components of the operculum are shown. Abbreviations: OP- marcated dorsally by the posterior parolfactory sulcus (Figs.
operculum; Cruc S-cruciate sulcur;; I.at-lateral. 4lA.B) from the subcallosal gyrus (limes hippocampalis
parolfactorius) and anteriorly by the short but constant
the hippocampal sulcus joined in Cetacea by the rhino- anterior parolfactory sulcus (Figs. 41A.B) from the splenial
hippocampal sulcus (Figs. 54B, 55, 58), (3) the circular sul- (superior fornicate or calloso-marginal) gyrus of the “grand
cus of the insula (temporal sector, Fig. 48) and (4) the lobe limbique” of Broca. The anterior parolfactory sulcus
parahippocampal suIcus* joined in Cetacea by a constant varies in prominence and in configuration in different species
connecting groove designated in our figures as the in- of Cetacea (Figs. 15-19). In different mammals its
suloparahippocampal groove (or sulcus) and described by prominence appears to depend on the development of the
Beauregard [2] in Balaenoptera sibbaldii as “pli de passage parolfactory lobule (Figs. 30. 31, 32). In all mammals the
limbique” (Figs. 54.55). A small annectant gyrus is often septal area and the parolfactory lobule constitute the region
present in the floor of the groove between the parahip- from which three long limiting sulci arise and extend in con-
pocampal gyrus and the temporal pole. The posterior rhinal centric arcs onto the medial and onto the basal wall of the
and hippocampal sulci, (Figs. 48, 54B. 58). demarcate the hemisphere. These three fissures are the splenial sulcus. the
periamygdalar limes (threshold) of the rhinencephalon (un- sulcus of the corpus callosum, and the anterior rhinal sulcus.
cus) from the parahippocampal limbus of the hemisphere the latter two sulci joined by the posterior parolfactory sul-
comprising the parahippocampal gyrus and the temporaI sec- cus (Figs. 4lA,B). The pattern of sulcation of the medial wall
tor of the transverse insular gyrus. The circular SUICUS of the of the orbital lobe in Cetacea is remarkably similar to that in
insula (temporal sector) and parahippocampal sulcus (Figs. primates (Fig. 31) and, particularly. to that in man (Fig. 32) if
54B. 55, 66) demarcate the insula and the parahippocampal the frontal lobe of the latter be shorn of the superior frontal
gyrus (limbus of the hemisphere) from the operculum and gyrus (Figs. 32. 41A, B).
from the temporal pole. The latter formations represent thu\ The gyrus proreus extends anteriorly from the Junctron 01
the paralimbic borders of the supralimbic mass of the hemi- the parolfactory lobule and transverse orbital gyrus and
forms the orbito-medial cant of the hemisphere (Figs. 40.
*It should be emphasized that the parahippocampal sulcus in our 4lA. B). Extending from the transverse orbital gyms lateral
terminology forms the external boundary of the parahippocampal and parallel to the gyrus proreus. n variable medial orbital
gyrus of the limbic lobe. Filimonoff and others refer to this sulcus a\ SIJ~CUSor sulcus olfactorius (Figs. 40. 4lA) demarcates the
the rhinal sulcus. We would emphasize that their designation “rhi-
gyrus proreus from the medial orbital gyrus and the orbital
nal” should not be confused with our anterior and posterior rhinal
sulci which have been described in detail in this paper. It also ap-
pole of the hemisphere (Figs. 25-D. 4lA. B, 54.4).
pears that in primate material the sulcus we refer to a~ parahip- The lateral orbital sulcus demarcates the lateral orbital
pocampal is often designated as the collateral sulcus. gyrus from the orbital operculum. This sulcus is variable in
position and is often interrupted by annectant gyri but is cular groove, uninterrupted by any intervening gyri (Fig. 58).
found in all odontocete whales (Figs. 25-28). It could not be We designate this entire circular complex of cardinal sulci as
identified in the mysticete whale Balaenoptera physalus the rhinic cleft, and the rhinencephalic formations demar-
(Fig. 29). As has been described earlier, in most specimens cated bilaterad by these sulci from the hemispheres proper,
the lateral orbital sulcus represents the rostral continuation as the rhinic lobe (Fig. 39).
of the circular sulcus of the insula. It extends onto the an- The cerebral hemispheres in Tursiops truncates and in
terior wall of the orbital lobe and continues dorso-mediad as other Cetacea are invested, as in all mammals, by the corti-
the sulcus cruciatus of Leuret (Figs. 9-13, 66). The sulcus cal plate (mantle, pallium). As seen in Nissl sections, this
cruciatus is found in all Cetacea we have studied. It is a deep plate has sharp clinoid borders along the limbic banks of the
and constant sulcus extending dorsad from the ventro-lateral rhinic cleft which demarcate each hemisphere from the
to the dorso-medial border of the orbital lobe at a sharp angle rhinencephalon (olfactory lobe and hippocampus), the for-
to the interhemispheric cleft sometimes reaching the mations of which are devoid of such a plate.
antero-medial cant of the hemisphere. The sulcus cruciatus The cortical mantle (pallium) of Cetacea is finely sulcated
was designated by Beauregard [2] in Balaenoptera sibbaldii or gyrencephalic in type [47]. The relatively narrow and
as the sulcus of Roland0 of other mammals. On the grounds finely convoluted gyri on the free surfaces of the hemi-
of architectonic evidence (see below) we accept this spheres are crowded against each other between sulci which
homology. are tightly closed and shallow. This impression of shallow
sulci between the narrow, randomly convoluted gyri is dis-
5. Opercular Lobule pelled, however, when the pia is removed and the depths of
the sulci are revealed (Figs. 67,68). The cortical plate and
Dorso-laterad from the cruciate sulcus rises the massive gyral alba are narrow (Fig. 67). The surface area of the cor-
bulk of the hemisphere bisected ventro-laterad by the Syl- tex buried deep in the banks of the sulci greatly exceeds that
vian cleft into an antero-dorsal (parietal) and a larger of the free surface on the crowns of the narrow gyri between
postero-lateral (temporo-occipital) part (Figs. 61A-65). the tightly closed sulci. We have observed three features in
Above the Sylvian cleft the two parts blend without de- the mode of the infolding of the cerebral cortex of Cetacea, as
marcation. The sulcation pattern of the free dorso-lateral compared to that of primates and man, that deserve to be
convexity of the hemisphere, dorso-lateral and posterior to noted: (1) in perfused specimens the banks of the deep sulci
the cruciate sulcus, is characterized by a prevalence of ar- are in tight apposition being separated by thin layers of pia-
cuate longitudinal sulci [32,48] which conform essentially arachnoid without free subarachnoid space; (2) the banks of
with the arc of the hemisphere. Of these roughly concentric sulci meet in the floors of sulci at a sharp angle; and (3) the
arcuate sulci, the deepest and most constant is the inferior T-shaped opercularization of the floor of the sulci, prominent
lateral fissure (Figs. 61A-65), which forms a virtual cleft. in the parietal and occipital lobes of anthropoids and man,
This sulcus duplicates in its course and position on the lateral does not occur anywhere in the deep vertical sulci of the
surface of the hemisphere the course and position of the
cortical mantle of Cetacea.
circular sulcus of the insula under the operculum (Figs. 50, When the intact specimens are hardened in formalin and
61B, 66). The inferior lateral fissure (cleft) corresponds in then dehydrated in alcohols, the sulci are opened and the
Cetacea to the “ectosylvian” fissure of ungulates and cami- orderly pattern of their orientation and disposition on the
vores [47,58]. In all five species of Cetacea we have studied, surface of the brain is revealed (Figs. 4,7, 13, 16, 17, 19,24,
the inferior lateral sulcus demarcates a discrete formation in
29, 46, 49, 50, 51, 52, 62, 63, 65). As a general rule the
the lateral wall of the hemisphere designated here as the
fissures running in antero-posterior (longitudinal) arcs are
opercular lobule (Fig. 61A). longer and deeper than the sulci orientated in transverse
planes [32]. The sulcation becomes finer and the depth of the
IV. THE MANTLE (PALLIUM) OF THE CEREBRAL HEMISPHERES longitudinal fissures increases from the orbital to the occipi-
PROPER
tal lobes (Figs. 59, 67, 68). On the medial walls, above the
Summarizing the foregoing description of the more salient corpus callosum, the gyri are widest and tend to be disposed
features of the surface configurations of the cerebral hemi- in concentric tiers separated by deep fissures (Figs. 15-19,
spheres of Cetacea, special attention must be drawn to the 59). The sulcation becomes finer and shallower and the gyral
clarity and constancy of demarcation between the rhinen- pattern more variable in different species on the free medial
cephalon and the borders of the hemisphere proper. In all 5 borders and on the dorso-lateral and posterior convexities of
species of Cetacea examined the anterior and posterior rhi- the hemispheres (Figs. 61A-65).
nal sulci form together a shallow, yet continuous groove, Two exceptionally deep longitudinal fissures arise as shal-
least variable in its position and course in different species of low sulci in the orbital lobe and extend dorsally and
whales. The identity of this cardinal groove with the anterior Posteriorly. The first of these two cardinal longitudinal fis-
and posterior rhinal sulci of other mammals does not permit sures of the hemisphere proper is the sulcus splenialis
any reasonable doubt. The groove demarcates the olfactory (“scissure limbique” of Broca which is designated the lim-
lobe (paleocortex) from the parolfactory, orbital, insular, bit cleft in this paper) which arises from the parolfactory lob-
and the medial temporal borders of the epistriatal walls of the ule on the medial wall of the orbital lobe (Figs. 15-19, 40,
hemisphere. Similarly, the sulcus of the corpus callosum and 41A, 55,59,66,69A, B, 70). The second cardinal fissure, the
the hippocampal sulcus in Cetacea are also clearly homolo- sulcus confinis of Guldberg or paralimbic cleft (Figs. 15-19,
gous to the cardinal grooves of the same designations in the 4% 59, 69A, B, 70), arises on the anterior wall of the orbital
brains of all other mammals. These two sulci demarcate the lobe and extends posteriorly onto the medial wall parallel to
hippocampus (archicortex) from the pericallosal, retrosplen- the dorsal and posterior free borders of the hemisphere.
ial and parahippocampal borders of the epihippocampal These two deep and constant fissures are outstanding fea-
walls of the hemisphere (Figs. 39, 40, 58). These sulci form tures of the surface configurations of the hemispheres of
with the anterior and posterior rhinal sulci a continuous cir- Tursi0p.r truncatus (Fig. 15) and of the other three odonto-
12 MORGANE. JACOBS AKD MC FARLAND
FIG. 67. Photograph of sagittal brain section in dolphin (Tursiops rruncofrrsl indicating the extreme depth of the sulc~, particularly m rhc
dorsolateral areas. Note the vertical orientation of both the Sylvian fissure and inferior horn of the lateral ventricle. Compare with Rg. 68.
Loyez myelin stain. (approx. 1.2X).
SURFACE ANATOMY OF WHALE BRALN 73
SUPRALiM~lC LOBE SUPRALIMBIC LOBE
LIMBIC LOBE SUPRALIMBIC L&E
FIG. 68. Phot~gmph of medial aspect of rig& cerebral hemisphere of the dolphin ~T~rsiups f~u~c~~u~i. The dorsal posterior parts of the
paralimbic lobe (oval lobule and lingual lobule) have been removed exposing the medial wall of the supralimbic lobe of the hemisphere. Note
the extraordinary depth of the parahmbic cleft and the long, closely spaced radial gyri regutariy disposed along its lateral bank. The amount
of neocortex buried in the depths of the cfefts, fissures and sulci in all the Cetacea we have examined exceeds that on the exposed surface of
the hemispheres. This free cortical surface consists of the crowns of very slender gyri whose close spacing is associated with tightly closed
sulci (Cf. Fig. 47). (approx. IX).
MORGANE, JACOBS AND McFARtAND
CALCAR_INE SULCUS0)
_IMBIC
:FT
‘PARkbH~F”~AL SULCUS
-\,‘~~
SULCUS
I RH~INALIS POSTERIOR
PAROLFACI-ORY LOBULE OF BROCA
INFUNDfBULUM
OPTIC CHIASM
RCULUM OLFACI-ORIUM
‘LATERAL OLFACTORY GYRUS
SULCC JS PAROLFACTORIUS POSTERIOR SULCUS RHlNALlS ANTERIOR

Sl JLCUS PAROLFACTORIUS ANTERIOR
FIG. 69A. Interpretive drawing of the medial wall of the right hemisphere in the dowin f7ir~siups tru~ufus/ showing the major S&I
(clefts), gyri, and lobular formations. Note especially the continuity of the deep clefts (limbic and paralimbic) separating. respectively. the
limbi~~imbic and p~~imbic~sup~imbic l&es.
INFEE
MARGlNAL
EXTERNAL
INTERCALATE -
I_,MI~C GYRUS
INTERMEDIATE
(NTERCALATE --
LIMB~C GYRUS
INTERNAL /
INTERCALATE
UMBIC GYRUS
FIG. 69B. Interpretive drawing of the medial wail of the right hemisphere in the dolphin (Ttirsiops fruncarusj showing the major sulci (clefts),
gyri, and lobular formations. This figure shows the various intercalate sulci and gyri of the limbic and paralimbic lobes (Cf. Fig. 70).
MORGANE. JACOBS AND Mt~FARI.ANU
inferior bank of porolimbic cfefi OVAL COBWE
rnfeiics book of
porokmbic cleft
-CALC AWNE SULCUS ( ? )
>LI~uGUAL IX)IIULE
of limbic RE‘rROSPLLNTAl LOMILE

OF LIMBIC LOBE
of limbic clef ,HIPPOCAMPAL SUlCUS
SUPRACALLWAL LO&WE
OF LlMBtC lo&
EXTERNAL MTSRCALATE
tturiltt GWWS
~N~~R~~CT~S~A~Al~ \ limbir honk‘ of rhinic cl& GYhUS SEMILUNARlS

IN
~~~~ATE
FIG. 70.Photograph of dissected Iimbic (below) and paralimbic (above) lobes (minus gyrus proreus) and uncw. In the paralimhic i&c the
oval lobule and its boundaries are well shown as is the Iinguai lobde. fn the limbic lobe the three intercalate gyri comprising the supracaltosai
sector are indicated. Note the narrowing of this lobe in the retrospienial area to form the retrosplenial lobule of the limb& Me. In the
temporal sector the limbic lobe continues as the parahippocampal gyrus beyond the oblique sulcus. (appnw 1. 1
cete Cetacea used in the present study (Figs. 16, 17, 18). 66) and consists of two sectors: ii dorso-medial or parahip-
These two fissures were also found in 3u~~en~~te~~ p~~~~f~~~~~ pocampal se.ctor on the medial wall of the hemisphere, ithove
(Fig. 19), although in this mysticete whale the fissural pattern and around the corpus callosum (Figs. 15-19. 69A, B, 701,
of the hemispheres presents many departures in details from and a ventrolateral or orbito-insular sector (Figs. 48,661.
thaf of the odontocete Cetacea. Because of the exceptional The parahippocampal sector of the limbic cleft arises on
depth of these two fissures (Fig. 68), we sha21 hereinafter the medial wall of the orbital lobe from the anterior parolfac-
refer to them, respectively, as the limbic and the paralimbic tory sulcus (Figs. 418, Bt 66. 69A. 70) as a shallow. and in
clefts. They subdivide the hemispheres into three arcuate some species discontinuous, sufcus interrupted by annectant
lobar formations-the limbic, the paralimbic, and the supra- gyri and extends dorsad as the splenial sulcus of Krueg or
limbic lobes, rising in tiers above the corpus callosum “scissure limbique” of Broca (cingular of caIloso-marginal
(Figs. 15-19, 59, 69A, B, 70). In Cetacea the orbital lobe fissure of primates) (Fig. 66). Above the corpus callosum,
wedged between the parolfactory lobule and the insula ap- the “scissure limbique” extends towards the splenium and.
pears to be a common hub from which these Iobar formations gaining in depth, gives off at nearly right angles two short
of the hemisphere expand dorsad and posteriorly over the branches, an ascending one towards the dorsal cant of the
rhinic lobe (Fig. 59). We shall describe the surface config- hemisphere, the superior marginal sulcus, and. slightly
urations of these three lobar formations of the cetacean posteriorly, a shallower and shorter descending branch, the
cerebral hemispheres and define their boundaries. inferior margitial sulcus, oriented ventrad towards the srkus
of the corpus callosum (Figs. 15. 16, 17. 698). The cleft
continues posteriorly and ventrad around the splenium as the
1. titnbir Cleft uad Limbic L&c retrosplenial sulcus towards the oblique SU~CUSof the
The limbic cleft forms a nearly closed ring of continuous par~~ppocamp~ gyrus {Fig. 66). From the oblique SU~US
grooves concentric with the limbus of the hemisphere (Fig. the Limbic cleft coniinues onto the haso-medial wall of the
hemisphere as the parahippocampal sulcus to the pole of the intercalate gyri (Fig. 70). The external intercalate sulcus
temporal lobe, where the fissure meets the temporal end of branches off from the suprasplenial sulcus of the limbic cleft
the circular sulcus of the insula in the sulcus insulo- near its origin from the anterior parolfactory sulcus (Figs.
parahippocampalis over a short annectant gyrus (“pli de 15,69B). The latter is the small limiting sulcus of the parol-
passage limbique” of Beauregard, Figs. 54B, 55, 66). The factory lobule or area and in Tursiops truncatus does not
parahippocampal, i.e., the dorso-medial sector of the limbic communicate with the sulcus of the corpus callosum (rhinic
cleft, demarcates the dorso-medial, epihippocampal limbus cleft). In Delphinupterus leucas (Fig. 16) the supracallosal
of the hemisphere bordering on the archicortical sector of the lobule exhibits only a single long axial sulcus bordered and
rhinic lobe (hippocampus, fasciola cinerea, supracallosal enclosed at each end by two intercalate gyri. Also, in this
stria [taeniae tectae] induseum griseum, gyrus subcallosus) species the anterior parolfactory sulcus is deep and com-
from the olfactory lobe to the “pli de passage” of municates with the sulcus of the corpus callosum (Fig. 16).
Beauregard (sulcus insulo-parahippocampalis). This sector In Phocuenu phocuenu (Fig. 17) a single intercalate sulcus is
of the limbic lobe as here described corresponds to the pallial present. It arises from the deep anterior parolfactory sulcus
(neocortical) part of the “grand lobe limbique” of Broca which also opens into the sulcus of the corpus callosum. In
(parolfactory lobule, the splenial or “superior fornicate” Znia geoffrensis (Fig. 18) the pattern of sulcation resembles
gyrus, the retrosplenial gyrus, and the parahippocampal or more that of Tursiops truncutus than that of Delphinupterus
“inferior fornicate” gyrus and uncus). The hippocampus leucus or Phocuenu phocuenu. In Znia two intercalate sulci
represents the rhinencephalic “archicortical” part of the are present: the suprasplenial sulcus of the limbic cleft and
“grand lobe limbique,” a term conceived by Broca in 1878, the short internal intercalate sulcus. They are, however,
before the tectogenetic and architectonic differences be- broken in this species by prominent annectant gyri. The
tween the hippocampus and the hemisphere had been clearly internal intercalate sulcus arises from the anterior parolfac-
defined [8, 10, 16, 531. tory sulcus which does not communicate with the sulcus of
The circular sulcus of the insula (Figs. 48, 49, 50, 51, 52, the corpus callosum (Fig. 18).
53,54B, 55,61B, 66) represents the orbito-insular orepistriat- In Buluenopteru physulus (Fig. 19) the pattern of sulca-
al sector of the limbic cleft and may be regarded as the con- tion presents a distinct departure from that seen in the odon-
tinuation of the epihippocampal sector of the cleft on the tocetes. It has some resemblance to that in Zniu geojfiensis.
lateral wall of the hemisphere. The circular sulcus demar- In Baluenoptera physulus the suprasplenial sulcus of the
cates the orbito-insular or epistriatal sector of the hemi- limbic cleft is interrupted by a massive annectant gyral for-
spheric limbus bordering on the olfactory lobe (paleocortex) mation which blends with the orbital lobe mediad to the
and covering the striatum from the operculum. The entire cruciate sulcus (Figs. 13,19). The external intercalate limbic
complex of constant grooves (Fig. 66) begins with the shal- sulcus branches from the suprasplenial sulcus near the lat-
low anterior parolfactory sulcus on the medial wall of the ter’s origin from the anterior parolfactory sulcus and joins
orbital lobe, and extends posteriorly as a continuous and, in the internal intercalate sulcus above the splenium and,
Cetacea, exceptionally deep “scissure limbique” of Broca thence, extends to the oblique sulcus as a continuous groove
(splenial, retrosplenial, and parahippocampal fissures) to the (Fig. 19). The internal and intermediate intercalate gyri of the
temporal pole. It then continues as the circular sulcus under supracallosal lobule form in Buluenoptera physulus two
the operculum to the lateral orbital sulcus and demarcates the overlapping crescents lodged within the concavity of the
entire circumference of the limbus of the hemisphere around massive external intercalate gyrus. The inferior marginal
the hemispheric stalk. In Cetacea the limbus of the hemi- sulcus is not present (Fig. 19) and the supracallosal lobule is
sphere represents a discrete lobar formation and we have thus not demarcated from the retrosplenial sector of the
designated this entire paramedian zone of the hemisphere as limbic lobe.
the limbic lobe. In Tursiops truncutus, Delphinupterus leucus, and
The limbic lobe, bounded by the rhinic and limbic clefts, Phocaenu phocaenu the retrosplenial lobule of the limbic
is contoured by the branches of the latter and is subdivided lobe is clearly demarcated by the inferior marginal and the
into five lobules: (1) the parolfactory, (2) supracallosal, oblique sulci. This lobule is best developed in Tursiops trun-
(3) retrosplenial, (4) parahippocampal lobules of the dorso- cutus and is grooved by an inter&ate axial sulcus (Figs. 15,
medial limbus, and (5) the insular lobule of the ventrolateral 69B, 70). An intercalate sulcus is also present in Baluenop-
limbus of the hemisphere. The parolfactory lobule and the teru physulus (Fig. 19). No such intercalate sulcus is present
insular lobule are joined by the transverse orbital gyrus, thus inDelphinupterus leucus (Fig. 16), Phocuenu phocuenu (Fig.
closing the ring of the limbic formations (Figs. 40, 55, 66, 17), and Zniu geoffrensis (Fig. 18). In Delphinupterus leucas
69A, B). (Fig. 16) and in Zniu geofjensis (Fig. 18) the retrosplenial
In the five species of Cetacea we have studied the parol- lobule is a slender, narrow and smooth gyrus which blends
factory lobule on the medial wall of the orbital lobe and the with the parahippocampal gyrus.
parahippocampal lobule (gyrus) on the baso-medial wall of
the temporal lobe exhibit the least variation in size and sur-
2. Puralimhic Cleft and Puralimbic Lobe
face configuration. The insular lobule of the limbus is more
variable as to size and number of radial sulci that groove it in The second and deepest of the two cardinal longitudinal
different species (Figs. 48, 49, 50, 51, 52). The supracallosal fissures in the wall of the hemisphere of Cetacea is concen-
and the retrosplenial lobules exhibit the widest variations in tric with the dorsomedial (epihippocampal) sector of the
surface configurations. In Tursiops truncatus the supracal- limbic cleft (Figs. 9, 15, 16, 17, 18, 19, 20, 68, 69A, B, 71).
losal lobule of the limbic lobe (Fig. 40). extends from the We designate the fissure as the paralimbic cleft and the large
anterior parolfactory sulcus to the inferior marginal sulcus gyral formation intercalated between the limbic and
and is grooved by two intercalate axial sulci, the internal and paralimbic cleft on the medial wall of the hemisphere, as the
external (Figs. 15, 69B), which subdivide the lobule into paralimbic lobe (Fig. 69A, B, 70). The cleft arises in all 5
three gyri, the internal, the intermediate, and the external cetacean species in this study in a shallow and constant
the orbital lobe and expands posteriorly from the gyrus pro-
reus (Figs. 40, 4lA, B. 55. 72B) to the superior marginal
sulcus. The lobule is demarcated from the orbital operculum
by the cruciate sulcus and from the supracallosal lobule of
the limbic lobe by the splenial sulcus (limbic cleft): (2) the
oval lobule, demarcated anteriorly and ventrally. respec-
tively, by the superior marginal sulcus and retrosplenial
sulci, is bounded dorsally and posteriorly by the paralimbic
cleft and its recurrent branch (Figs. 15, 69, 70. 71): and (3)
the lingual lobule comprises the third division of the
paralimbic lobe and is wedged between the recurrent branch
(calcarine sulcus) and the posterior extension of the
paralimbic cleft on the baso-medial wall of the temporal lobe
(Figs. 40, 55. 69A, B, 71. 72A, B).
The configurations and boundaneb of the three subdivi-
sions of the parahmbic lobe are explicit in Tcrr.siop.ctrwtmtu~
(Figs. 15,70) and in Phocuencr phm u~n~t (Fig. 17). They arc
somewhat less clear but recognizable in DC,lplti,lNptc,rlc.\
/cccws (Fig. 16). In Iniu ,qcwjJrvt~ri~ (Fig. 18) and in
Bdtrcnopterct physalus (Fig. 19) the surface configurations of
the paralimbic lobe depart the farthest from the standard
described in Twsiops trunwtu.\. The variations of the sur-
face configurations in the paralimbic lobe are determined bq
PARALIMBIC CLEFT (Configuration)
the variations in the positions and prominence of the limiting
FIG. 71. Schematic representation of the sulci which form the
and intercalate sulci. In Inio gmffrcncis and in Bnlac~noptcrcr
paralimbic cleft (solid lines) in the cetacean brain. The limbic cleft is physdus the superior marginal and the recurrent sulci could
indicated by dashed lines for orientation. not be identified with certainty. In Inicl gec~:frcrr.si.\ and
Bduenopterct physalus no oval lobule as such is defined
since we cannot locate either its anterior (superior marginal
T-shaped sulcal configuration (Figs. 9-13, 48) on the sulcus) or posterior (calcarine sulcus) boundaries. In the oval
antero-dorsal surface of the hemisphere, some distance dor- lobule of Ttrr.siop.struncutus (Figs. IS. 69A, B. 70), Drlphiw
sad from the orbital pole, mediad from the end of the cruciate trptertts 1euc~1.s(Fig. 16). and Phocl~wcc phoc~cr~~nn (Fig. 17) a
sulcus and about 2 cm laterad from the medial, free border or single intetcalate axial sulcus is present,
cant of the hemisphere. The long stem of the T, designated Because of the great depth and antero-posterior extent of
by Guldberg [2S] as the entolateral sulcus (or fissure), and by the parahmbic cleft (Figs. 68, 72A, B) the cleft and the mas-
Kiikenthal and Ziehen [34] as the sulcus confinis (Fig. 48). sive and elaborate gyral formations intercalated between it
extends dorso-mediad in a broad arc nearly parallel to the arc and the limbic cleft represent together a unique feature of the
of the dorsal cant of the hemisphere and continues surface configuration of the medial wall of the cetacean
posteriorly and ventrad onto the baso-medial wall of the hemisphere not found, to our knowledge, in the hemispheres
temporal lobe (Figs. 15, 16, 17, 18, 19,20,21,22,23, 24,40. of any other mammal. The posterior horn of the lateral ven-
69A, B, 70,71,72A). From about the point at which the cleft tricle is absent or greatly reduced in Cetacea (Fig. 35 and
cuts the dorsal cant of the hemisphere, it gives off mediad. at McFarland, Morgane, Jacobs. 1421. The occipital lobe ih
a sharp angle open posteriorly, a deep and long recurrent rounded and the blunt occipital pole appears shifted in
branch (labelled calcarine sulcus and qualified with a ques- Cetacea mediad and towards, rather than posterior (i.e..
tion mark in Figs. IS, 16, 17,40,69A, B, 70). The main and away from), the splenium as in primates. The occipital pole
deeper branch of the paralimbic cleft, designated the sulcus corresponds in Tursiops trunc utu.5 to the oval and lingual
entolateralis or entolateral fissure (25, 32, 341, extends lobules separated by the recurrent fissure (Figs. 15. 69A. B.
posteriorly and ventrad in a broad arc concentric with the 71). This fissure appears to arise virtually as a shallow sulcus
recurrent or calcarine sulcus (Fig. 69A, B, 70, 71) and, be- from the external (paralimbic) lip of the retrosplenial fissure
coming shallow, terminates near the limbic cleft at the level (limbic cleft). Growing deep it extends in a loop dorsad and
of the oblique sulcus (Fig. 40, 69A, B, 71). The superior anteriorly to open into the paralimbic cleft and so appears as
marginal branch of the limbic cleft cuts into the paralimbic a branch of the latter (Fig. 71). The cytoarchitectonic evi.
lobe and demarcates the anterior paralimbic lobule from the dence. to be detailed further elsewhere, seems to support
characteristic oval-shaped and finely sulcated, rosette-like this inference. For purposes of this paper some of this evi-
gyral formation which we designate as the oval lobule of the dence is given below and represents our preliminary ar-
paralimbic lobe (Figs. 40, 69A, B, 70, 71). The recurrent chitectural assessments.
branch of the paralimbic cleft demarcates the club-shaped Anteriorly, in the paralimbic wall of the gyrus proreus
posterior lobule (lingual lobule) of the paralimbic lobe, be- (blocked field A. Fig. 73 and Fig. 73A) the cortical Plate is
tween the posterior (entolateral) and the retrosplenial sulcus wide, the outer granular layer is dense, and the third layer
(limbic cleft), from the oval lobule. Thus the paralimbic lobe, is wide and clear, containing mostly small and medium
intercalated between the limbic and massive supralimbic pyramids and is nearly devoid of granule cells. The pyramids
lobes, is clearly demarcated by the superior marginal and the rapidly gain in size in the fifth layer while the sixth layer is
recurrent fissures into three discrete lobular formations poorly demarcated from the subcortical alba. This Cortex is.
(Figs. 40,69A, B, 70.71): (1) The anterior paralimbic lobule, thus, an agranular macropyramidal cortex. Further dorsad.
narrow anteriorly, occupies the antero-medial free border of in the medial wall of the anterior paralimbic lobule (blocked
! S:YIcontinued on page XX
OVAL LQBULE PARIETAL. LOBE PARALIMBIC CLEFT

1
LOBE
superior
marginal rulcus~
inferior
marginal sulcus :
LINGUAL LOBULE
LIMBICCLEFT
(ruprarplsniol
rulcur)
PARALIMBIC CLEFT
,TEMPORAL LOBE
RHINIC CLEFT {rulcur of corpus callorum) \ calcorine sulcus ? \ LIMBIC CLEFT (parahippocompal sulcur)
\ (branch of paralimbic cleft) .
PARAHlPPOCAMPAL GYRUS
OLFACTORY TUBERCLE
(LOBULE DkSERT)
FIG. 72A. Postero-medial view of right cerebral hemisphere in the dolphin (Tursiops truncatusj illustrating the posterior course of the
paralimbic cleft. Note the tongue-shaped lobular formation (lingual lobule) defined by the paralimbic cleft and what we term the calcarine
fissure. The broad arc of the paralimbic cleft extending posteriorly and ventrad does not break into the limbic cleft at its termination (See
Figs. 40, 69A, B, 71). The banks of the paralimbic cleft and of the calcarine sulcus have been separated artificially to emphasize their great
depth. (approx. )/4x).
80 MORGANE, JACOBS AND McFAKLAND
OVAL LOBULE,
: CLb 1
GYRUS CR~EUS\
LATERAL QlltarpILt G=US I. L ORBITAL o?ucuLuM
FIG. 72B. Baso-medial-posterior view of right cerebral hemisphere in the dolphin (Tursiop~ rrunc~~rus~illustrating certain details of the
paralimbic lobe and orbital area. (approx. %x).
SULCUS ENTOLATERALIS
(Paralimbic cleft)
SUPERIOR MARGINAL
SULCUS
ULCUS SPLENIALIS
SWLCUS, TELO-DIENCEPHALICUS
FIG. 73. Schema of parasagittal section of brain of Z’ursiops truncatus 4.4 mm from the midsagittal plane showing locations of the sequence
of cellular field (Boxes A-F) studied architectonically and illustrated in Figs. 73A, B, C, D, E and F. Major sulci labelled at this level indicate
the lobar and lobular region with which each cortical sample is associated.
FIG. 73A. Histologicat section through tha cerebml cortex of the bottlenuse dofphirx in the pmdimbiz wail &ii& gyr-us prureua~t t3ox ii, l”‘ig.
73). Note the width of the cortical plate (layers IL-WI, dense,outer granular layer, wide less dense third layer cirntaining mttstly small and
medium sized pyramids, labge pyramids in the fifth layer, and a sixth layer containing small and medium size &fk of varying sh&e which is
pooriy demarcated from the subcortical ah. The cytoarchitecture af this fidd is th:it -crfan agran~~lar. ~i~e~~~~~~-~~?~~~~
corC.kS type. Nisd
stain, 100x
83
FIG. 73B. Histological section through the cerebral cortex of the dolphin in the medial wall of the anterior pkalimbic lobule further dorsad
than Figure 73A (Box B, Fig. 73), Although the cortical plate is somewhat wider and more celhdar, especially layer III, than in Fig. 73A the
architectonic pattern is essentially the same, Also, taller pyramidal cells with long apical dendrites appear in cksters in the fifth Iayer. khis
cortex is strongly suggestive of the “motor” agranular gigantapyramidal type. Nissl stain, 100x.
FIG. 73C. Histologicai section through the cerebral cortex of the dolphin at a location (Box C, Fig. 73) that IS sub more dorsad and postcmr
in the anterior paralimbic lobule than is Fig. 73B. In this cortical region, very near the superior marginal sulcus. the cortical cellular pattern
changes drastically. Here, the somewhat narrower cortical plate is far more ceflular than anteriorly in the pardhmbic lobe (Fig. 734 and
contains predominantly smah ceils which give to this cortical region a definite granular appearance. The large pyramids have disappeared rn
the deep layers except for a few medium sized triangular cells. Such an architectonic pattern suggests a highly pranu)ar sensory type cortex.
Nissl stain, 100x.
8.5
FIG. 73D. Histological section through the cerebral cortex of the dolphin in the medial wall of the paralimbic lobe behind the superior
marginal sulcus in the oval lobule (Box D, Fig. 73). In this region the cortical plate is narrower than in anterior paralimbic fields, the width of
the Iamina zonahs (molecular layer) has become greatly reduced, and the pyramids of the third layer in Figs. 73A, B and C have become
replaced by densely distributed, small granule cells. In the fifth layer a clearing is present bordered superficially by a narrow, sir&e layer of
large multipolar, dark staining cells. This cortical pattern is reminiscent of the modified special heterotypical granulose isocortex (koniocor-
tex) of the occipital lobe of other mammals. Nissl stain, 100X.
86 MORGANE, JACOBS AND WFARLAND
FIG. 73E. Histological section through the cerebial cortex of the dolphin shawing cortical organization at the antenor end of the superlot
surface of the lingual lobule of the pa~imb~c lobe (Box E, Fig. 731. The architectu~~ pattern is quite similar to that of Fig. 73D. Niscl stain,
100X
FIG. 73F. Histological section through the cerebral cortex of the dolphin showing cortical organization at the posterior surface of the lingual
lobule of the paralimbic lobe (Box F, Fig. 73). The architectural pattern is essentially unchanged from that seen in Figs. 73D and E and is
strongly suggestive of a modified special heterotypical granular type of isocortex (koniocortex). Nissl stain, 100x.
field B, Fig. 73 and Fig. 73B) the same architectonic pattern of its surface configuration and sulcal patterns In different
persists. However, the cortical plate (layers II-VI or cellular species (Figs. 9. 10, II, 12. 13,20.21.22,23. 24.61A. 62.63.
region minus layer I) is somewhat wider and more cellular, 64, 65). In contrast to the paralimbic lobe, its surface is rela-
especially layer III. Slightly larger and taller pyramidal cells tively homogenous and presents no deep sulci (Figs. 9. 10.
with long apical dendrites appear in clusters in the fifth layer. II. 12, ]3,20.21,22.23,24,61A,62.63.64.65).Anteriorly.
Even though the general appearance of the cortical plate is the supralimbic lobe is wedged between the sulcus cruciatus
different from that in the cat, dog and primates, the pattern is and the orbital operculum (Figs. 9. 10. I I. I?. 13,61A. 62.63.
strongly suggestive of the “motor” agranular giganto- 64,65, 75). It expands from the orbital lobe dorsolaterad and
pyramidal type of cortex. Still further dorsad and posteriorly in an arc to the temporal lobe, between the oper-
posteriorly, near the superior marginal sulcus, but still in the cular lobule and the paralimbic lobe (Figs. 75.76). In it%
anterior paralimbic lobule (blocked field C, Fig. 73), the ap- topographic position in the globoid hemisphere of Cetacea
pearance of the cortical plate changes radically (Fig. 730. the supralimbic lobe corresponds primarily to the free sur-
The cortical plate narrows slightly, becomes distinctly granu- faces of the parietal and temporal lobes of the hemisphere of
lar in all layers, the large pyramids disappear in the deep primates.
layers except for a few medium sized triangular cells, and the In Tursiops the supralimbic lobe. bounded by the
demarcation of the cortical plate from the alba becomes paralimbic cleft and by the inferior lateral (ectosylvian) fis-
sharper than in Figs. 73A and B. Such an architectonic pat- sures, is subdivided (Figs. 61A. 62. 63. 64. 65) by two
tern suggests a highly granular, sensory (somesthetic) cor- variable fissures-the intermediate lateral (suprasylvian) and
tex. Beyond the superior marginal sulcus, in the oval lobule the superior lateral fissures (or sulci) into three arcuatc
(blocked field D, Fig. 73), and in the lingual lobule (blocked lobule-the inferior, the intermediate and the superior sup-
fields E and F, Fig. 73 and Figs. 73E and 73F). the architec- ralimbic lobules (Figs. 75,76). These supralimbic lobules and
tonic physiognomy of the cortex assumes a distinctive and sulci exhibit the widest variations in position on the convex-
altogether special appearance. The cortical plate narrows. ity of the hemisphere and in configuration in different spe-
the width of the lamina zonalis (molecular layer) becomes cies, in different specimens of the same species. and even in
greatly reduced, the pyramids of the third layer become re- the right and left hemisphere of the same specimen (Figs. 9.
placed by densely distributed, small granule cells, while in IO, 11, 12. 13. 20. 21. 22. 23. 24. 6lA. 62. 63, 64. 6.51.
the fifth layer a clearing appears bordered by a narrow. The superior lateral fissure is designated variously in
single layer of large multi-polar, dark staining cells. Although different whale species as the lateral and as the ectolateral
granularity of this cortex is not as apparent as in other mam- fissure [25,47,55]. The fissure appears to begin in the orbital
mals, the pattern is reminiscent of a modified special lobe near the medial cant of the hemisphere (Figs. 15. 16. 17.
heterotypical granulose isocortex or “koniocortex” of the 18, 19, 75, 76) either from (or lateral to) the paralimbic cleft
occipital lobes of other mammals. or sulcus confinis 125,551. In its posterior course it parallels
Posteriorly, on the baso-medial wall of the temporal lobe, the paralimbic cleft (sulcus entolateralis) from which it de-
the lingual lobule is not sharply demarcated from the baso- flects gradually laterad, onto the hasolatera] surface of the
lateral and anterior parts of the temporal lobe (Figs. 72A,B). temporal lobe (Figs. 75.76). It i:, a discontinuous groove,
The architectonic pattern of the cortex of the lingual lobule interrupted by the intervening gyri which convey IO it a
in the bank of the paralimbic cleft (Fig. 73E) is similar in “zig-zag” course.
many ways to that seen in the wide expanse of the cortex in The intermediate lateral (SUpr’dSyiVian) sulcus appears in
the baso-lateral wall. This latter type of cortex is shown in some specimens of Tursiops trwxxr/ur bmin to arise in the
Fig. 74A. Here the cortex is very narrow and is also granu- orbital lobe above and extends in a broad arc more or less
lar, and the fifth layer contains large multi-polar cells. How- parallel to the ectolateral (superior lateral) fissure, onto the
ever, in the anterior transverse temporal gyrus the cortex lateral surface of the temporal lobe below and behind the
assumes a different appearance (Fig. 74B). It is more densely posterior (temporal) part of the inferior lateral fissure (Figs.
cellular, more granular, and exhibits a clearing in the deep 61A, 75, 76). It is resolved on the lateral wall of the temporal
cell zone in which smaller and fewer pyramidal and multi- lobe in the network of shallow sulci.
polar cells are present. Following the cortical plate from the The opercular lobule, sharply demarcated from the in-
lingua1 lobule (Fig. 72) onto the baso-lateral and supero- ferior supralimbic lobule by the constant inferior lateral fis-
medial walls of the temporal lobe one has, nevertheless, sure, a virtual deep cleft, appears in Cetacea as a discrete
the impression that the architectonic areal differentiation is cortical formation, distinct from the supralimbic lobe (Figs.
greatly attenuated between the baso-medial (lingual lobule) 61A. 62, 63. 64, 65. 75). It caps the Sylvian cleft and is
and the baso-lateral and superior temporal cortical regions of grooved by the transverse sulci which radiate from the Syl-
the hemisphere. It may be that nearly all the temporal lobe vian cleft. The inferior lateral fissure duplicates in its arcuate
baso-laterad from the foreshortened and blunt occipital pole course and position on the lateral wall of the hemisphere the
to the temporal pole represents an extension, across the course and position of the circular sulcus beneath the oper-
paralimbic cleft, of a type of modified acoustic analyzer area. culum (Figs. 48,61B).
The cytoarchitectonic pattern of the cortex anterior to the
Sylvian cleft is that of a special heterotypical cortex of granu-
lar somatomotor type with large, squat pyramids in the deep
The wide expanse of the free dorso-lateral and posterior layer (Fig. 74C). In the posterior (temporal) bank of the Syl-
convexity of the hemisphere, bounded mediad by the vian cleft, the cortex is narrow and more granular, with scat-
paralimbic cleft from the paralimbic lobe and, laterad, by the tered small pyramids in the deep layer (Fig. 74D). Thus. the
inferior lateral (ectosylvian) fissure from the opercular lobule entire circumference of the arcuate supralimbic lobe is bor-
is designated as the supralimbic lobe (Figs. 20-24, 61A, 62, dered by the heterotypical orbital, paralimbic, and opercular
63,64,65,68,69A, B, 75). This largest of the three divisions cortex which is predominantly granular posterior to the
of the cetacean hemisphere exhibits the greatest variability plane of the Sylvian cleft.
FIG. 74. Schema of right posterior quadrant of horizontal section through brain of Tursiops truncates at a
depth of 58.0 mm from the dorsalmost surface. Shown are the locations of cortical fields in the opercular
region and the basolateral wall of the temporal lobe (Boxes A-D) that are illustrated in Figs. 74A, B, C and D.
Labelled circular sulcus and Sylvian cleft indicate the topographic relationships of these fields to the insula.
FIG. 74A. Histological section through the cerebral cortex of the dolphin in the posterior lateral wall of the hemisphere (Box A, Fig. 74~.
Note the following cytoarchitectonic similarities of the cortex in this region to that of the lingual lobule shown in Figs. 73E and F: quite
considerable narrowing of the cortex, overall granularity of cells of the cortical plate and presence of large multipolar cells in the fifth layer.
Nissl stain, 100x.
FIG. 74B. Histological section through the cerebral cortex of the dolphin in the anterior transverse temporal gyrus (Box B, Fig. 74). In
contrast to the cortex in the posterolateral wall of the hemisphere (Fig. 74A), the cortex here is wider and exhibits an increased cell density
that further emphasizes the overall granular appearance of the cortical plate. Moreover, fewer large pyramidal and multipolar cells are
present in the deeper layers. Nissl stain, 100x
FIG. 74C. Histological section through dolphin cerebral cortex of opercular lobule anterior to the Sylvian clcf~. I-he cortrcal platc ha\ an
overall granular appearance but is not as densely cellular as in granular paralimbic sectors (Cf. Figs. 73C and 7311). The fifth corttcal layer
contains scattered, squat, large and intermediate size pyramids present superficial to a relative Lone of clearing which separate\ the lifth
from the sixth cortical layer. This cytoarchitectonic pattern represents that of a special heterotypical cortex of R granular sorn~l~~rn~~t~r type.
Nissl stain. 100x
FIG. 7483. ~isto[ogicaI section through dolphin cerebral cortex in the temporal part of the opercular lobule along the posterior bank of the
Sylvian cleft. As compared to the operculum anteriorty (Fig. 74C), the posterior opercular cortex is slightly narrower, far more granular with
small cells predominating, and contains only small pyramids scattered in the deep layer. This is a heterotypical cortex of granular type. Nissl
stain, 100x.
MORGANE, JACOBS AND Mcl;ARtAND
\
\\
-____--
SURFACE ANATOMY OF WHALE BRAIN 9.5
PARALIMUIC CLEFT
(ENTOLATERAL FISSURE; Suws CON~~NIS) SUPERIOR LATERAL FlSSuffE (RCTO~ATERAL sumis)
SW< :lJSCRUCIATUS
1INFFRIOR LATIIRA1
SULCUS
SUPRALIMBIC LOUE
ORBITAL OPiRCUlUM SYiVlAN CLEFT TEMPORAL LO5E
FIG. 76. Photograph of the left cerebral hemisphere of the dolphin (Tursiops rruncatus) orientated in the dorso-frontal and lateral direction to
illustrate the main mass of the supralimbic lobe and some of the principal sulci, fissures and clefts. The massive dorsolateral and posterior
expansion of the sup~imbic lobe is indicated in this view. The sup~imbic lobe in Cetacea corresponds primarily to the free surfaces of the
parietal and temporal lobes of the hemisphere of primates, though its homology with the latter lobes of primates is doubtful. (approx. r/z x).
96 MORGANE, JACOBS~AND MCFARLAND
ANTERIOR ECTOSYLWAN SYWAN CLEFT

POST-SYLVIAN SULCUS
STERlOR ECTOSYLVIAN GYRUS
&ZTOSYlVtAN SUlCUS
FK. 77A. Drawing of brain of the pig (lateral view) indicating the main gyri and sulci in the left hemisphcrt-
S~A~YlVlAN SUWJS
SY MAN GYRUS
.ECT~~~~tAN ~~A~YlVlAN GY RUS
\ w,-*__I .’ / SULCUS ECTOLATERALIS
ECTUfYlVtAN SULCUS
I
OLFACTORY
BULB
ANTERIOR
7
RHtNAl FISSURE
PYRIFORM LOBE
DOG BRAIN
FIG. 7713. Drawing of brain of the dog (lateral view) indicating the main gyri and zulci in the left hrmisphetv
The supralimbic lobe above this paralimbic fringe repre- relative to the greatly increased size of the hemispheres,
sents the fundus of the hemisphere. The architectonic pat- while the olfactory lobes became conspicuously reduced in
tern of the cortical plate in the fundus changes gradually size.
from the heterotypical patterns in the inferior supralimbic In Cetacea it is the olfactory lobe that has maintained
lobule nearest the opercular lobule (Fig. 78A), and from the better its relative size than the hippocampus which, con-
superior supralimbic lobule nearest the paralimbic lobe (Fig. trariwise, has become greatly reduced in size. When compar-
78C) to an increasingly uniform, homogeneous pattern of ing relative sizes of the hippocampus and cerebral hemi-
homotypical cortex of the intermediate supralimbic lobule spheres, the Cetacea would be placed on the level with pri-
(Fig. 78B). mates. With regard to relative sizes of the olfactory lobes
and hemispheres, however, they would be placed on a level
with that of ungulates and carnivores. In this conflict of ap-
pearances, it is noteworthy that the rule of inverse relations
DISCUSSION
between relative sizes of the rhinencephalon and hemisphere
The surface configurations of the telencephalon of is not breached if separate bases of comparison are used in
Cetacea present all the general features of the mammalian Cetacea, one which compares the size of the olfactory lobe
brain, the basic plan being essentially the same. However, relative to the epistriatal part of the hemisphere, and the
there are considerable extrinsic and intrinsic peculiarities of other which compares the size of the hippocampus relative
the cetacean brain and it departs from the surface configura- to the epihippocampal part of the hemisphere. The epistriatal
tion of land-living mammals in many respects. In certain or frontal lobes, large in primates, are reduced in Cetacea to
aspects the brain of the bottlenose dolphin and other Cetacea small “orbital” lobes and overlie relatively large olfactory
follow the patterns found in Camivora, in others it resembles lobes, which are greatly reduced in primates. The epihip-
the sub-human primate brain, while in still other features it pocampal part (parietal, occipital and temporal lobes) which
approximates, and even exceeds in development and promi- are, in Cetacea, proportionately even larger than in primates,
nence, certain features of the human brain. It has a median overlies a greatly reduced hippocampus.
rhinencephalon and paired hemispheres joined by a massive When the olfactory lobes and epistriatal part of the hemi-
corpus callosum and covered by the convoluted gray mantle spheres (frontal lobes in Primates, orbital lobes in Cetacea)
of the cerebral cortex (palhum). However, beyond these and the hippocampus and epihippocampal part of the hemi-
general features, the configurations of the brain of these sphere (parietal, temporal, and occipital lobes) are envisaged
aquatic mammals present considerable extrinsic and intrinsic as morphologically distinct formations, then in Cetacea it is
peculiarities and depart from those of land-living mammals in the epihippocampal part that seems to represent the
many significant details. These departures are subtle in the phylogenetically most advanced part of the telencephalon,
rhinencephalic (median) zone and are more prominent in the while the epistriatal part has remained on a lower rung of the
limbic (paramedian) zones. The departures from the general scale of phyletic evolution. From this it seems to follow that
plan become outspoken in the dorsolateral and posterior the “encephalization” of the nervous system of the exclu-
zones of the fundus of the hemispheres and exhibit widest sively aquatic Cetacea expressed itself in the course of evo-
interspecies as well as considerable individual variations lution in a great elaboration of the epihippocampal portion of
within the same species of Cetacea. the cerebral hemispheres at the expense of the epistriatal
From the comparative study of the general configuration portion which, as compared to that of land-living mammals,
and relative proportions of the ventral, or orbital, part of the has lagged in development.
hemisphere of the dolphin, it appears rather evident that this “Encephalization” of Cetacea seemingly has taken a
orbital extremity of the hemisphere corresponds at most to different course in evolution than in land-living mammals
the inferior or orbital part of the frontal lobe of carnivores, and was apparently dictated by differences in the evolution
primates, and man shorn of the “prefrontal” part. This ros- of the physical conditions of the environment to which they
tral extremity of the hemisphere of the bottlenose dolphin became adapted. These environmental conditions empha-
and other cetaceans is thus greatly reduced when compared sized the development of special sensory inputs of stimuli-
to the hemisphere of man, anthropoid apes, and even that of physical agents and of the special cerebral apparatus for
carnivores and ungulates (fig. 8). The foreshortening and differentially analyzing the information provided by the
blunting of the occipital pole of the cerebral hemisphere and stimuli-signals of these physical agents. At the same time,
the absence of the occipital horn of the lateral ventricle (Fig. these environmental conditions de-emphasized the devel-
34) suggest that this most dorsal and caudal part of the dol- opment of the cortical apparatus concerned with somatic
phin brain is developed according to a plan different from responses to the stimuli-signals. Such a view may raise ob-
that which determined the form of the occipital lobe in the jections on the grounds of the apparently great versatility of
brain of land-living mammals, particularly primates and man. Cetacea in their mode of performance in water. This ver-
It is generally accepted that the size of the rhinencephalon satility is achieved, however, with a minimum inventory of
(as originally defined by Turner, [57]) and the size of the effector organs (skeletal musculature and joints) present in a
hemispheres (pallium) exhibit an inverse relation in the as- strongly axiate, streamlined body form that is almost devoid
cending series of mammals from monotremes, marsupials, of useful extremities.
and rodents to primates and man-the larger the hemi- The epistriatal part of the cerebral hemispheres develop,
spheres, the smaller the rhinencephalon. This statement, in mammalian phylogeny and ontogeny, upon the olfactory
however, holds true mainly with regard to the size of the lobes as their pediment in intimate structural relation to the
olfactory lobes or paleocortical division of the rhinencepha- hypothalamus and viscerosomatic motor apparatus (effector
ion [28]. The hippocampus (archicortical division of the organs) of the head regions and forelimbs. The epihippocam-
rhinencephalon) follows less closely the rule of reduction of pal part of the hemispheres, overlying the hippocampus as
its size with the increase in size of the hemisphere [29]. In their pediment, develop in an intimate structural and con-
primates, including man, the hippocampus maintains its size nective relation to the thalamus and sensory analyzer appa-
98
SUPERIOR LATERAL SULC CLEFT
INTERMEDIATE
SULCUS
(Fissure ectosyh4a)
SYLVIAN CLEFi
i
_J
_’
,i’
‘,._.,
FIG. 78. Schema of frontal section of brain of Tur.riops ~rumx~us at a level 37.8 mm caudai to anterior brain surface ~~ati~su~~~b~c
cortical areas (Boxes A-C) illustrated in Figs. 78A, B and C.-Major sulci are labeled to indicate r&ion on the dorso-later’&wait oi’ the
hemisphere (supraiimbic lobe) with which each cortical sample is associated.
ratus (receptor organs) of the head region (sensory Broca on the dorsomedial limbus of the hemisphere is a for-
placodes). mation distinct from the hippocampus and is continuous with
The surface demarcation of the rhinencephalon from the the insula and the orbital limbus of the hemisphere. The
paired hemispheres and of cleavage of each hemisphere into paralimbic lobe on the medial wall and the opercular lobule
three concentric tiers, the limbic, the paralimbic and the sup- on the lateral wall, respectively, overlie and opercularize the
ralimbic, are marked in the cetacean brain and are not seen limbic lobe as we propose to redefine it here. The similarity
in any other mammalian brain. As noted, the paralimbic for- of the architectonic pattern of the heterotypical cortex in the
mation appears to represent a cardinal feature of the ceta- anterior (orbital) part of the paralimbic lobe (Fig. 73B) and in
cean brain. This segregation of the paralimbic lobe in the the opercular lobule (Fig. 74C) suggests that the former may
cetacean brain is represented schematically in Fig. 79. The be the area of cortical representation of the flippers, trunk,
pattern in the primate brain is shown for comparison in Fig. and flukes, while the opercular lobule may be the area of
80. The segregation of the special heterotypical cortex of cortical represenation of the trigeminal (beak) and glos-
motor, tactile, and visual types in the cetacean paralimbic sopharyngeal (blowhole) innervation. The opercular lobule
lobe between the arcuate longitudinal clefts which separate may be regarded as representing on the lateral wall of the
this formation from the limbic and supralimbic divisions of hemisphere the lateral couriterpart of the paralimbic lobe on
the hemisphere is, so far as we have been able to determine, the medial wall.
not found in any other mammal. These special heterotypical The limbic lobe of Cetacea is small in relative surface area
areas in primates are contiguous with the general heterotypi- as compared to the great surface area of the supralimbic part
cal cortex of the limbus of the hemisphere and are, in gen- of the hemisphere. According to Filimonoff (171, the surface
eral. separated from each other by a wide expanse of area of the limbic lobe (mesocortex) of Defphinrts ddphis
homotypical cortex (association cortex) present in the su- represents only 0.4% of the telencephalic surface as against
pralimbic fundus of the hemispheres. In Cetacea, however, 1.3% in man, 2.0% in monkeys, 2.7% in dog, 7.1% in kan-
these intimately thalamus-bound special heterotypical areas garoo, and 17.6% in hedgehog. The surface area of the hemi-
remain on the dorsomedial wall, and all border on the sphere (neocortex), outside the limbic lobe, is 97.9% in the
epihippocampal limbus of the hemisphere. They do not ex- dolphin as against 95.9% in man, 93.3% in monkeys, 84.2% in
tend onto the supralimbic fundus of the hemisphere. dog, 62.2% in kangaroo, and 32.4% in hedgehog.
All three tiers or lobes of the hemisphere arise in the It is significant that the parahippocampal lobule of the
orbital lobe as their common hub and expand posteiioi-ly limbic lobe in Tursiops and in the other cetacean brains
over the hippocampus to converge upon the temporal pole of examined is, in sharp contrast to the pitiform lobe of cami-
the hemisphere (Figs. 15.59). The “grand lobe limbique” of vores and the hippocampal gyrus of primates, greatly re-
‘rc‘cxtcontinued on page 101
99
FIG. 78A. Histological section through the cerebral cortex of the dolphin on the superior lateral wall of the fundus of the hemisphere.
Cortical field is located in the inferior suprahmbic lobule very near the opercular lobule (Cf. Fig. 78, Box A and Fig. 61). The cortical plate is
densely cethtlar and contains predomin~tly small cells. The fifth cortical layer containing sparsely scattered large pyramidal and multipolar
cells is indistinct owing to their admixture with small cells. This represents a heterotypical type of cortex. Nissl stain, 100x.
FIG. 788. Histological section through the cerebral cortex of the dolphin on the superior lateral wall of the hemisphere (fundus) III the
intermediate supralimbic lobule. Observe the decreased cell density, especially in the third layer, and the relative distinctness of all cortical
layers as compared to the cortex in the inferior supralimbic lobule (Fig. 78A). Layer five contains scattered clusters of large and intermediate
size pyramidal cells, and a discontinuous clearing zone is present at the junction of layers V-VI. This cytoarchitectonic pattern is that of a
homotypical cortex and may be regarded as the closest approximation in the cetacean brain of the “associationid” cortex of the primate
hemisphere. Nissl stain, IOOx
101
FIG. 78C. Histological section through the cerebral cortex of the dolphin on the superior lateral wall of the hemisphere (fundus) in the
superior supralimbic lobule nearest the paralimbic lobe. Approaching the paralimbic lobe, the cortex of the superior supralimbic lobule
exhibits a more granular appearance and the layers of its cortical plate are less easily separable than in the homotypical cortex present in the
intermediate supralimbic lobule (Fig. 78B). The cellular pattern here is that of a heterotypical cortex. Nissi stain, 100X.
DOLPHIN FOREBRAiN
D0RC;OtA-i ERAL ZONE

(LATERAL TELENCEPHALON,
MEDIAN ZONE
(MEDIAN TELENCEPHALONI
INTERCAL.ATE ZONE
‘(INTERCALATE TELENCEPHALON)
PARAMEDlAN LONE
(PARAMEDIAN TELENCEPHALONI’
FIG. 79. Schema, based on our present studies, to illustrate the basic organization of the dolphin forebram. Note that in the dolphin
forebrain the parahmbic lobe is interposed between the supralimbic lobe and the limbic lobe (Cf. Fig. 80). Three major zones of the
telencephalon can be defined (Yakovlev, 1968): (1) the median rhinencephalon or rhinic lobe (telencephalon medium or impar) which is
unpaired and lies between the hemispheres and includes the septal area, the paleocortex of the olfactory lobes and the archicortex of the
hippocampus; (2) the partially paired limbic lobes or telencephalon semipar; and (3) the completely paired supralimbic lobes or telencepha-
Ion totopar. The above morphogenetic definitions of the three zones of the telencephalon (impar. semipar and totopar) are based on
abundant evidence of differences in the tectogenesis. ultimate cyto- and myeloarchitecture and in the pattern of organization of each zone
with the corresponding tectogenetic zones of the thalamus. To our knowledge the separation of the forebrain zones into an additional lobe.
the paralimbic. is unique among mammals to the cetacean brain. Abbreviations: Hypoth-hypothalamus; hypoph-hypophysis.
duced below the oblique sulcus and rolled inward with the challenging problem. We can, however, venture the follow-
hippocampus deep into the transverse cleft of Bichat. while ing general conjectures. The topographic relations of the
the basal paralimbic wall of the temporal lobe is, on the most anterior part of the paralimbic lobe to the sulcus
contrary, greatly expanded. All of the cetacean brains we cruciatus and gyrus proreus, between which sulcus and
have examined have revealed with almost textbook clarity, gyrus it blends with the orbital lobe, suggest possible homol-
clear-cut development of the rhinic and limbic lobes with the ogy of this anterior part to the anterior cruciate gyrus and
features of these two lobes being generally constant (but with proreus of the dog’s frontal lobe and, therefore. to the
individual variations). The position and surface confuura- somato-motor area of the agranular macropyramidal frontal
tions of the limbic lobe of Broca in the medial wall of the cortex of primates. In addition, the extraordinary depth and
dolphin hemisphere are unmistakably comparable to those of position of the posterior extension (entolateral fissure) of the
the gyrus fomicatus (superior limbic or cingular, and parahip- parahmbic cleft and its recurrent branch on the posterior and
pocampal or inferior limbic gyri) of the cerebral hemisphere basomedial wall of the hemisphere suggest that in Cetacea
in primates and man. There are, however, some notable this cleft may be comparable to a greatly modified parieto-
peculiarities of these medial limbic formations in the dolphin occipital fissure present in the medial wall of the primate
brain, e.g., the anterior (supracallosal) part of the limbic lobe hemisphere, while its recurrent branch may represent the
in the dolphin is much larger than the superior limbic (cingu- calcarine fissure of primates. Such an interpretation implies,
lar) gyrus in carnivores and primates. of course, that there occurred a profound evolutionary mod-
Because of the great depth and antero-posterior extent of ification of the topographic pattern of the posterior wall of
the paralimbic cleft (Figs. 68,72), this cleft and the massive the cetacean hemisphere and, correspondingly, a shift in the
and elaborate gyral formations intercalated between it and position of the occipital pole in Cetacea as compared to pri-
the limbic cleft represent together a feature peculiar to the mates. Based on our architectonic evidence the paralimbic
surface configuration of the medial wall of the cetacean lobe, intercalated into the anterior, medial and basomedial
hemisphere, not found, to our knowledge, in the hemi- wall of the hemisphere, may represent the special heterotyp-
spheres of any other terrestrial or aquatic mammal. The ical cortex of the motor and some somatic sensory analyzer
homology of this great fissure and of the lobar formation areas, segregated into a discrete lohar formation between the
which it cleaves off from the supralimbic lobe presents a limhic lobe and the free dorsolateral, posterior and basolat-
PRIMATE FOREBRAIN
DORSOLATERAL ZONE
RHINIC LOBE
STRIATUM
LiMBlC LOBE SEMIPAR
TELENCEPHALON
SUPRALIMBIC TOTOPAR I + 11
LOBE
FIG. 80. Schema to illustrate basic organization of the primate forebrain (Cf. dolphin forebrain, Fig. 79). Note the basic organization of
rhinic lobe, limbic lobe and supralimbic lobe. Abbreviations: Hy~th-hypothalamus; hypoph-hypophysis.
erd convexities of the hemisphere. It is of great interest that the region of the recurrent branch of the paralimbic cleft.
these cortical areas of the paralimbic lobe border the epihip- Thus, the paralimbic cleft and its recurrent branch in
pocampal sector of the limbic lobe and extend in an antero- Cetacea may then be comparable to the parieto-occipital
posterior sequence from the orbital to the occipital poles. cleft and calcarine sulcus of primates.
Summarizing our description of the paralimbic lobe, the The foreshortening and blunting of the occipital pole of
two outstanding features of this formation are: (1) its con- the cerebral hemisphere and the absence of the occipital
stancy, in terms of existence as a distinct lobar formation horn of the lateral ventricle in Cetacea suggest that this most
separated by deep clefts from the limbic and supralimbic dorsal and caudal part of the dolphin brain is developed ac-
lobes in the four species of odontocete Cetacea and in the cording to a plan different from that of the occipital lobe of
mysticete, Balaenoptera physalus; and (2) the great variabil- the brain of terrestrial mammals, particularly primates and
ity in the pattern of surface configurations of its anterior, man. In this same regard, it is important to point out some of
intermediate, and posterior parts in different representatives the evidence by which the homology of the recurrent branch
of the 5 species studied. In all Cetacea the paralimbic lobe of the paralimbic cleft in Cetacea with the calcarine fissure in
appears as a direct dorsal and posterior extension of the anthropoids was established. In part this was based on de-
orbital lobe, intercalated between the limbic lobe on the me- scriptive anatomical relationships showing the sulcal branch
dial wall and the main mass of the hemisphere on the free of the paralimbic cleft cutting into the posterior part of the
dorsolateral wall. The architectonic evidence suggests that paralimbic lobe in a recurrent movement and terminating
the paralimbic lobe represents the so-called motor, somes- near the limbic cleft without, however, establishing con-
thetic, and visual projection areas segregated in Cetacea into tinuity with it (Figs. 15, 17, 32, 40, 69A, B). The other
a discrete longitudinal lobar formation which, posteriorly in grounds on which we accepted the designation of this sulcus
the massive temporal lobe, is expanded into the acoustic as the “calcarine sulcus” will be discussed in a future paper
projection area without a sharp architectonic demarcation of dealing with the architectonic considerations of the paralim-
the latter from the visual area. It is suggested that in Cetacea bit lobe. Turner [57] remarked that it is the calcarine sulcus
the occipital lobe and visual cortex have been greatly mod- which “determines the presence of an occipital lobe,” but in
ified and shifted onto the medial wall, so that the part of the the case of the present paper it is more convenient to desig-
lobe co~esponding to the occipital pole has come to occupy nate the lobular area to which the visual radiations project by
104 MORGANE. JAC‘OHS AND Mc F4RL.4Nl)
the term “oval lobule” and establish its more posterior patterns in the supralimbic lobe5 01’ the cetacean brains
boundary as the calcarine sulcus. Surface homologies are examined. The variation in lobular patterns of the supralim-
best established by the relationships these structures have bit lobe among the brains in the present study is rather IX-
with the underlying ventricular system (McFarland CQ~1.. markable but obviously reflects the neocortex (cortical man-
[42]) and to neighboring brain regions, as well as to their tle) as being the most plastic substrate of adaptive behavior
precocious appearance in ontogeny, and to their constancy even among the different cetacean species. It has been of
under varying conditions in mammals which have diverged special value. however, to separate descriptions of the sur-
widely from a primitive stock (Elliot Smith, [Ill). Elliot face features of the cetacean brain into the two major
Smith presents strong evidence that the calcarine fissure and taxonomic groups of Cetacea. Odontoceti and Mysticeti.
the retrosplenial part of the splenial sulcus are identical. He This has been necessary because mysticete (baleen) whales
points out that the splenial sulcus is the most constant and were the group originally discussed in the literature by
precocious neopallial furrow in the mesial wall of the hemi- Beauregard [2]. Guldberg 1251, Kiikenthal and Ziehen 1341
sphere in the Chiroptera, Camivom, Ungulata, and the and Rawitz [55l and from which the descriptive terminology
Cetacea and that the calcarine fissure in primates develops in for the cetacean brain has been developed. It is, therefore. of
a position which exactly corresponds to that occupied by the great value to compare our baleen whale fH&~,ro~~~r~
splenial sulcus in other mammals and at the same epoch in phy.srr1u.s) with those described by these authors. In such ;I
development. The present studies indicate that on primtr comparison we have generally agreed with the configura-
firck evidence of disposition and reciprocal relations of the tions established in these earlier studies. and we have clearly
oval and lingual lobules, these two formations of the identified in our specimen of Halc~ot~~tptrrtr pl~~~.utlus the
paralimbic lobe in Cetacea seemingly correspond, respec- superior lateral, intermediate lateral. and inferior lateral gyri.
tively, to the cuneus and the lingual gyrus of the occipital On the other hand. the odontocete brains have other specific
lobe in anthropoid apes and man. differences in their gyral and sulcal patterns and are best
In particular, the surface contigurations of the supralim- compared with the study of Kojima 1321 who described the
bit zone of the cerebral hemispheres in Cetacea exhibit con- brain of the sperm whale, which is the largest of odontocete
siderable differences as compared to their configurations in brains. It is of special interest to note that the entolateral
other mammals. This is not surprising since the cortical sulcus of conventional cetacean nomenclature I?. 25, 34) ap-
mantle (neocortex) is the most plastic substrate of adaptive pears to have no furrow homologous to that in the brain of
evolution of mammals. We have identified homotypical cor- land-living and non-cetacean aquatic mammals. The topo-
tex mainly on the dorsolateral convexity of the hemisphere graphic ambiguity of the term “entolateral” and the chasmal
in the intermediate lobule of the supralimbic lobe (Fig. 78B). depth of this furrow (Figs. 48. 49, SO. 511 caused u4 to desig-
The cortex of the intermediate supralimbic lobule of the nate it as the “paralimbic” cleft. rhis latter designation
cetacean brain may be regarded as the closest approximation bears upon a particularly important aspect of the fundamen-
to the “associational” cortical areas of the primate hemi- tal arrangement of the cetacean brain, namely. the separa-
sphere. The transition from the heterotypical cortex fringing tion of primary projection areas into distinct lobules entireI)
the anterior (orbital), medial (paralimbic), and lateral (oper- separated from surrounding association areas by chasmal
cular) borders of the supralimbic lobe to the homotypical clefts. In the hemispheres of non-placental mammals the
cortex on its dorsal convexity is gradual. The cortex is pre- primary projection areas overlap and occupy almost the
dominantly agranular in type anterior to the plane of the entire convexity as well as the basomedial wall of the hemi-
Sylvian cleft (epistriatal cortex) and predominantly small cel- sphere. However. in placental ungulates, carnivores, and
led (granular) behind this plane (epihippocampal cortex). primates these areas of special heterotypical cortex become
The homology of the suprahmbic lobe of the cetacean increasingly separated from each other by the expansion of
brain with the parietal and temporal lobes of primates is areas of homotypical isocortex intervening between and sur-
doubtful. It is not demarcated by a transverse Rolandic (cen- rounding them. Nevertheless, these special hetrrotypical
tral) fissure such as dominates and defines the configuration areas never lose their topographic contiguity with the general
of the free dorsolateral surface of the frontal and parietal heterotypical isocortex (mesocortex) of the paramedian zone
lobes of the primate hemisphere. The cetacean supralimbic bordering on the hippocampus. In Cetacea, however, the
lobe resembles somewhat the longitudinal gyri on the dorsal primary projection areas are formed into distinct lobules.
convexity of the hemisphere of ungulates and carnivores. each clearly separated from the lateral convexities (associa-
Hence, their nomenclature [33] has been transferred to tion areas) of the hemispheres by deep clefts. This pattern ot
Cetacea [25,34]. The individual variability of sulcation in the arrangement is unique to Cetacea and contrasts sharply with
supralimbic lobe of Cetacea is comparable to the variability the surface arrangements of the primary projection areas and
of the individual pattern of sulcation in the so-called “aso- association areas found in the brains of terrestrial and other
ciational” areas of the convexity of the human hemisphere. aquatic mammals such as Pinnipedia (seals, sea-lions. wal-
In the case of the supralimbic lobe, the intermediate lat- ruses) and Sirenia (manatees, dugongs. sea-cows). Thus.
eral (suprasylvian) and the superior lateral (ectolateral) lon- three basic distribution patterns of isocortical areas exist
gitudinal sulci are extremely variable among the cetacean amongst Mammalia. In non-placental mammals. the cortical
species we have studied and vary from specimen to speci- pattern in the supralimbic zone is a confluence of association
men in the same species, from species to species and even in and primary projection zones. In primates and carnivores the
the two hemispheres of the same specimen (Figs. 1, 10, 11, primary projection zones exist as islands surrounded by
12. 13, 21, 22, 24, 48, 53A, 54A, 61A. 62, 63, 64. 65). The association cortex. A third pattern may exist only in Cetacca
variability of position, of continuity, and of configuration of in which there is a unique separation of the primary prqjec-
these two sulci is reflected in the wide inconsistencies to be tion zones from association areas. Thus. in the dolphin the
found in their labeling in extant illustrations of closely re- somesthetic, visual, and acoustic cortex develop within. and
lated cetacean brains IS, 12, 32, 341. In addition, there is a form special individual lobules of the paralimbic lobe. It fol-
great deal of variation in practically all the gyral and fissural lows then that the supralimbic region may have expanded
entirely outside of the primary receptor zones in the ceta- tion of afferent paths from the peripheral sensory receptors
cean brain. It is of special interest that primates have a very of mainly the head region. The cortical projection areas of
reduced paralimbic zone as compared to the dolphin where these recipient surfaces of the body (cortical analyzers)
most of the analyzers appear to be arranged serially in the differentiate in mammalian embryos in the paramedian zone
paralimbic zone surrounding the limbic gyrus. In the dolphin of the cerebral hemisphere near and about the hippocampus.
it then appears that the areas above the limbic cleft are zones The great isocortical expansion and proplasia occurring in
of special elaboration, and, as emphasized, a most interest- the dorsolateral zone of the hemisphere in primates and man
ing feature is the intercalation of these zones by the deep may be regarded as the evolutionary product of a progressive
fissures of the limbic and paralimbic clefts between the lim- elaboration and differentiation of the cortical ends of sensory
bit and supralimbic lobes. It is also noteworthy that in the analyzers of these areas. The so-called “primary” projection
dolphin the primary projection areas appear to occupy a rel- areas extend onto the convexity of the hemisphere from the
atively greater surface area of the hemisphere than in the heterotypical isocortex of the paramedian zone and exhibit a
human and anthropoid apes, while the “association areas” special heterotypical structure such as enhancement of the
develop as an entirely special and separate supralimbic lobe. granular layers, or, in the opposite direction, reduction of the
In one respect, the primary projection areas in the dolphin granular layers and enhancement of the pyramidal cell
are generally like those of Carnivora in disposition, i.e., layers. These special heterotypical areas exhibit konio-
are paramedian, but are, as indicated, much more greatly granulose or giganto-pyramidal cytoarchitectonic features.
elaborated. As has been stated previously, in the hemispheres of non-
The opercular lobule about the Sylvian cleft appears to be placental mammals these areas overlap; they occupy almost
intercalate cortex between the insula and the supralimbic the entire convexity as well as the basomedial wall of the
lobe. It will be of special interest to determine if this is a hemisphere. However, in placental ungulates, carnivores
paralimbic zone related to the paralimbic areas on the medial and primates these areas become increasingly separated
wall of the hemisphere. Such determinations involve cyto- from each other by the expansion of areas of homotypical
architectural analyses and, ultimately, a study of the isocortex intervening between and surrounding them. Nev-
thalamus and its projection systems. Similarly, the inferior ertheless, these special heterotypical areas never lose their
supralimbic lobule (ectosylvian gyrus of conventional usage) topographic contiguity with the general heterotypical isocor-
is demarcated on the lateral convexity of the supralimbic tex (mesocortex) of the paramedian zone bordering on the
lobe by the deep inferior lateral sulcus and may be regarded hippocampus.
as a distinct formation transitional from the supralimbic lobe It is interesting to speculate why the primary projection
to the orbital extremity of the paralimbic lobe. areas of the dolphin are so huge and form specific lobules in
In sharp contrast to the dorsal and caudal part of the their own right. As noted, the dolphin exhibits huge dorso-
hemisphere is the massive proplasia of the parietal and tem- lateral parietal and temporal expansions of the hemispheres.
poral regions which, in the dolphin’s brain, occupy almost It will require a study of the thalamic projection systems to
the entire expanse of the lateral and basal wall of the hemi- decide whether these supralimbic areas are “association
sphere. The massiveness of these regions and the promi- areas” that have not elaborated concentrically, but rather
nence of the sella media and of the inferior horn of the lateral “outside” the primary projection zones.
ventricle (Fig. 34) suggest a special development of these On the basis of our studies we have concluded that the
parts which correspond most closely to and exceed in rela- lobular formations in the dolphin brain are organized in a
tive proportions, the temporal and parietal lobes in carnivora pattern fundamentally different from that seen in the brains
and primates, including man. Of the 5-6 transverse temporal of primates or carnivores. This fundamental difference in
gyri facing the 3-4 posterior radial insular gyri, which cut basic arrangement makes certain of the homologies ex-
across the superomedial wall of the temporal lobe and extend tremely tenuous. These latter will have to be established by
onto the free lateral wall of this lobe, two are especially cytoarchitectural studies, comparative studies of the
prominent and are bounded by deeper transverse sulci. The thalamocortical relations, and ultimately by electro-
configuration and topographic position of these two large physiological mapping of the sensory zones of the cortex.
transverse gyri indicate a possible homology with the
anterior transverse temporal gyri (Heschl’s convolution) of
ACKNOWLEDGMENTS
the superior plane of the temporal lobe of primates.
It is generally accepted that the elaboration of the isocor- The authors wish to thank Prudence Carter and Marcelina Obaya
tex (neocortex, neopallium) was guided in the evolution of for the bioillustration work. We also thank Dr. Paul I. Yakovlev for
the mammalian endbrain by the convergence and concentra- his many helpful suggestions during the course of this project.
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The Anatomy of the Brain of the Bottlenose

Dolphin (Tursiops trunca . Surface tus)

Configurations of the Telencephalon of the

Worcester Foundation for Experimental Biology, Shrewsbury, MA,

Received 30 November 1979

‘Supported by NSF grant BNS 77-08660.

rostrum of corpus callosum Chop

BOD; AXIS / ah& I m&encephalic flsxure

LATERAL ORBITAL -SULCUS SYLVIAN CLEFT

CALLITHRIX ARGENTATA CANIS FAMILIARIS

PARIETAL LOBE pAmt~htc CLEFT (entolakral fissure ; rulcur confinis)

SUPERIOR LATERAL FISSURE (ectoht-1 SUlCUd

ORBITAL LOBES coronal sulcus

SYLVIAN CLEF i CRUCIATE SULCUS ORBITAL LOBES

superior loteral fissure (ectoloterol sulcur)

(ectolateraf sulcur) (ectolaterol sulcur)

FIG;. 13. Frontal view ofl3ultri,r?oprrrtr)pf[~r~~

external intwcalate tasnSa Ierta

LfMBIC CLEFT - LAMtNA ROSTRALJS

INTERCALATE SULCUS OF THE OVAL LOBULE INFERIOR MARGINAL WLCUS

PAROLFACTORY LOBULE OF BRtJCA OLFACTORY LOBE MASSA INTERMEDIA THALAMI

cokarine *uikw (?i

farciolo cinema PARALLMBIC CLEFT (sntokzterof fissure)

OLFACTORY LOBi hippccompol rutcur UBXUI

ini ternal i ntercal ote

Pa rolfacto ry sulc :us\ \ oblique I sulcus

iporohi ppocampS al sulcus

FRANSWERSE CLEFT OF IlKHAT

PARALIMBIC CLEFT (entolateral sulcus) superior lateral fissure (ectoloteral sulcu 4

TRANSVERSE CLEFT OF BfCHAT fNTERH~MfSPHERfC CLEFT

PARALCMW CLEFT (ontofated fissure) superior lahrol firsufrr (etctolaf~al sukus]

TRANSVERSE CLEFT OF illCHAT -INTERHEMISi’HERIC CLEFT

TRANSVERSE Ct.WT OF BtCNAT

ORBITAt LOBES medial

OLFACTORY LOBE n/h

anterior rhino1 IUICUS medial orbital SUICUS ORBITAL LOBE

gyrus proreus parolfactory lobule of Broco

foramen of Monro BODY OF THE LATERAL VENTRICLE

INFERIOR HORN lateral rec*ss of IV ventricle

olfactory bulb olfactory tract temeoral Dole

ANTERIOR PERFORATE SUBSTANCE (olfactory lobe) uncus

RCtDY INFERIOR HORN

FORCEPS ANTERIOR (minor] caudots nucleus

OVAL LOBULE OCCIPITAL LOBE iNTERHEMISPHERIC CLEFT (supracaliorof part)

ORBITAL LOBE INTERHEMISPHERIC CLEFT (rubcollorol part)

SUBCALLOSAL GYRI 310LA CINEREA

IJATUS anterior rhinal sulcus oifoctory gyrus transverse orbtital 9y,,,,

optic trot p+arior rhino1

hypophysis rplmiem of corpus culiorum gyrus samtlunoris

““CU. posterior arcuate sulcus olfactory tubercle rlobule dbrsrt’)

tranrvorre ins vlw

outer banks of the

transverse central rulcur

cruciate sulcuS insular gyros

gyrus ambiens parahippocampal g;rus parahipbocampal dcus~parterior rhino1 sulcus

outer bank of the inferior lotoral fiisuro (cleft) (ectosylvian rulcur)

cirr~ular~ r&em of thkr inrula (orbitof roctor)

sulcus confinis) ~sulcus of insula

~NRCT~S~A~Al~ \ limbir honk‘ of rhinic cl& GYhUS SEMILUNARlS